Distribution and diet of chamois (

Changes in Hieracium abundance in Eastern Otago tussock grassland were examined by sampling 163 sites in 1982 and again in 1992. For Hieracium pilosella, H. praealtum and H. lepidulum, as well as Agrostis capillaris for comparison, colonisation of new sites was recorded, as well as extinction of species from sites over the 10 years, and changes in cover. H. pilosella colonised the majority of sites from which it had been absent in 1982; it disappeared from only a few sites where it had been present at very low cover. In sites where it remained over the decade, its cover increased by 50%. Its rate of increase was very similar to that recorded for North Canterbury, but with no indication of the 34% stabilisation point that has been hypothesised for the latter area. H. praealtum and H. lepidulum, in contrast, remained absent from most sites, and colonised only a few; in very few sites was their cover greater than 1%. Agrostis capillaris was widespread and abundant, but increased little over the period. The sites where H. pilosella colonised tended to be degraded, with a greater, and increasing, percentage of bare ground and low tussock cover. The few sites where it disappeared were ones where pasture improvement had occurred, as seen in the increase of species indicative of pasture development. Several explanations are considered for the increase in H. pilosella in recent decades. The increase of this species in Eastern Otago has been greater than that of most other exotic species. Changes in land management have contributed to its increase, but probably changes caused indirectly by government policy, not gradual degradation. Genetic changes in the species, and metapopulation dynamics, have not been investigated, but they could well have contributed to Hieracium invasion.     

Rapid short-tussock grassland decline with and without grazing, Marlborough, New Zealand

Species abundance, species richness, and ground cover were measured over 10 years on nine paired grazed and exclosure plots in short-tussock grassland in the early stages of invasion by Hieracium species. With and without grazing, H. pilosella and H. caespitosum increased markedly and H. lepidulum increased locally. In contrast, 50% of all other common species and species groups, and total, native, and exotic species richness declined significantly. Exclusion increased or had no effect on rates of increase in Hieracium species and rates of decline in short tussocks, and did not reduce rates of decline in other species. Exclusion had no effect on decline in native species richness, but mainly accelerated declines in total and exotic richness. Declines in 13 key vegetation variables were significantly predicted by increase in Hieracium abundance, suggesting competitive exclusion. With or without grazing, Hieracium species will become more dominant and other species will continue to decline. The effects of large herbivores on plant species diversity can often be predicted from site productivity. Our results indicate the need also to account for species origin, spatial scale, time, and exotic invasion.     

The distribution and abundance of

We examined the distribution and abundance of the invasive Hieracium species (hawkweeds) in the dry grasslands of the Upper Waitaki Basin (Canterbury) and Otago, using measures of Hieracium species frequency and hawkweed cover from 301 vegetation plots. Average hawkweed cover was significantly less in Otago than Canterbury. Hawkweed cover was also lower on drier sites, with hawkweeds having less cover at lower elevation, on more xeric sites and, in Canterbury, on soils with a lower moisture holding capacity. Environmental variables (elevation, moisture index, soil class, and measures of rabbit activity and the amount of recent pasture development) explained 43% of the variation in hawkweed cover in Canterbury, but only 25% in Otago. Furthermore, having controlled for environmental differences among plots, hawkweed cover in Otago was significantly spatially autocorrelated such that plots on the same property had similar hawkweed cover, but tended to differ from plots on other properties. This pattern is consistent with the hypothesis that variation in property management influenced the degree of hawkweed infestation. However, the same pattern was not found in Canterbury. We propose a general model to explain the observed patterns. The probability that a site will be invaded by hawkweeds is a function of: (1) the suitability of the site for hawkweed establishment (a function of the environment and past management), and (2) the size of the hawkweed propagule rain. The probability of hawkweed invasion is high when one or both of these variables have high values. Following initial invasion, there will be more local propagules coming from newly established plants and hawkweed cover at a site will increase, eventually stabilising at a level predictable from the environmental conditions at the site. This model predicts that the sites most suited for hawkweed establishment would be invaded first and fill the quickest. These susceptible sites would then serve as foci, providing propagules for subsequent hawkweed spread into adjacent, less invasion prone areas. The model predicts that the initial increase in hawkweed cover at a site should be exponential, and that there should be a lag in hawkweed spread whereby less susceptible sites are invaded later or at a slower rate. Both predictions are supported by data from monitoring trials. The model can explain the lower abundance of hawkweeds, the less predictable distribution of hawkweeds in relation to environmental variables, and the significant spatial autocorrelation in hawkweed cover in Otago compared to Canterbury, if hawkweed invasion in Otago is less complete than in Canterbury.     

Influence of pastoral management on plant biodiversity in a depleted short tussock grassland, Mackenzie Basin

This study investigated the effects of different management inputs (fertiliser and seed) and grazing patterns on plant biodiversity in a short tussock grassland with a strong Hieracium pilosella component. Cover abundance of vascular and non-vascular plants and environmental variables were measured in 32 10×10-m plots located in five blocks with different management treatments. Ordination of the floristic data separated the block with the highest management inputs from other blocks. Several adventive species were significantly more abundant in this block, while several native species were either absent or uncommon, but were significantly more abundant in other study blocks. H. pilosella was significantly more abundant in blocks with lower management inputs. Diversity was significantly higher in the block with the highest management inputs. The native tussock Poa colensoi had significantly greater cover abundance while Festuca novae-zelandiae tussocks were significantly taller in this block. Our results suggest that high management inputs reduce the abundance of H. pilosella and diversity of native species, but increase the abundance of other adventive species and the cover and vigour of native tussocks. Our results highlight an interesting management conundrum for short tussock grasslands. No-input management is likely to result in a decline in native biodiversity, as well as production values, as H. pilosella mats deplete soil nutrients and restrict regeneration of native species. However, input of fertiliser and adventive seeds to enhance production values, although resulting in an increase in the vigour and abundance of some native species (mainly tussocks) and a reduction in H. pilosella abundance, will also result in a decline in overall native species richness.     

Inferring process from pattern in plant invasions: a semimechanistic model incorporating propagule pressure and environmental factors

Propagule pressure is intuitively a key factor in biological invasions: increased availability of propagules increases the chances of establishment, persistence, naturalization, and invasion. The role of propagule pressure relative to disturbance and various environmental factors is, however, difficult to quantify. We explored the relative importance of factors driving invasions using detailed data on the distribution and percentage cover of alien tree species on South Africa's Agulhas Plain (2,160 km2). Classification trees based on geology, climate, land use, and topography adequately explained distribution but not abundance (canopy cover) of three widespread invasive species (Acacia cyclops, Acacia saligna, and Pinus pinaster). A semimechanistic model was then developed to quantify the roles of propagule pressure and environmental heterogeneity in structuring invasion patterns. The intensity of propagule pressure (approximated by the distance from putative invasion foci) was a much better predictor of canopy cover than any environmental factor that was considered. The influence of environmental factors was then assessed on the residuals of the first model to determine how propagule pressure interacts with environmental factors. The mediating effect of environmental factors was species specific. Models combining propagule pressure and environmental factors successfully predicted more than 70% of the variation in canopy cover for each species.     

Changes in the Structure of Tall Tussock Grasslands and Infestation by Species of

A plant sociological survey of tall-tussock grasslands in the Mackenzie country was repeated after an interval of 26-28 years. Changes in physiognomy of the grasslands which have been inferred from earlier studies have been found to be continuing on many sites. A noteworthy feature of most sites has been a reduction in number of indigenous species found. An increase in abundance of Hieracium pilosella or H. praealtum has occurred at most sites. About 140 species and 9 variables from 53 sites were interrelated in a direct unimodal ordination (canonical correspondence analysis). Ordination results of the 1960s and the 1989 data were compared and interpreted with respect to spread of Hieracium pilosella and H. praealtum. The dynamics of Hieracium infestation was studied within changing community structure. A distinct trend is demonstrated of increasing infestation with increasing grassland degradation.     

Interspecific hybridization among Hieracium species in New Zealand: evidence from flow cytometry

Hieracium pilosella: (Asteraceae) was accidentally introduced to New Zealand about 100 years ago. Since then it has become an aggressive weed, and an unexpected degree of genetic and genome size variation has been detected; features that might result from interspecies hybridization. We investigated the possibility that H. pilosella has hybridized with related taxa. Of the four other subgenus Pilosella species introduced to New Zealand, H. praealtum is the most abundant and, on morphological and distributional evidence, most likely to be the other parent. Flow cytometry was used to estimate relative genome size for 156 Hieracium plants collected from the wild. Plants assigned to either parental or hybrid morphotypes were found to comprise tetraploid and pentaploid individuals using genome size measurements, and this was confirmed with direct mitotic chromosome counts for a subset of plants. The haploid DNA content of H. praealtum was approximately 22% larger than that of H. pilosella. Putative hybrids that were tetraploid had mean genome sizes equivalent to two H. pilosella and two H. praealtum haploid chromosome sets, implying they were hybrids arising from the fertilization of two reduced gametes. Similar results were obtained from tetraploid hybrids produced by controlled pollination. However, the majority of field hybrids were pentaploid with a genome size equivalent to four H. pilosella and one H. praealtum haploid chromosome sets. We infer that these are not first-generation hybrids but represent successful backcrossing with H. pilosella and/or hybrid-hybrid crossing, and that sexual tetraploid hybrids have been the parents. We note that populations putatively of H. pilosella often comprise apomictic pentaploid hybrids. Significantly, our data indicate the emergence of sexual hybrids that provide further opportunity for gene flow among taxa in this complex.     

Chloroplast DNA diversity of Hieracium Pilosella (Asteraceae) introduced to New Zealand: reticulation, hybridization, and invasion

The European hawkweed Hieracium pilosella is a successful invader and a troublesome weed in New Zealand. The systematics of the genus Hieracium is extremely complex and contentious, probably due to recent speciation, hybridization, polyploidy, and diverse reproductive strategies. In the first chloroplast DNA survey of the group, we sequenced 285 plants (including H. pilosella and 12 other species of subgenus Pilosella) from New Zealand and Europe for 900 bp of trnL-trnF. Eleven haplotypes were identified with much sharing among species. Three haplotypes (A, D, G) were found in seven, three, and four species, respectively, but two species (H. lactucella and H. auricula) had single, private haplotypes. Our cpDNA data for subgenus Pilosella are consistent with the group's having incomplete lineage sorting and/or recent reticulate evolution. Six haplotypes were identified in H. pilosella, four of these unique to this taxon in our sample. In New Zealand, haplotype A was common and occurred in plants of different ploidy (i.e., 4×, 5×, 6×), whereas haplotypes C, B, and M were restricted to 4×, 5×, and 6× plants, respectively. The distribution of haplotype variation suggests that some or all of the H. pilosella seeds accidentally introduced into New Zealand probably came from east Europe rather than the United Kingdom and that a minimum of four lineages were introduced. Within New Zealand, hybridization of H. pilosella with a related taxon (probably H. praealtum) has occurred at least three times, involving both obligate sexual tetraploids and facultative apomictic pentaploids of H. pilosella.     

Functional and performance comparisons of invasive Hieracium lepidulum and co‐occurring species in New Zealand

Abstract One of the key environmental factors affecting plant species abundance, including that of invasive exotics, is nutrient resource availability. Plant functional response to nutrient availability, and what this tells us about plant interactions with associated species, may therefore give us clues about underlying processes related to plant abundance and invasion. Patterns of abundance of Hieracium lepidulum, a European herbaceous invader of subalpine New Zealand, appear to be related to soil fertility/nutrient availability, however, abundance may be influenced by other factors including disturbance. In this study we compare H. lepidulum and field co‐occurring species for growth performance across artificial nutrient concentration gradients, for relative competitiveness and for response to disturbance, to construct a functional profile of the species. Hieracium lepidulum was found to be significantly different in its functional response to nutrient concentration gradients. Hieracium lepidulum had high relative growth rate, high yield and root plasticity in response to nutrient concentration dilution, relatively low absolute yield, low competitive yield and a positive response to clipping disturbance relative to other species. Based on overall functional response to nutrient concentration gradients, compared with other species found at the same field sites, we hypothesize that H. lepidulum invasion is not related to competitive domination. Relatively low tolerance of nutrient dilution leads us to predict that H. lepidulum is likely to be restricted from invading low fertility sites, including sites within alpine vegetation or where intact high biomass plant communities are found. Positive response to clipping disturbance and relatively high nutrient requirement, despite poor competitive performance, leads us to predict that H. lepidulum may respond to selective grazing disturbance of associated vegetation. These results are discussed in relation to published observations of H. lepidulum in New Zealand and possible tests for the hypotheses raised here.     

Reproduction mode in the allopolyploid facultatively apomictic hawkweed Hieracium rubrum (Asteraceae, H. subgen. Pilosella)

The versatility of the breeding system in the hybridogenous hexaploid, Hieracium rubrum, was demonstrated in emasculation and crossing experiments. The flow-cytometric ploidy analysis of 1095 seedlings amongst its' progeny enabled the determination and quantification of the reproductive pathway (apospory, haploid parthenogenesis, cross-fertilization of both reduced and unreduced female gametes) responsible for each progeny class. The progeny profiles were stable between two successive years. The percentage of hybrids arisen from crosses with tetraploid sexual H. pilosella was 7.9%. The rate of trihaploids generated by H. rubrum when crossed to H. pilosella was 3.95%, similar to that of pentaploid (n + n) and octoploid (2n + n) hybrids (3.95% and 3.39%). Unreduced pollen from H. pilosella contributed to hybridization much less frequently (0.56% of all progeny), than unreduced eggs of H. rubrum. The increased frequency of trihaploids formed by crossed compared to emasculated plants suggests an interaction between the presence of pollen and the autonomous development of meiotic embryo sacs. Although the environmental conditions (garden compared to unheated glasshouse) influenced the progeny following the emasculation, no corresponding response to environment was detected amongst the autonomously derived progeny following pollination with H. pilosella. However, there was an influence of pollen parent on progeny which varied under the different environmental treatments. The importance of residual sexuality in reproduction of apomicts is evident. The non-maternal progeny was generated with highly variable frequency especially in the glasshouse, with some capitula reaching up to 50%. Having an autonomous endosperm development, Hieracium subgen. Pilosella is almost unique among other aposporous genera, in which such variable reproduction mode is connected with pseudogamy.     

Changes in vegetation states in grazed and ungrazed Mackenzie Basin grasslands, New Zealand, 1990-2000

Changes in vegetation from 1990 to 2000 were examined at 10 high country localities, representing four grassland types: fescue tussock (Festuca novae-zelandiae), snow tussock (Chionochloa rigida), red tussock (C. rubra), and silver tussock (Poa cita). At each locality, three treatments were established: ambient sheep+rabbit grazing, rabbit grazing only, and no grazing. The mutivariate methods of classification and ordination were used on individual-quadrat cover data to define vegetation states and to examine transitions between them over time. Vegetation states in quadrats already dominated by Hieracium pilosella(> 50% cover) in 1990 showed little change in species composition regardless of grassland type and grazing treatment. In fescue tussock grassland, H. pilosellaincreased regardless of grazing treatment in states with low initial H. pilosellacover (< 5%), while the cover of Carex colensoi, Aira caryophyllea and Rumex acetosella decreased. In the single silver tussock locality, Poa citadecreased markedly in the ungrazed treatment as adventive species such as Dactylis glomerataand Echium vulgare increased. However, Poa citaalso decreased, probably due to drought, in the grazed treatment. Snow tussock and red tussock grassland states were more stable than those in short tussock grasslands, but there was also a general trend towards increasing H. pilosellacover in intertussock vegetation regardless of treatment. However, at one snow tussock locality, transitions from H. piloselladominated to C. rigida-dominated states occurred in ungrazed quadrats, while the reverse occurred in grazed vegetation. Implications for the management of tussock grasslands for conservation are discussed.     

Interaction between Some Pasture Species and 2

Possible control options are investigated for the introduced Hieracium weeds, particular problems in South Island high country, New Zealand. In a pot experiment regression of input to output ratios of above ground biomass over successive harvests, from binary mixtures was used to determine the competitive interaction between 13 pasture species and two Hieracium species, H. pilosella and H. praealtum, in a low fertility soil. Treatments also included a factorial of presence or absence of compartments separating root and shoots of species. Species differed in their mean growth rate, relative to Hieracium species. The ranking of mean growth rate relative to H. pilosella was Trifolium repens (best), Bromus inermis, Sanguisorba minor, Festuca novaezelandiae, F. rubra, Arrhenatherum elatius, Anthoxanthum odoratum, Lotus corniculatus, Hypochoeris radicata, Trifolium repens, T. hybridum, T. medium and Astragalus cicer. Shoot partitions decreased Hieracium's interaction with white clover while root partitions increased interaction with B. inermis. However, the rates were not related to the proportion of Hieracium in the mixtures, indicating a general lack of specific competitive effects against Hieracium.     

Catchment land cover as a proxy for macroinvertebrate assemblage structure in Carpathian Mountain streams

We compared land cover, riparian vegetation, and instream habitat characteristics with stream macroinvertebrate assemblages in 25 catchments in the Carpathian Mountains in Central Europe. This study area was particularly selected because of its diverse history of forest and agricultural ecosystems linked to geopolitical dynamic, which provide a suite of unique landscape scale, land cover settings in one ecoregion. Canonical Correspondence Analysis (CCA) showed that variation in composition and structure of macroinvertebrate assemblages was primarily related to four land cover types, and not to riparian or instream habitat. These were the portions in the catchment areas of (1) broadleaved forest, (2) fine-grained agricultural landscape mosaic with scattered trees (e.g., pre-industrial cultural landscape), (3) mixed forest, and (4) natural grassland without trees. Principal Component Analysis (PCA) suggested that land cover types and stream channel substrates co-varied. The PCA also showed that chemical variables, including organic carbon, had higher values in the agricultural landscape compared to natural forests. The major source of variation among taxa in streams was higher abundance of Diptera in agricultural landscapes and of Plecoptera, Coleoptera, Trichoptera, and Amphipoda in forests. Gastropoda and Oligochaeta were more abundant in open, fine-grained agricultural landscape mosaics with scattered trees. Ephemeroptera taxa were quite indifferent to these gradients in catchment land cover, but showed a tendency of being more abundant in the pre-industrial cultural landscape. Our findings suggest that land cover can be used as a proxy of the composition and structure of macroinvertebrate assemblages. This means that land use management at the catchment scale is needed for efficient conservation and recovery of stream invertebrate communities.     

Intraspecific adoption and foster feeding of fledglings in the North Island robin

We sampled soils and vegetation within and outside two sheep and rabbit exclosures, fenced in 1979, on steep sunny and shady slopes at 770 m altitude on seasonally-dry pastoral steeplands. The vegetation of sunny aspects was characterised by higher floristic diversity, annual species, and low plant cover. Here the exotic grass Anthoxanthum odoratum dominated on grazed treatments, and the exotic forb Hieracium pilosella on ungrazed. Shady aspects supported fewer, and almost entirely perennial, species. Here Hieracium pilosella dominated grazed treatments, but co-dominated with the exotic forb H. praealtum and the native grass Festuca novae-zelandiae on ungrazed treatments. There was 43% more biomass in exclosures (P < 0.01). Most of the biomass difference (4285 kg/ha) was from greater root mass (2400 kg/ha). 1385 kg/ha of the difference was from herbage and the remainder (500 kg/ha) from litter. Exclosures had 50 to 100% more Ca, Mg, K and P in the biomass (P < 0.05), but the effect on soils was limited to significantly higher concentrations of total N (P < 0.05) and exchangeable Mg (P < 0.01) in 0-7.5 cm soils. We conclude that stopping grazing for 16 years on seasonally-dry steeplands results in greater plant cover, approximately double the biomass of standing vegetation, greater biomass in roots, and more biomass nutrients relative to grazed areas. However, it does not favour native species and has little effect on soil nutrients or soil carbon. Stopping grazing alone therefore cannot be regarded as a comprehensive short- or medium-term vegetation or soil rehabilitation option.     

Disturbance affects short-term facilitation, but not long-term saturation, of exotic plant invasion in New Zealand forest

We investigate the spread of an exotic herb, Hieracium lepidulum, into a New Zealand Nothofagus forest with the aim of understanding how stand-development of tree populations, propagule pressure and invader persistence, affect invasion across the landscape and within communities. Using data repeatedly collected over 35 years, from 250 locations, we parametrize continuous-time Markov chain models and use these models to examine future projections of the invasion under a range of hypothetical scenarios. We found that the probability of invasion into a stand was relatively high following canopy disturbance and that local abundance of Hieracium was promoted by minor disturbances. However, model predictions extrapolated 45 years into the future show that neither the rate of landscape-level invasion, nor local population growth of Hieracium, was affected much by changing the frequency of canopy disturbance events. Instead, invasion levels were strongly affected by the ability of Hieracium to persist in the understorey following forest canopy closure, and by propagule supply from streams, forest edges and plants already established within the stand. Our results show that disturbance frequency has surprisingly little influence on the long-term trajectory of invasion, while invader persistence strongly determines invasion patterns.     

Resource availability and plant diversity explain patterns of invasion of an exotic grass

Aims In this study, we examine two common invasion biology hypotheses—biotic resistance and fluctuating resource availability—to explain the patterns of invasion of an invasive grass, Microstegium vimineum.Methods We used 13-year-old deer exclosures in Great Smoky Mountains National Park, USA, to examine how chronic disturbance by deer browsing affects available resources, plant diversity, and invasion in an understory plant community. Using two replicate 1 m 2 plots in each deer browsed and unbrowsed area, we recorded each plant species present, the abundance per species, and the fractional percent cover of vegetation by the cover classes: herbaceous, woody, and graminoid. For each sample plot, we also estimated overstory canopy cover, soil moisture, total soil carbon and nitrogen, and soil pH as a measure of abiotic differences between plots.Important findings We found that plant community composition between chronically browsed and unbrowsed plots differed markedly. Plant diversity was 40% lower in browsed than in unbrowsed plots. At our sites, diversity explained 48% and woody plant cover 35% of the variation in M. vimineum abundance. In addition, we found 3.3 times less M. vimineum in the unbrowsed plots due to higher woody plant cover and plant diversity than in the browsed plots. A parsimonious explanation of these results indicate that disturbances such as herbivory may elicit multiple conditions, namely releasing available resources such as open space, light, and decreasing plant diversity, which may facilitate the proliferation of an invasive species. Finally, by testing two different hypotheses, this study addresses more recent calls to incorporate multiple hypotheses into research attempting to explain plant invasion.     

Gradients in stream fish assemblages across a Mediterranean landscape: contributions of environmental factors and spatial structure

1. Despite wide recognition that fish assemblages are influenced by factors operating over a range of spatial scales, little effort has been devoted to quantifying large‐scale variation and the multiscale dependencies of assemblage patterns and processes. This is particularly true for Mediterranean streams, where seasonally predictable drying‐up may lead to a strong association between assemblage attributes and large‐scale factors affecting the distribution of population sources and extinction likelihood. 2. The contribution of large‐scale factors to stream fish assemblage variation was quantified across a Mediterranean landscape, in south‐west Portugal. Fish abundance and species composition were estimated at 166 sites across third‐ to sixth‐order streams, in March–July 1998. Variance partitioning by redundancy analyses was used to analyse assemblage variation against three sets of predictor variables: environmental (catchment position, and geomorphic and hydrological factors), large‐scale spatial trends and neighbourhood effects. 3. Environmental variables and spatial trends accounted for 34.6% of the assemblage variation across the entire region, and for 36.6 and 57.8% within the two largest catchments (Mira and Seixe). Neighbourhood effects were analysed at the catchment scale, increasing the explained variation to 56.1% (Mira) and 70.7% (Seixe). 4. A prevailing environmental gradient was reflected in an increase in the abundance of all species and size‐classes in relation to catchment position, with more fish present in larger streams and in downstream reaches. Variables describing geomorphic and hydrological settings were less important in explaining assemblage variation. 5. Spatial trends always accounted for the smallest fraction of assemblage variation, and they were probably associated with historical barriers to fish dispersal. The strong neighbourhood effects may be related to spatially autocorrelated habitat conditions, but they are also a likely consequence of fish emigration/extinction and colonisation processes. 6. These results emphasise that a substantial proportion of fish assemblage variation in Mediterranean streams may be explained by large‐scale factors, irrespective of microhabitats and local biotic interactions. It is suggested that this pattern results to a large extent from the seasonal drying‐up, with the summer shortage of surface water limiting fish occurrence in headwaters, and consequently the key core areas for fish concentrating in larger streams and tributaries adjacent to large streams because of neighbourhood effects.     

The Influence of Landscape Position and Catchment Characteristics on Aquatic Biogeochemistry in High-Elevation Lake-Chains

To examine the influence of landscape characteristics and landscape position on aquatic biogeochemistry, we sampled a total of 76 lakes within 14 different lake-chains spanning the latitudinal extent of the high-elevation Sierra Nevada (California). We measured water chemistry, dissolved organic matter (DOM), nutrients, and biotic variables in study catchments that encompassed representative ranges of area (3–22 km²), elevation (2,200–3,700 m.a.s.l), elevation change (50–700 m), and average slope (13°–26°). Hierarchical models were used to account for variability in biogeochemistry because they explicitly maintain the framework of lakes within individual lake-chains while accounting for variation among lake-chains. Unconditional means models, where lake-chain was a random effect, revealed significant differences among lake-chains for nearly all biogeochemical variables. Models explained 42–95% of this variability, with the majority of the variation (70%) explained by the among lake-chain component. To explore the amount of additional variation explained by lake landscape position, we added lake network number (LNN) to models. LNN explained a significant amount of additional variation (7% average) in 8 of 23 biogeochemical parameters. However, it explained more variability within individual lake-chains (75%), where among lake-chain differences did not obscure patterns. Patterns of increase with LNN were found for dissolved organic carbon and nitrogen, fluorescence of DOM, alkalinity, and bacterioplankton abundance, whereas nitrate and nitrogen to phosphorus nutrient ratios decreased. LNN explained variation because it served as a proxy for underlying catchment characteristics that changed consistently along downstream flow paths. To characterize the amount of variation explained by catchment characteristics alone, we fit a third model that included lake-chain as a random effect and landscape or lake morphometry attributes as fixed effects. Catchment characteristics explained about as much additional variation (6%) as LNN, but for substantially more biogeochemical parameters (18 out of 23). The catchment characteristics most predictive of biogeochemistry were land-cover factors delineating alpine and subalpine zones (elevation, slope, or proportions of rock or shrub cover). In general, catchment characteristics were stronger predictors of biogeochemistry than characteristics of lake morphometry, emphasizing the relative importance of landscape processes in snowmelt-dominated lake ecosystems.     

Sexual and vegetative reproduction of Hieracium pilosella L. under competition and disturbance: a grid-based simulation model

We used a spatially explicit simulation model to examine the relative importance of vegetative and sexual reproduction in Hieracium pilosella L. Based on an understanding of the complex life cycle of this species and on data from in situ population dynamics in a calcareous grassland in NW Switzerland, we simulated growth and the relative contribution of clonal reproduction by stolons and reproduction by seeds across a gradient of increasing soil fertility. Competition by a clonal grass resulted in nearly complete exclusion of H. pilosella from the more fertile part of the simulation plot. Under low soil fertility, when grass could not survive, H. pilosella largely persisted by vegetatively produced rosettes. This pattern of a sharp separation of both species was shifted slightly in favour of H. pilosella by introducing random disturbances. Only by adding: (1) long-distance seed dispersal, and (2) facilitation of seedling establishment in the vicinity of grass tussocks in vegetation gaps was a more realistic representation of field observations realised, with rosettes of H. pilosella grown from seeds occasionally distributed within dense grass vegetation. Phenotypic plasticity of stolon length was a decisive factor for the maintenance of H. pilosella populations. We conclude that a mixed strategy of clonal growth and reproduction by seeds in H. pilosella is necessary to maintain populations of this species in the presence of high interspecific competition and a shortage of open space.