Effects of climate and snow depth on Bromus tectorum population dynamics at high elevation

Invasive plants are thought to be especially capable of range shifts or expansion in response to climate change due to high dispersal and colonization abilities. Although highly invasive throughout the Intermountain West, the presence and impact of the grass Bromus tectorum has been limited at higher elevations in the eastern Sierra Nevada, potentially due to extreme wintertime conditions. However, climate models project an upward elevational shift of climate regimes in the Sierra Nevada that could favor B. tectorum expansion. This research specifically examined the effects of experimental snow depth manipulations and interannual climate variability over 5 years on B. tectorum populations at high elevation (2,175 m). Experimentally-increased snow depth had an effect on phenology and biomass, but no effect on individual fecundity. Instead an experimentally-increased snowpack inhibited population growth in 1 year by reducing seedling emergence and early survival. A similar negative effect of increased snow was observed 2 years later. However, a strong negative effect on B. tectorum was also associated with a naturally low-snow winter, when seedling emergence was reduced by 86%. Across 5 years, winters with greater snow cover and a slower accumulation of degree-days coincided with higher B. tectorum seedling density and population growth. Thus, we observed negative effects associated with both experimentally-increased and naturally-decreased snowpacks. It is likely that the effect of snow at high elevation is nonlinear and differs from lower elevations where wintertime germination can be favorable. Additionally, we observed a doubling of population size in 1 year, which is alarming at this elevation.     

Effects of resource availability on seedling recruitment in a fire-maintained savanna

The herbaceous ground cover of the longleaf pine ecosystem harbors the highest plant species richness in North America, with up to 50 species per square meter, but the mechanisms that regulate this diversity are not well understood. In this system, variability in seedling recruitment events may best explain the extremely high small-scale species richness and its relationship to soil moisture and system net primary productivity. To understand the potential mechanistic controls on species richness, we used a long-term resource manipulation study across a natural soil moisture gradient to assess environmental controls on seedling recruitment. We considered the availability of resources to be an indicator of seedling safe-site supply, and also manipulated seed availability to examine the relative importance of recruitment limitations on seedling diversity. We found that water availability regulated the number of species in the seedling community regardless of the underlying natural moisture gradient, and that this effect may result from differential responses of seedling guilds to resource availability. Water supply was more important than seed supply in determining seedling establishment, suggesting that appropriate sites for regeneration are a factor limiting seedling success. This is the first study that shows that the episodic supply of microsites for recruitment could influence species richness in the highly threatened and biodiverse longleaf pine savanna.     

Differences in nitrogen use strategies between native and exotic tree species: predicting impacts on invaded ecosystems

Background and aim

Exotic plant species can alter the nitrogen cycle in invaded ecosystems. We assess the differences in nitrogen use strategies and litter production and dynamics among three native riparian trees (Fraxinus angustifolia, Populus alba and Ulmus minor) and three co-occurring exotics (Ailanthus altissima, Robinia pseudoacacia and Ulmus pumila), currently spreading throughout river banks in inner Spain. We aim to predict the ecological consequences of a replacement of the natives by the exotics.

Methods

We compared the leaf lifespan, nitrogen resorption efficiency in leaves, nitrogen mean residence time, amount and timing of litter production and amount of nitrogen returned to soils between these native and exotic species.

Results

We found differences among species in all the variables measured, but not between native or exotic origins. Species were ranked from the most to the least conservative nitrogen use strategy as follows: U. pumila was the most conservative species, followed by the three natives (with an intermediate strategy), A. altissima and finally by the nitrogen-fixer R. pseudoacacia. The studied exotic species would produce contrasting impacts on the nitrogen cycle upon invasion.

Conclusions

On the basis of our results, we predict little impacts on the nitrogen cycle if U. pumila dominates the landscape. Despite being nitrogen-fixer R. pseudocacia would not increase soil nitrogen availability in the study area due to its low litter production and litter decomposition rates. In contrast, we predict an increase in nitrogen availability of soils upon A. altissima invasion, as this species produces a high amount of nitrogen rich and labile litter. This study offers a striking example of the contingencies involved in predicting the ecosystem impacts of exotic plant invasion.     

Effects of native species diversity and resource additions on invader impact

Theory and empirical work have demonstrated that diverse communities can inhibit invasion. Yet, it is unclear how diversity influences invader impact, how impact varies among exotics, and what the relative importance of diversity is versus extrinsic factors that themselves can influence invasion. To address these issues, we established plant assemblages that varied in native species and functional richness and crossed this gradient in diversity with resource (water) addition. Identical assemblages were either uninvaded or invaded with one of three exotic forbs: spotted knapweed (Centaurea maculosa), dalmatian toadflax (Linaria dalmatica), or sulfur cinquefoil (Potentilla recta). To determine impacts, we measured the effects of exotics on native biomass and, for spotted knapweed, on soil moisture and nitrogen levels. Assemblages with high species richness were less invaded and less impacted than less diverse assemblages. Impact scaled with exotic biomass; spotted knapweed had the largest impact on native biomass compared with the other exotics. Although invasion depressed native biomass, the net result was to increase total community yield. Water addition increased invasibility (for knapweed only) but had no effect on invader impact. Together, these results suggest that diversity inhibits invasion and reduces impact more than resource additions facilitate invasion or impact.     

Effects of body size and resource availability on dispersal in a native and a non-native estuarine snail

I manipulated snail densities of two coexisting species of salt marsh snail, Cerithidea californica Haldeman (native) and Batillaria attramentaria Sowerby (non-indigenous) to investigate how resource levels set by intraspecific competition may influence dispersal rates. I used two distinct size classes of the snails (mature and immature) to determine if the effects of competition on dispersal differed between developmental stages of a consumer. Dispersal attempts were measured within enclosure pens by counting snails climbing the sides of the enclosure. The influence of snail density per se and resource levels (which were set by snail densities) on dispersal rates were separated by comparing responses of snails to density before and after resources became depleted. For large snails of both species, dispersal increased as resource levels decreased, supporting the hypothesis that competition influences dispersal rates. Small snails of both species, in contrast, always dispersed at relatively higher rates than larger individuals, but were not influenced by variation in resource levels. This result corroborates other studies that have shown reduced competition in these species at smaller size, and suggests that another mechanism, such as genetically coded behavior to disperse when young, influences their behavior. Previous experiments demonstrated Batillaria's superior resource conversion efficiency; therefore, I had hypothesized that for any given resource level, Cerithidea would disperse more, because it was more affected by resource availability. Adult Batillaria, however, responded more sensitively to resource levels (i.e., dispersed more at any given resource level) than Cerithidea. This counter-intuitive result illustrates the potential importance of genetic limitations on behavioral responses available to a species. Constraints on behavioral responses may have been accentuated since Batillaria is a non-indigenous species whose evolved behavioral responses are not necessarily adapted to its present, non-native environment.     

Effects of resource availability on plant recruitment at the community level in a Mediterranean mountain ecosystem

Coexisting plant species usually differ in resource requirements, which may also vary within species at successive demographic stages. Such differences become extremely important during the early life stages, since these are the most critical phases in woody-species recruitment, they depend heavily on resources, and they may determine future community composition. Under a global-change scenario, where climatic conditions, nutrient availability, and habitat characteristics are expected to be altered, it is difficult to predict the way in which plant recruitment will be affected. To understand the impact of different global-change drivers on community recruitment, we sowed a set of species representative of the different successional groups of a complete Mediterranean woody community under field conditions, and studied their emergence, growth, and survival along the main resource gradients of light, water, and nutrients. The light and nutrient gradients followed the natural range of conditions in the study area, but water availability was manipulated to simulate three contrasting climatic scenarios: wetter, drier, and current conditions. Structural equation modelling was used to provide a comprehensive analysis of the factors and relations governing plant recruitment. Overall, seedling emergence was determined directly by light; growth was determined by light and summer soil moisture; and survival was determined by summer soil moisture. Light was the main factor indirectly affecting the demographic stages of all species. However, the magnitude of the direct and indirect relationships varied among species. Particularly, species differed in their response to the expected drier climatic conditions, some (e.g. Pinus sylvestris, Acer opalus) being much more vulnerable than others (e.g. Cytisus scoparius, Salvia lavandulifolia). These differential responses could translate as major shifts in the structure of the overall plant community. Our results support the idea that the analysis of complex relations among essential resources is critical for accurate forecasts of the impact of climate change on community dynamics.     

Effects of simulated herbivory and resource availability on the invasive plant, Alternanthera philoxeroides in different habitats

In biological control programs, the insect natural enemy’s ability to suppress the plant invader may be affected by abiotic factors, such as resource availability, that can influence plant growth and reproduction. Understanding plant tolerance to herbivory under different environmental conditions will help to improve biocontrol efficacy. The invasive alligator weed (Alternanthera philoxeroides) has been successfully controlled by natural enemies in many aquatic habitats but not in terrestrial environments worldwide. This study examined the effects of different levels of simulated leaf herbivory on the growth of alligator weed at two levels of fertilization and three levels of soil moisture (aquatic, semi-aquatic, and terrestrial habitats). Increasing levels of simulated (manual) defoliation generally caused decreases in total biomass in all habitats. However, the plant appeared to respond differently to high levels of herbivory in the three habitats. Terrestrial plants showed the highest below–above ground mass ratio (R/S), indicating the plant is more tolerant to herbivory in terrestrial habitats than in aquatic habitats. The unfertilized treatment exhibited greater tolerance than the fertilized treatment in the terrestrial habitat at the first stage of this experiment (day 15), but fertilizer appears not to have influenced tolerance at the middle and last stages of the experiment. No such difference was found in semi-aquatic and aquatic habitats. These findings suggest that plant tolerance is affected by habitats and soil nutrients and this relationship could influence the biological control outcome. Plant compensatory response to herbivory under different environmental conditions should, therefore, be carefully considered when planning to use biological control in management programs against invasive plants.     

Fungal endophytes of native Gossypium species in Australia

Fungal endophytes of 17 genera were found in stems of four native Gossypium species (G. australe, G. bickii, G. nelsonii, G. sturtianum) collected from inland areas in Queensland, the Northern Territory, and South Australia in 2001. Phoma, Alternaria, Fusarium, Botryosphaeria, Dichomera, and Phomopsis were common, accounting for 58, 18, 11, 3, 1, and 1 % of the 281 recovered isolates, respectively, and occurring in 47, 29, 19, 5, 5, and 4 % of the 79 sampled populations. Among the four Gossypium species in Queensland and the Northern Territory, Alternaria spp. and Fusarium spp. had the greatest recovery frequency in G. bickii stems. The recovery frequencies of Phoma spp. and Alternaria spp. were significantly greater in the G. sturtianum stems collected from South Australia than in those from Queensland and the Northern Territory. Pathogenicity of 42 representative isolates was tested on cultivated cotton (G. hirsutum). All isolates caused some localized discoloration in stem tissue when inoculation was conducted with the stem puncturing method, but none of the isolates could induce any foliar symptoms during the five-week experimental period by either inoculation method (root dipping or stem puncturing), suggesting that the endophytic fungi of native Gossypium species are unlikely sources of cotton pathogens.     

Dispersal and recruitment of a canopy-forming intertidal alga: the relative roles of propagule availability and post-settlement processes

The daily settlement of eggs and zygotes of the monoecious brown alga Pelvetia compressa (J. Agardh) De Toni was measured on artificial substrata in areas inside and outside patches of adults in the high intertidal zone of central California. Settlement was generally 1–2 orders of magnitude higher under the adult canopy. This pattern seems to be due to the synchronous release of gametes during the daytime low tide. The release of gametes also appears periodic over longer time scales (e.g., 3- and 14-day cycles). In spite of the high availability of propagules under the adult canopy, juveniles were most abundant outside patches, where propagule availability was lower. In both areas, juveniles were disproportionately associated with patches of a red algal turf [primarily Endocladia muricata (Postels & Ruprecht) J. Agardh and Masticarpus papillata (C. Agardh) Kützing]. The turf, which is less common under the P. compressa canopy, may offer protection from dislodgment, grazing, and/or desiccation and thus facilitate recruitment at this site. Overall, post-settlement processes appear more important in determining population structure than does the availability of propagules in areas in and around patches of adults. However, the apparent small range of dispersal of P. compressa may make propagule availability an important limitation to the establishment of new populations and may restrict gene flow between populations. Received: 31 October 1997 / Accepted: 31 August 1998     

Impact of resource availability on species composition and diversity in freshwater nematodes

This study investigates the long-term effects of resource availability in a freshwater nematode community. We carried out a mesocosm experiment where natural nematode communities were exposed to nutrient addition/depletion over 2 years. Compared to the nutrient-addition treatment, species richness and diversity were strongly reduced upon nutrient depletion. The functional group of bacterial feeders particularly suffered severely from nutrient depletion. The decrease in diversity of bacterial feeders was linked to reduced species richness and diversity of large omnivorous species, as predicted by trophic-dynamic models. Tilmans (1976) statement, that under low nutrient levels the best competitor dominates the system, was applicable in our system. Upon nutrient depletion, resource depletion led to a monoculture of 1 small bacterial feeder, but even after 2 years of resource depletion, up to 16 species still coexisted. Our results provide strong evidence that freshwater nematode systems can be regulated by nutrient competition.     

Exotic plants increase and native plants decrease with loss of foundation species in sagebrush steppe

Dominant plant species, or foundation species, are recognized to have a disproportionate control over resources in ecosystems, but few studies have evaluated their relationship to exotic invasions. Loss of foundation species could increase resource availability to the benefit of exotic plants, and could thereby facilitate invasion. The success of exotic plant invasions in sagebrush steppe was hypothesized to benefit from increased available soil water following removal of sagebrush (Artemisia tridentata), a foundation species. We examined the effects of sagebrush removal, with and without the extra soil water made available by exclusion of sagebrush, on abundance of exotic and native plants in the shrub steppe of southern Idaho, USA. We compared plant responses in three treatments: undisturbed sagebrush steppe; sagebrush removed; and sagebrush removed plus plots covered with “rainout” shelters that blocked winter-spring recharge of soil water. The third treatment allowed us to examine effects of sagebrush removal alone, without the associated increase in deep-soil water that is expected to accompany removal of sagebrush. Overall, exotic herbs (the grass Bromus tectorum and four forbs) were 3–4 times more abundant in shrub-removal and 2 times more abundant in shrub-removal + rainout-shelter treatments than in the control treatment, where sagebrush was undisturbed. Conversely, native forbs were only about half as abundant in shrub removal compared to control plots. These results indicate that removal of sagebrush facilitates invasion of exotic plants, and that increased soil water is one of the causes. Our findings suggest that sagebrush plays an important role in reducing invasions by exotic plants and maintaining native plant communities, in the cold desert we evaluated.     

Effects of resource pulse magnitude on nutrient availability,productivity, stability,and food web dynamics in experimental aquatic ecosystems

Oceanographic processes play a key role in influencing the structure of the marine planktonic ecosystems. Taking advantage of the quasi-simultaneous collection of a large ichthyoplanktonic dataset in different regions of the Central Mediterranean Sea (Italian/Maltese, Tunisian and Libyan waters), this study aimed at the identification of the main environmental drivers that control the structure of the larval fish assemblages. Spatial distribution and taxa composition were related to physical forcings (geostrophic currents and wind stress) and environmental conditions (bottom depth, temperature, salinity, chlorophyll-a concentration). ANOSIM and SIMPER identified contribution of fish taxa to the average Bray–Curtis dissimilarity among regions. In Italian and Libyan waters, two assemblages (neritic and oceanic) were identified, while a mixed assemblage characterized only some stations. Two neritic and one oceanic assemblages were discriminated in Tunisian waters. Random Forest classification model highlighted the essential role of the bathymetry, while Lagrangian simulations evidenced the action of the hydrodynamics in mixing neritic and oceanic assemblages in the Italian/Maltese and partially in Libyan waters. These findings highlighted the importance of the multidisciplinary approach and shed light on the potential value of the ichthyoplanktonic surveys for the assessment of the state of the marine ecosystem and the conservation of the fishery resources.     

Global resource acquisition patterns of invasive and native plant species do not hold at the regional scale in Mediterranean type ecosystems

Invasive species may outperform native species by acquiring more resources or by efficiently using limited resources. Studies comparing leaf traits as a metric of carbon capture strategies in native and invasive species have come to different conclusions. Some studies suggest that invasive species are better at acquiring resources, but that native and invasive species use resources similarly. Other studies have found that native and invasive species differ in resource use efficiency, which implies different biochemical or physiological mechanisms of carbon capture. To resolve this debate, we examined relationships among four leaf traits (photosynthetic rate, specific leaf area, foliar nitrogen, foliar phosphorus) in co-occurring native and invasive species from eight plant communities across five Mediterranean-climate ecosystems. We performed standardized major axis regression for all trait combinations within and across sites, testing for slope homogeneity and shifts in elevation (y-intercept) or along a common slope between species groups. Across the global dataset, native and invasive species had similar carbon capture strategies (i.e., similar slopes), with invasive species occupying a position of greater resource acquisition. However, these patterns did not hold when regions were analyzed individually. Regional differences may be driven by differences in life form between native and invasive species, and variation in soil resource availability among regions. Our context-dependent results reveal not only that management of invasive species will differ across regions but also that global comparisons of invasive and native species can be misleading.     

Honeydew feeding in the solitary bee Osmia bicornis as affected by aphid species and nectar availability

Like honey bees (Apis mellifera), non-Apis bees could exploit honeydew as a carbohydrate source. In addition to providing carbohydrates, this may expose them to potentially harmful plant products secreted in honeydew. However, knowledge on honeydew feeding by solitary bees is very scarce. Here we determine whether the polylectic solitary bee Osmia bicornis (=O. rufa) collects honeydew under semi-field conditions, and whether this is affected by aphid species and presence of floral nectar. Bees were provided with oilseed rape plants containing flowers and/or colonies of either Myzus persicae or Brevicoryne brassicae. We used the total sugar level of the bee crop as a measure of the individual's nutritional state and the oligosaccharide erlose as indicator for honeydew consumption. Erlose was present in honeydews from both aphid species, while absent in oilseed rape nectar, nor being synthesized by O. bicornis. When bees were confined to a single honeydew type as the only carbohydrate source, consumption of M. persicae honeydew was confirmed for 47% of the bees and consumption of B. brassicae honeydew for only 3%. Increased mortality in the latter treatment provided further evidence that B. brassicae honeydew is an unsuitable food source for O. bicornis. All bees that were given the choice between honeydew and floral nectar showed significantly increased total sugar levels. However, the fact that no erlose was detected in these bees indicates that honeydew was not consumed when suitable floral nectar was available. This study demonstrates that honeydew exploitation by O. bicornis is dependent on honeydew type and the presence of floral nectar.     

Effects of resource availability on integration and clonal growth inMaianthemum bifolium

The aim of this study was to investigate whether resource availability affects the degree of physiological integration and the growth pattern of interconnected ramets in the clonal plantMaianthemum bifolium (L.)F.W. Schmidt (Liliaceae), a rhizomatous herb of European forests, by studying it at two contrasting South Swedish beech forest sites termed “poor” and “rich”. The degree of physiological integration was studied by tracing the pattern of¹⁴C translocation and in a cutting experiment involving rhizome severing and defoliation treatments. The size of the plants, growth of new rhizomes, branching frequency, distance between shoots and the internode length were compared. The plants were larger, rhizomes had greater specific mass (mg mm^?1), internodes were shorter and branching frequency higher at the rich site. The cutting treatments reduced the growth of new rhizomes at both sites, and new rhizome segments had lower specific mass in treated plants than in controls, showing the importance of physiological integration for new growth. Translocation of¹⁴C in May showed that the young rhizome tip was a strong sink for carbon. Basipetal translocation to older portions of the rhizome system was greater at the rich site than at the poor site. In September, four months after labelling, the rhizome tips were still strongly labelled with¹⁴C and basipetal translocation had increased at both sites. Plants at the rich site appeared to translocate larger amounts of¹⁴C basipetally than plants at the poor site. It is concluded thatM. bifolium shows a plastic response to resource availability by varying rhizome growth and branching frequency, but the degree of physiological integration is probably only indirectly affected through an increased number of sinks (new rhizome branches) along the ramet system at the rich site.     

Effects of nutrient supply,light availability and herbivory on the growth of heather and three competing grass species

Grasses are becoming more abundant in areas in NE Scotland which until recently were dominated by heather (Calluna vulgaris). However, it is not clear if grasses are aggressive competitors which are now able to outcompete the dwarf shrub due to changes in environmental factors (such as grazing pressure and increasing nutrient inputs), or just opportunistic invaders, occupying gaps in the canopy which occur when heather reaches the degenerate stage. Experiments in turves and in field plots were carried out in order to investigate the performance of three grass species, Nardus stricta, Deschampsia cespitosa and Deschampsia flexuosa growing in competition with heather. These three species were selected because they differ in their nutrient requirements, palatability to herbivores and tolerance of shading. The grasses were planted in heather canopies of different structure, either turves of heather of different height and age, or moorland plots with or without heavy grazing by sheep and deer. Fertiliser (NPK) was applied to half the experimental plants. The growth of the grass species and the heather in response to the fertiliser and grazing treatments was measured, together with the light levels penetrating the canopy and received by the grass plants.Results indicated that heather was likely to be outcompeted by grasses only when there are gaps in the canopy, resulting either from heavy grazing or from the heather being in the mature or degenerate phase. Fertiliser enhanced plant growth whereas fencing out herbivores led to strong competition for light as the heather canopy closed. It is concluded that grasses require gaps in the canopy to successfully invade heather moorland, or they tend to be shaded out. Thus better management of heather moorlands to maintain a dense canopy structure may help to preserve heather cover even under increasing nutrient inputs.     

Canopy cover type,and not fine-scale resource availability,explains native and exotic species richness in a landscape affected by anthropogenic fires and posterior land-use change

Anthropogenic fires and land-use change, including the conversion from native to exotic species canopies, are two major types of disturbances that strongly affect the functioning of forest ecosystems around the world. These disturbances alter the resource availability for plants, which may lead to changes in species richness. Here we examined the relative effects of canopy cover type, light availability and soil nutrient (N and P) availability on species richness, including invasive species, at different post-fire plant systems. Additionally, we tested the resource heterogeneity hypothesis (RHH) for plant diversity, which proposes that diversity is higher in habitats with spatially heterogeneous resources. We evaluated four different canopy cover types, including mature and second-growth Nothofagus pumilio forests, treeless prairie, Pinus sylvestris afforestations, all of which were converted from mature N. pumilio forests. Using generalized mixed-effects model correlations, we determined (1) the relative influence of canopy cover type, light and soil nutrient availability on understory species richness and (2) the relationship between species richness and resource heterogeneity. We found that canopy cover type was the factor that best explained species richness, much more than fine-scale light and soil nutrient availability. Additionally, we found that the more homogeneous the light environment the higher the number of exotic species (mainly found in the prairie where the highest light intensity occurred), which is contrary to what the RHH states. In conclusion, canopy cover type, a stand-scale driver, and not fine-scale resource (light, N and P) availability, was most important for explaining native and exotic (including invasive) species understory richness in a landscape affected by anthropogenic fires and posterior land-use change.