Propagule pressure governs establishment of an invasive herb

The success of plant invasions may be limited by the availability of propagules and/or of suitable microsites, with microsite availability being affected by, for example, disturbance and interspecific competition. A mechanistic understanding of the contributions of propagule pressure and microsite limitation to plant invasions is therefore required to minimise future invasions. Here, we investigated the relative roles of propagule pressure, the availability of microsites, and their interaction on the establishment of an invasive herb, Lupinus polyphyllus, in two geographic regions representing different climate and growth conditions in Finland (a more productive southern region and a harsher central region). We carried out a field experiment in 14 L. polyphyllus populations, in which we manipulated both propagule pressure and disturbance. In a complementary greenhouse experiment, we manipulated propagule pressure and interspecific competition. Seedling establishment of L. polyphyllus was higher in the more productive southern region than in the harsher central region. The number of L. polyphyllus seedlings increased with increasing propagule pressure regardless of disturbance or interspecific competition. However, the number of L. polyphyllus seedlings per sown seed (relative establishment) tended to decrease with increasing propagule pressure, indicating that the positive effect of propagule pressure on early invasion is partially counteracted by density-dependent mortality at high seed densities. Our results highlight the dominant role of propagule pressure over disturbance and interspecific competition in the establishment of L. polyphyllus, suggesting that the early stage of invasion is limited by the availability of propagules rather than the availability of suitable microsites.     

Linking carbon balance to establishment patterns: comparison of whitebark pine and Engelmann spruce seedlings along an herb cover exposure gradient at treeline

There is increasing evidence that landscape vegetation patterns near species?? range limits are associated with positive biotic interactions, such as in the alpine-treeline ecotone. In the northern Rocky Mountains, whitebark pine (Pinus albicaulis) is considered an early-successional species, able to establish in exposed microsites, while late-successional species such as Engelmann spruce (Picea engelmannii) are more dependent on neighboring vegetation to facilitate establishment. We compared ecophysiological traits associated with carbon balance of newly germinated seedlings of whitebark pine and Engelmann spruce along an herb cover gradient to (1) infer which ecophysiological properties explain the establishment success of seedlings, and (2) to assess differences in establishment patterns with respect to distance from neighboring vegetation. We measured survival over 2 years, and concurrently measured gas exchange and water relations (photosynthesis, respiration, and transpiration), morphology [specific leaf area (SLA)], and biochemistry [chlorophyll fluorescence (F _v /F _m) and nonstructural carbohydrates]. Both species initially established in the most exposed microsites away from vegetation during their first growing season, but only pine persisted in exposed microsites to the end of the second growing season. Pine exhibited phenotypic traits to increase stress tolerance (e.g., higher soluble sugar concentrations, lower SLA) and improve carbon balance (e.g., greater water use efficiency, lower respiration, higher F _v /F _m) compared to spruce in exposed sites, but had lower carbon balance under herb cover. Superior establishment success of pine in exposed microsites at treeline could thus be attributed to a suite of intrinsic physiological advantages that are apparent at the earliest stage of development.     

Population structure analysis provides insights into the infection biology and invasion strategies of Kretzschmaria deusta in trees

Infection biology and invasion strategies of Kretzschmaria deusta were investigated through the analysis of patterns of colonization and population diversity in three case studies, comprising different host species. Molecular analysis and isolation assays performed on stem sections at different heights indicated a prevalent heart rot mode of expansion. Random amplified microsatellites – RAMs and somatic incompatibility assays on isolates allowed detection, in one case, of a genet occupying the entire decay column and, in the other cases, of several different genets in each individual tree. Hypotheses on modes of arrival and entrance of K. deusta in trees are discussed on the basis of the distribution of genets and population genetics analyses. Significant correlation (Spearman ρ = 1.0; p < 0.001) between the number of genets and the number of areas delimited by pseudosclerotial plates (PSPs) on the decayed portions of stem sections suggests PSPs are interaction zone lines between different individuals of K. deusta occupying adjacent decay columns.     

Migration of herbaceous woodland flora into post-agricultural black alder woods planted on wet and fertile habitats in south western Poland

Because post-agricultural black alder woods have not been the subject of studies on forest recovery, the aim of the research was to investigate the process of colonization of their herb layer by woodland flora. Migration rates of 51 forest plant species in the herb layer of alder woods planted on abandoned meadows, bordering meso- and eutrophic forests, were studied. Mean migration rates calculated for typical wet alderwoods reached 1.20–1.60 m yr⁻¹, for oak-hornbeam forests 1.17–1.63 m yr⁻¹, and for alder-ash carrs 0.79–1.26 m yr⁻¹, exceeding those reported in studies conducted in less fertile, drier sites. Although migration rates for many species exceeded 2 or 3 m, there was a group of slow colonizing species, mainly ancient woodland indicators. Species migration patterns fitted either the model based on the establishment of isolated individuals, or the model of a wave front. The age of the recent forests and the presence of Urtica dioica in their herb layer are the main factors explaining the species composition in post-agricultural black alder woods.     

Determination of tipping points for aquatic plants and water quality parameters in fish pond systems: A multi-year approach

High levels of nutrients in fish ponds by fish farming may cause significant eutrophication leading to a loss in species richness and a decrease of cover of aquatic plants to phytoplankton dominance. This shift can be represented by a tipping point where a significant change in the state of the ecosystem is observed such as a change from high to low aquatic plants species richness and cover. A total of 100 fish ponds were studied during five years in the Dombes region, France, to determine tipping points in aquatic plant richness and cover using chlorophyll α (CHL), water transparency, Total N (TN) and Total P (TP) gradients with two statistical methods. The relationships between tipping points, nutrient loads and yearly variations in weather conditions were also evaluated. Looking at the five years data, tipping points were observed in aquatic plant richness at 6 and 60 μg/L for CHL, and at 3.90 mg/L for TN concentration; as well as at 70 cm for water transparency, but no tipping point was found with TP. For aquatic plant cover, tipping points were observed at 11 μg/L for CHL, 2.42 mg/L for TN, 0.05 mg/L for TP, and at 62 cm for water transparency. These tipping points showed a significant decrease of aquatic plant species richness and cover, linked to the nutrient concentrations which drive the competition between the primary producers phytoplankton and aquatic plants. However, tipping points could vary significantly between years. The inter-annual variability may be due to an early occurrence of phytoplankton blooms in some ponds in a year preventing the establishment of aquatic plants, and thus influencing the value of tipping points. Weather conditions influence the competition between primary producers by impacting chlorophyll α and nutrients concentrations. When weather conditions supported increased nutrient concentrations, the development of phytoplankton and aquatic plants was facilitated and tipping points in aquatic plant richness and cover occurred with relatively high values. Thus, a significant decrease of plant cover and richness occurred at higher level of nutrients compared to the other years. In these cases, aquatic plants dominated over phytoplankton for the spring period, and also often during summer. In conclusion, tipping points observed are mainly linked to the competition between aquatic plants and phytoplankton. In shallow and eutrophic systems like fish ponds where nutrients are not a limiting resource, weather conditions act temporarily during spring as the main regulator of this competition.     

Spatial distribution pattern of a clonal species: effects of differential production of clonal and sexual offspring

The spatial distribution patterns of genets and ramets within populations are expected to change as a function of the frequency with which clonal species recruit different types of offspring (sexual and clonal). We used an integrated approach to study the spatial arrangement of clonal plants by combining molecular and ecological data using Opuntia microdasys as a study system. The species is able to produce two types of clonal (plantlets and cladodes) and one type of sexual (seeds) offspring. Additionally it is found in three habitats that cause differences in the ability of each type of offspring to establish. In 2007, all individuals in the three habitats (162 in BH = bajada, 264 in IDH = hill-piedmont, and 136 at HPH = interdunes) were tagged and mapped. Amplified inter-simple sequence repeats (ISSR’s) were used to determine the multilocus genotype and relatedness of each individual ramet using 120 polymorphic bands (104 in BH, 128 in HPH and 180 in IDH). The spatial distribution pattern of genets and ramets was analyzed with the Hopkins test and spatial autocorrelation analysis. For all habitats we found that O. microdasys displayed a spatial distribution characterized by clumps of aggregated ramets, but habitats differed in the number of genets present. As for other clonal species a strong positive spatial autocorrelation exists within 20 m, although all analyses suggest that adjacent ramets are genetically less related to each other or belong to different genets, that is, ramets of different genets are intermingled. The spatial arrangement of genets and ramets in O. microdasys between habitats closely matches the frequency of establishment of each type of offspring (e.g. the more clonal areas are clumped groups of related individuals). These results confirm that in two habitats (BH and IDH) clonal recruitment had been more common than in the other habitat (HPH).     

Regeneration of N.W. African Pinus halepensis forests following fire

Pinus halepensis forests of N.W. Algeria are subjected to frequent fires. During the fire the aboveground parts of plants are completely burned but only a few species are killed. Most perennial herb and shrub species survive owing to their underground organs and regenerate vegetatively in the next moist period. The semi-shrubs regenerate both vegetatively and from seeds. The most intensive growth of the shrub layer occurs during the first 2 years and in the 5th year, it reaches a height of 1–1.5 m. Pinus halepensis is completely killed by the fire and it regenerates from seeds only. The regeneration is retarded during the first 2–3 years, apparently by competition of the rapidly developing shrubs and semi-shrubs with P. halepensis. In the following years, there is a more rapid increase in both density and height, although by the 5th year after fire, the height does not exceed 0.5 m. The young trees overtop the shrub layer between 10 and 15 years after fire. The increase in density and cover supress the lower layers, in particular the herb layer. The reduction in density of trees in the following decades enables the herb layer to reconstitute its composition and cover.This process of regeneration resembles forest growth cycles rather than a secondary succession. The shrub and herb layers maintain their identity as they are mostly formed of the same individuals as before the fire; they merely regenerated their aboveground organs. Only the tree layer regenerates anew after the fire.     

Influences of the genetic neighborhood on ramet reproductive success in a clonal desert cactus

Clonal structure in clonal plants can affect sexual reproduction. Individual ramets can decrease reproduction if their neighbors are ramets of the same genet due to inbreeding depression or self-incompatibility. We assessed ramet reproductive success in the partial self-incompatible Ferocactus robustus (Cactaceae) as a function of floral display size in focal ramets and floral display size and clonal structure of their reproductive neighborhoods. Ramets were labeled, sized in number of stems, mapped and genetically identified through RAPD markers in one population. A pollen dispersal area of 15-m radius was established for each ramet to determine the clonal diversity in the neighborhoods. Flower production and fruit set were counted on a monthly basis during one reproductive season as a surrogate of ramet fitness. We expected a decrease in individual ramet reproductive success as a function of the number of reproductive ramets of the same genet in the neighborhood. A total of 272 sampled ramets revealed 116 multilocus genotypes, showing high clonal diversity in the population (G/N = 0.43, D = 0.98). Clonal diversity of neighborhoods ranged from 0.06 to 1 and fruit set varied from 0 to 76.9%. Individual ramet reproductive success was influenced by (1) mate availability, (2) floral display size of a genet within the reproductive neighborhood, and (3) the proportion of distinguishable genotypes. Floral display size of genets and ramets coupled with the genetic diversity within the reproductive neighborhood determines the low sexual reproduction in F. robustus.     

Population structure and ecology of a tropical rare rhizomatous species of teosinte Zea diploperennis (Gramineae).

One of the objectives of the Sierra de Manantlán Biosphere Reserve (Jalisco, México) is the conservation in situ of the teosinte Zea diploperennis Iltis, Doebley, Guzman & Pazzi. Zea diploperennis is perennial, shade intolerant and its 1-3 m shoots are architecturally similar to maize. Clonal growth is of the phalanx type. Genets are iteroparous (modules semelparous). The demography of seven module and genet populations was studied in seven sites representing three stages of old-field succession. Seven permanent one-meter-square plots were randomly established in each site. All genets initially present and those that became established during our study were mapped and labeled according to year of establishment. The magnitude of demographic fluctuations was greater in module populations. Genet population dynamics followed a seasonal rhythm with a maximum population size obtained at the onset of the rainy season. A relation was documented between percent annual mortality of a cohort and its age: the younger the cohort, the greater the mortality. This was a statistically significant relationship, Y = [sin(-0.288x + 1.657)]2 (r = 0.92, p < 0.01), where is proportion annual mortality of genets and is the age of the cohort. The maximum rates of genet mortality occurred during the rainy season when population densities were greatest. As a consequence, we postulate that competition occurs principally during the rainy season. Linear relationships were observed between rate of population increase of genets versus old-field successional stage and soil type. Those areas with poor soil (Ultisol), degraded soil or soils with similar physical characteristics could be rehabilitated by introducing Z. diploperennis. Such rehabilitation would achieve two distinct objectives, promote propagation of this rare endemic and reclaim areas that are susceptible to erosion and further degradation.     

Taxonomy and distribution of Allium ursinum (Liliaceae) in Poland and adjacent countries

The paper presents distribution data on Allium ursinum L. in Poland, with additional records from the other European countries. The study comprised 367 herbarium specimens, drawn from most of Polish herbaria and individuals obtained during field investigations. Two subspecies have been recorded: A. ursinum L. subsp. ursinum and A. ursinum subsp. ucrainicum Kleop. et Oxner. The study also documents the presence of intermediate populations at the border of the distribution ranges for both subspecies. In addition, the paper presents the pedicel structure and pollen grains ultrasculpture of each subspecies under a scanning electron microscope (SEM). Finally, questions in respect of the distribution data and habitat preferences are discussed.     

High phenotypic variability in the wood decay fungus Phellopilus nigrolimitatus

Genetic and phenotypic variation among individuals increases population resilience to external disturbances and enables quicker adaptation to changing environments. Since the predominant phase in the fungal lifecycle occurs belowground or within substrates, it is difficult to assess the level of variation in functional traits occurring within species in natural environments. In this study, we investigated phenotypic variation in the wood-decay fungus Phellopilus nigrolimitatus and related this to genet size, measured as vegetative compatibility groups (VCGs). Out of 321 wood samples and sporocarps collected on six logs of Picea abies, we isolated 230 dikaryotic isolates that were grouped into 53 VCGs based on culturing experiments. The number of VCGs per log varied from six to twelve, with large variation in sizes; the smallest VCG was derived from a single wood sample, while the largest was isolated across 31 wood samples stretching over 15 m. The size of the VCGs was positively correlated with the area of the spore-producing layer of the sporocarps, implying that larger genets produce more spores. Additionally, the phenotypic variation among isolates of P. nigrolimitatus was high, both in terms of growth morphology and coloration, and consistent with the delimitation of isolates into VCGs. Isolates could to a large extent be correctly assigned to the VCGs a priori based purely on culture phenotype. This study reveals that extensive phenotypic variation can be observed between fungal genets at very fine spatial scale, and calls for more studies looking at intraspecific phenotypic variation, in line with assessing genetic diversity in fungi.     

Using phytostabilisation to conserve threatened endemic species in southeastern Democratic Republic of the Congo

Outcrops in the southeastern Democratic Republic of the Congo (DRC) are recognized as some of the largest copper–cobalt orebodies in the world. They support a unique vegetation with nearly 600 metallophytes that include rare and endemic species. Mineral exploitation has increased considerably in the region since the 1900s, affecting both environmental and public health. Phytostabilisation of polluted areas represents an opportunity to decrease the bioavailability of heavy metals in the highly polluted soils that result from ore extraction. Such a technique has been successfully implemented near Lubumbashi with the grass Microchloa altera. However, long-term maintenance requires a good understanding of interspecific relationships, such as competition and facilitation. This study tested the establishment success of four herbaceous species from the Katangan Copperbelt by assessing the potential role of Microchloa altera as a nurse species. Two annual and two perennial species were sown in an experiment designed to study the influence of soil amendment and vegetation cover on seedling emergence, growth, and survival. These variables were monitored during the vegetation growing season as well as resprouting success for perennials. Microchloa altera showed a distinct effect on the emergence and survival of annual and perennial species and negatively affected the growth of individuals belonging to both groups of species.     

Study on ascending and descending vertical dispersal behavior of third instar larvae of Chrysomya megacephala (Fabricius) (Diptera:Calliphoridae): An evidence that blowflies survive burial

Although the pupation behavior of blowflies has been widely studied, this preliminary study was done on the vertical dispersal behavior (both ascending and descending) and fly emergence rate of third instar larvae of Chrysomya megacephala (Fabricius) to evaluate weather immature stages of blowflies survive burial and emerge out as adult. Third instar larvae of Chrysomya megacephala were placed at three different depths (5 cm, 25 cm and 45 cm) of soil under laboratory conditions to determine the impact of soil depth on the ascending and descending vertical dispersal behavior and the subsequent emergence of adults. The results of this study, although preliminary, but valuable to the field of forensic entomology because they provide new information about both ascending and descending vertical dispersal behavior of Chrysomya megacephala forensically important species of blowfly. In all the cases, maximum number of pupae recovered at the depth of 0 to 5 cm are 35.5 ± 4.5, 34 ± 1, 25 ± 5, when food was located at 5 cm, 25 cm and 45 cm depth respectively. This means that maximum no of larvae reached to the depth of 0 to 5 cm by ascending dispersal irrespective of at which depth they are placed. Paramount pupae were recovered from shallow burial depth of 0–5 cm in ascending dispersal and showed highest eclosion success i.e. 90.1% followed by 25 cm and 45 cm i.e. 71.7% and 55% respectively. While the number of pupae recovered as well as eclosion success was less in descending dispersal with an average of 62.8%, 39.25% and 33.9% at depths of 5, 25 and 45 cm respectively. This manifest if larvae disperse ascendingly, it increases their chance of survival.     

Dynamic process,spatial pattern and species coexistence in plants

The significance of dynamic processes of individual genets/ramets for the spatial pattern of plant species and community structure is discussed. It is suggested that under a different mode of competition (symmetric vs. asymmetric), spatial distribution of individuals, initial size distribution at the establishment stage and boundary conditions as recruitment influence differently the species coexistence pattern. It is therefore important to consider the mode of competition for the study of community structure. To know the mode and degree of intra- and interspecific competition, the dynamic processes of individual genets/ramets must be studied by following the growth, mortality and recruitment of each genet/ramet of each component species in a plant community. The models and methods of plant population ecology are therefore useful also for plant community ecology.     

Genetic population structure of three Armillaria species at the landscape scale: a case study from Swiss Pinus mugo forests

Armillaria species are plant pathogens that cause Armillaria root rot and are known to cause mortality of mountain pines (Pinus mugo) in the Swiss National Park in the Central Alps. The identity of isolates and the spatially explicit population structure of the Armillaria species were investigated in a 3.3 km² study area in the Swiss National Park. In total, 242 Armillaria isolates, 205 from wood samples and 37 from epiphytic rhizomorphs, were collected. Species were identified using haploid–diploid pairings and genets were determined using intraspecific somatic incompatibility tests. The population structure differed markedly among the Armillaria species. A. cepistipes and A. borealis mainly occurred as genets of small spatial extent (mean 0.2 ha, and 0.6 ha), whereas A. ostoyae formed significantly larger genets (mean 6.8 ha). The largest A. ostoyae genet extended over approx. 37 ha. Several disease centres associated with Heterobasidion annosum were found to be embedded within large Armillaria genets. The extension of large A. ostoyae genets suggests that forests that occupy the study area have developed in the presence of these Armillaria genets. The finding of large Armillaria genets supports the assumption that large genets occur in areas with cold climate and little precipitation.     

Genet dynamics of a regenerating dwarf bamboo population across heterogeneous light environments in a temperate forest understorey

Despite the advantage of plant clonality in patchy environments, studies focusing on genet demography in relation to spatially heterogeneous environments remain scarce. Regeneration of bamboos in forest understoreys after synchronous die‐off provides an opportunity for assessing how they come to proliferate across heterogeneous light environments. In a Japanese forest, we examined genet demography of a population of Sasa kurilensis over a 7‐year period starting 10 years after die‐off, shortly after which some genets began spreading horizontally by rhizomes. The aboveground biomass was estimated, and genets were discriminated in 9‐m² plots placed under both canopy gaps and closed canopies. Overall, the results suggest that the survival and spread of more productive genets and the spatial expansion of genets into closed canopies underlie the proliferation of S. kurilensis. Compared to canopy gaps, the recovery rate of biomass was much slower under closed canopies for the first 10 years after the die‐off, but became accelerated during the next 7 years. Genet survival was greater for more productive genets (with greater initial number of culms), and the spaces occupied by genets that died were often colonized afterward by clonal growth of surviving genets. The number of genets decreased under canopy gaps due to greater mortality, but increased under closed canopies where greater number of genets colonized clonally from outside the plots than genets died. The colonizing genets were more productive (having larger culms) than those originally germinated within the plots, and the contribution of colonizing genets to the biomass was greater under closed canopies. Our study emphasizes the importance of investigating genet dynamics over relevant spatiotemporal scales to reveal processes underlying the success of clonal plants in heterogeneous habitats.     

Grazing and neighborhood interactions limit woody encroachment in wet subtropical savannas

Woody encroachment in savannas is a worldwide concern, and there is growing consensus that anthropogenic activities play a central role in changing tree – grass interactions. We evaluated the influence of livestock grazing and neighborhood interactions on seedling emergence and survival of the native tree Acacia caven in wet savannas of northeastern Argentina. We hypothesized that grazing and grass competition act as biotic barriers limiting tree recruitment, but the relative magnitude of such barriers differs according to grass patch type. In two consecutive years (cohort 1 and 2) we sowed seeds and transplanted seedlings of Acacia in two grass patch types (prostrate/palatable and tussock/unpalatable grasses) in both, grazed and ungrazed plots. Each grass patch type was further manipulated to create three levels of grass competition (unclipped control, above-ground biomass removal and total biomass removal).Cattle grazing diminished seedling emergence of both cohorts and seedling survival of cohort 1. The effect of grass competition changed according to grass patch type. Prostrate grass cover enhanced emergence but lowered early survival, while tussock grass cover and also its total biomass removal facilitated early survival. During the second year, a severe drought drastically reduced Acacia recruitment, and it was strong enough to eliminate any grazing effects although the effect of grass competition on seedling establishment remained significant.Our results suggest that grazing and grass competition additively diminished the risk of woody establishment in this wet savanna. However, the stocking rate should be carefully balanced, thus contributing to the maintenance of a competitive grass cover to limit tree recruitment.     

Habitat selection and abundance of common genets <Emphasis Type="Italic">Genetta genetta</Emphasis> using camera capture-mark-recapture data

Using camera-trapping techniques, the present study, conducted from 2005 to 2007, provides common genet abundance estimates in Serra da Malcata Nature Reserve (central-eastern Portugal). We estimated genet abundance using the software CAPTURE. It was possible to obtain a capture success of 1.49 captures/100 trap-nights. Considering the heterogeneity model (M_h), which presents higher biological significance, the estimated density varied between 0.50 (95% CI = 0.43–0.56 genets/km²) to 0.92 (95% CI = 0.87–0.97 genets/km²) genets/km² with an average density value of 0.70 genets/km² (95% CI = 0.58–0.82 genets/km²). These estimates emphasized this technique as a reliable method for assessing average genet density over large spatial scales and for monitoring future changes in genet numbers. In terms of habitat selection, genets selected Quercus rotundifolia and Arbutus unedo woodlands and avoided Erica spp. and Cistus ladanifer scrubland and Eucalyptus stands. Considering the landscape heterogeneity outside the reserve, our study emphasizes the importance of the protected area for small carnivore conservation.     

Colonization of secondary woodlands by Anemone nemorosa

Migration of the herb species Anemone nemorosa from older woodlands into adjacent, recently established deciduous woodlands on former arable land was studied. The wood anemone had colonized part of all studied recent woods, varying in age between 30 and 75 years. Cover of A. nemorosa in the recent woods decreased with increasing distance from the older woodland at all sites but one, indicating dispersal limitation during colonization. The advancing edge of most populations was characterized by negative logarithmic decrease in cover with establishment of isolated pioneer individuals and later gradual infill between pioneers. Migration rates were calculated by three methods. The mean migration rate of A. nemorosa based on observed maximum cover in the recent woods was 0.20 m year¹, 0.40 m year^-1 based on half maximum cover and 0.85 m year¹ based on the individual found farthest from the former woodland border. The calculated migration rates were consistently higher than the rate of possible rhizome growth. Seed dispersal and establishment is thus very important for colonization of new woodlands. Migration rates increased with tree canopy cover–especially cover of broad-leaved species with quickly decomposing litter–and with soil pH. Migration rates decreased with increasing grass cover in the field layer. These differences in migration rates may be due to increased micro-site availability for establishment at high canopy cover, low grass cover and high soil pH. Our results show that the wood anemone generally colonizes recent woods from nearby source populations. However, colonization proceeds relatively slowly and is limited by both seed dispersal and availability of suitable micro-sites. Gradients in abundance of A. nemorosa within secondary woods may be detectable for long periods of time and indicate the recent origin of a woodland.     

Response of herbaceous layer species to canopy and soil variables in a central Appalachian hardwood forest ecosystem

Previous work has suggested that excess nitrogen (N) alters the degree to which forest canopy versus soil variables influence forest herb communities. This study tested the hypothesis that excess N would shift this influence on individual herb species from soil N availability to stand structural variables that determine light availability to the forest floor. Two watersheds at the Fernow Experimental Forest, West Virginia, USA were used: WS4 and WS3 as untreated reference and treatment watersheds, respectively. WS3 receives 35 kg N/ha/year via aerial application. Herb cover and composition was measured in seven permanent plots/WS from 1991, currently on-going. In 2011, soil moisture and N availability were measured in each plot, along with several variables of canopy structure. Backwards stepwise regression was used to determine relationships between herb cover/individual species and soil/canopy measurements. Herb cover varied spatially with soil resources on WS4, whereas cover varied spatially with canopy structure on WS3. Although results for total herb layer cover supported this hypothesis, results for individual herb species rejected it. This contrast was especially evident for Rubus allegheniensis (blackberry), a nitrophilic species which increased with increasing soil N on both watersheds, but was not correlated with canopy structure on reference WS4, while being correlated with canopy structure on N-treated WS3. Excess N from atmospheric deposition has been shown to decrease plant biodiversity of impacted forests, especially in its effects on herbaceous layer communities. This work demonstrates that one of the mechanisms of such response is in N-mediated changes in the response of herb communities to soil resources and light availability.