A global analysis of bidirectional interactions in alpine plant communities shows facilitators experiencing strong reciprocal fitness costs.

Facilitative interactions are defined as positive effects of one species on another, but bidirectional feedbacks may be positive, neutral, or negative. Understanding the bidirectional nature of these interactions is a fundamental prerequisite for the assessment of the potential evolutionary consequences of facilitation. In a global study combining observational and experimental approaches, we quantified the impact of the cover and richness of species associated with alpine cushion plants on reproductive traits of the benefactor cushions. We found a decline in cushion seed production with increasing cover of cushion-associated species, indicating that being a benefactor came at an overall cost. The effect of cushion-associated species was negative for flower density and seed set of cushions, but not for fruit set and seed quality. Richness of cushion-associated species had positive effects on seed density and modulated the effects of their abundance on flower density and fruit set, indicating that the costs and benefits of harboring associated species depend on the composition of the plant assemblage. Our study demonstrates 'parasitic' interactions among plants over a wide range of species and environments in alpine systems, and we consider their implications for the possible selective effects of interactions between benefactor and beneficiary species.

Ontogenetic shifts in plant interactions vary with environmental severity and affect population structure.

Environmental conditions and plant size may both alter the outcome of inter-specific plant-plant interactions, with seedlings generally facilitated more strongly than larger individuals in stressful habitats. However, the combined impact of plant size and environmental severity on interactions is poorly understood. Here, we tested explicitly for the first time the hypothesis that ontogenetic shifts in interactions are delayed under increasingly severe conditions by examining the interaction between a grass, Agrostis magellanica, and a cushion plant, Azorella selago, along two severity gradients. The impact of A. selago on A. magellanica abundance, but not reproductive effort, was related to A. magellanica size, with a trend for delayed shifts towards more negative interactions under greater environmental severity. Intermediate-sized individuals were most strongly facilitated, leading to differences in the size-class distribution of A. magellanica on the soil and on A. selago. The A. magellanica size-class distribution was more strongly affected by A. selago than by environmental severity, demonstrating that the plant-plant interaction impacts A. magellanica population structure more strongly than habitat conditions. As ontogenetic shifts in plant-plant interactions cannot be assumed to be constant across severity gradients and may impact species population structure, studies examining the outcome of interactions need to consider the potential for size- or age-related variation in competition and facilitation.

Horizontal, but not vertical, biotic interactions affect fine-scale plant distribution patterns in a low-energy system.

Studies of species range determinants have traditionally focused on abiotic variables (typically climatic conditions), and therefore the recent explicit consideration of biotic interactions represents an important advance in the field. While these studies clearly support the role of biotic interactions in shaping species distributions, most examine only the influence of a single species and/or a single interaction, failing to account for species being subject to multiple concurrent interactions. By fitting species distribution models (SDMs), we examine the influence of multiple vertical (i.e., grazing, trampling, and manuring by mammalian herbivores) and horizontal (i.e., competition and facilitation; estimated from the cover of dominant plant species) interspecific interactions on the occurrence and cover of 41 alpine tundra plant species. Adding plant-plant interactions to baseline SDMs (using five field-quantified abiotic variables) significantly improved models' predictive power for independent data, while herbivore-related variables had only a weak influence. Overall, abiotic variables had the strongest individual contributions to the distribution of alpine tundra plants, with the importance of horizontal interaction variables exceeding that of vertical interaction variables. These results were consistent across three modeling techniques, for both species occurrence and cover, demonstrating the pattern to be robust. Thus, the explicit consideration of multiple biotic interactions reveals that plant-plant interactions exert control over the fine-scale distribution of vascular species that is comparable to abiotic drivers and considerably stronger than herbivores in this low-energy system.

Climate change and elevational diversity capacity: do weedy species take up the slack?

Climate change leads to species range shifts and consequently to changes in diversity. For many systems, increases in diversity capacity have been forecast, with spare capacity to be taken up by a pool of weedy species moved around by humans. Few tests of this hypothesis have been undertaken, and in many temperate systems, climate change impacts may be confounded by simultaneous increases in human-related disturbance, which also promote weedy species. Areas to which weedy species are being introduced, but with little human disturbance, are therefore ideal for testing the idea. We make predictions about how such diversity capacity increases play out across elevational gradients in non-water-limited systems. Then, using modern and historical data on the elevational range of indigenous and naturalized alien vascular plant species from the relatively undisturbed sub-Antarctic Marion Island, we show that alien species have contributed significantly to filling available diversity capacity and that increases in energy availability rather than disturbance are the probable underlying cause.

Remnant fragments within an agricultural matrix enhance conditions for a rodent host and its fleas.

Habitat fragmentation can adversely impact biodiversity, although where remnant fragments of natural vegetation provide favourable conditions the negative effects of fragmentation may be mitigated. Host-parasite systems in fragmented areas have only recently been examined, with parasites generally showing higher prevalence and richness in fragments, mediated by changes in host density. However, the effect of fragmentation on parasite body size and fecundity remains poorly investigated. Thus, here we compared the body size and condition of a generalist rodent host, Rhabdomys pumilio and the body size of 2 common flea species between pristine natural areas and remnant fragments within agriculture areas. Host body length, weight and body condition values were significantly larger in fragments than in pristine natural vegetation. Listropsylla agrippinae fleas showed the same pattern, being significantly larger in fragments, while Chiastopsylla rossi fleas did not differ in size between fragments and natural areas. The differential response of the 2 flea species may reflect the strength of association between the host and parasite, with the former spending a greater proportion of its lifespan on the host. Therefore, in this study agriculture fragments provide better conditions for both an opportunistic rodent and a closely associated flea species.

Plant dispersal in the sub-Antarctic inferred from anisotropic genetic structure.

Climatic conditions and landscape features often strongly affect species' local distribution patterns, dispersal, reproduction and survival and may therefore have considerable impacts on species' fine-scale spatial genetic structure (SGS). In this study, we demonstrate the efficacy of combining fine-scale SGS analyses with isotropic and anisotropic spatial autocorrelation techniques to infer the impact of wind patterns on plant dispersal processes. We genotyped 1304 Azorella selago (Apiaceae) specimens, a wind-pollinated and wind-dispersed plant, from four populations distributed across sub-Antarctic Marion Island. SGS was variable with Sp values ranging from 0.001 to 0.014, suggesting notable variability in dispersal distance and wind velocities between sites. Nonetheless, the data supported previous hypotheses of a strong NW-SE gradient in wind strength across the island. Anisotropic autocorrelation analyses further suggested that dispersal is strongly directional, but varying between sites depending on the local prevailing winds. Despite the high frequency of gale-force winds on Marion Island, gene dispersal distance estimates (σ) were surprisingly low (<10 m), most probably because of a low pollen dispersal efficiency. An SGS approach in association with isotropic and anisotropic analyses provides a powerful means to assess the relative influence of abiotic factors on dispersal and allow inferences that would not be possible without this combined approach.

Ecologically relevant measures of tolerance to potentially lethal temperatures.

The acute thermal tolerance of ectotherms has been measured in a variety of ways; these include assays where organisms are shifted abruptly to stressful temperatures and assays where organisms experience temperatures that are ramped more slowly to stressful levels. Ramping assays are thought to be more relevant to natural conditions where sudden abrupt shifts are unlikely to occur often, but it has been argued that thermal limits established under ramping conditions are underestimates of true thermal limits because stresses due to starvation and/or desiccation can arise under ramping. These confounding effects might also impact the variance and heritability of thermal tolerance. We argue here that ramping assays are useful in capturing aspects of ecological relevance even though there is potential for confounding effects of other stresses that can also influence thermal limits in nature. Moreover, we show that the levels of desiccation and starvation experienced by ectotherms in ramping assays will often be minor unless the assays involve small animals and last for many hours. Empirical data illustrate that the combined effects of food and humidity on thermal limits under ramping and sudden shifts to stressful conditions are unpredictable; in Drosophila melanogaster the presence of food decreased rather than increased thermal limits, whereas in Ceratitis capitata they had little impact. The literature provides examples where thermal limits are increased under ramping presumably because of the potential for physiological changes leading to acclimation. It is unclear whether heritabilities and population differentiation will necessarily be lower under ramping because of confounding effects. Although it is important to clearly define experimental methods, particularly when undertaking comparative assessments, and to understand potential confounding effects, thermotolerance assays based on ramping remain an important tool for understanding and predicting species responses to environmental change. An important area for further development is to identify the impact of rates of temperature change under field and laboratory conditions.

Cryptic spatial aggregation of the cushion plant Azorella selago (Apiaceae) revealed by a multilocus molecular approach suggests frequent intraspecific facilitation under sub-Antarctic conditions.

PREMISE OF THE STUDY: In abiotically severe habitats, intraspecific aggregations can increase species' fitness by ameliorating stressful environmental factors. However, the difficulty of identifying individual plants in some growth forms makes the measurements of intraspecific aggregation, and therefore the assessment of intraspecific facilitation, problematic. In this study, we examined the genotype composition within cushions of Azorella selago, a sub-Antarctic cushion plant, to investigate the potential extent of intraspecific facilitation. METHODS: The study was performed on Marion Island, South Africa. Two to eight samples were collected from 42 A. selago cushions, comprising eight different growth forms. Samples were genotyped using seven microsatellite markers. KEY RESULTS: We showed that all cushion shapes, with the exception of small cushions, may be comprised of more than one genetically distinct individual. CONCLUSIONS: Under harsh sub-Antarctic conditions, intraspecific aggregation between A. selago individuals appears common and may be driven by the positive impacts of environmental amelioration.

Interspecific facilitation mediates the outcome of intraspecific interactions across an elevational gradient.

Where interspecific facilitation favors the establishment of high densities of a beneficiary species, strong intraspecific competition may subsequently impede beneficiary performance. Consequently, the negative influence of intraspecific competition between beneficiary individuals could potentially outweigh the positive influence of interspecific facilitation when, for example, higher densities of a beneficiary are negated by the negative effect of crowding on beneficiary reproduction. The aim of this study was, therefore, to examine the impact of an interspecific interaction on the outcome of intraspecific interactions within the context of plant-plant facilitation. We used the cushion-forming Azorella selago and a commonly co-occurring dominant perennial grass species, Agrostis magellanica, on sub-Antarctic Marion Island as a model system. We assessed the impact of an interspecific interaction (between A. selago and A. magellanica) on the outcome of intraspecific interactions (between A. magellanica individuals), by testing if the impact of A. magellanica density on A. magellanica performance is mediated by its interaction with A. selago. We observed evidence for competition among A. magellanica conspecifics, with a decreasing proportion of A. magellanica individuals being reproductive under higher conspecific density. This negative intraspecific effect was greater on A. selago than on the adjacent substrate, suggesting that the facilitative effect of A. selago changes the intensity of intraspecific interactions between A. magellanica individuals. However, experimentally reducing A. magellanica density did not affect the species' performance. We also observed that the effect of A. selago on A. magellanica was positive, and despite the negative effect of intraspecific density on the proportion of reproductive A. magellanica individuals, the net reproductive effort of A. magellanica (i.e., the density of reproductive individuals) was significantly greater on A. selago than on the adjacent substrate. These results highlight that, in abiotically severe environments, the positive effects of interspecific facilitation by a benefactor species may outweigh the negative effects of intraspecific competition among beneficiaries. More broadly, these results suggest that both positive inter- and intraspecific biotic interactions may be key to consider when examining spatial and temporal variation in species' performance.

Consistent trait-environment relationships within and across tundra plant communities.

A fundamental assumption in trait-based ecology is that relationships between traits and environmental conditions are globally consistent. We use field-quantified microclimate and soil data to explore if trait-environment relationships are generalizable across plant communities and spatial scales. We collected data from 6,720 plots and 217 species across four distinct tundra regions from both hemispheres. We combined these data with over 76,000 database trait records to relate local plant community trait composition to broad gradients of key environmental drivers: soil moisture, soil temperature, soil pH and potential solar radiation. Results revealed strong, consistent trait-environment relationships across Arctic and Antarctic regions. This indicates that the detected relationships are transferable between tundra plant communities also when fine-scale environmental heterogeneity is accounted for, and that variation in local conditions heavily influences both structural and leaf economic traits. Our results strengthen the biological and mechanistic basis for climate change impact predictions of vulnerable high-latitude ecosystems.

Extrapolating population size from the occupancy-abundance relationship and the scaling pattern of occupancy.

The estimation of species abundances at regional scales requires a cost-efficient method that can be applied to existing broadscale data. We compared the performance of eight models for estimating species abundance and community structure from presence-absence maps of the southern African avifauna. Six models were based on the intraspecific occupancy-abundance relationship (OAR); the other two on the scaling pattern of species occupancy (SPO), which quantifies the decline in species range size when measured across progressively finer scales. The performance of these models was examined using five tests: the first three compared the predicted community structure against well-documented macroecological patterns; the final two compared published abundance estimates for rare species and the total regional abundance estimate against predicted abundances. Approximately two billion birds were estimated as occurring in South Africa, Lesotho, and Swaziland. SPO models outperformed the OAR models, due to OAR models assuming environmental homogeneity and yielding scale-dependent estimates. Therefore, OAR models should only be applied across small, homogenous areas. By contrast, SPO models are suitable for data at larger spatial scales because they are based on the scale dependence of species range size and incorporate environmental heterogeneity (assuming fractal habitat structure or performing a Bayesian estimate of occupancy). Therefore, SPO models are recommended for assemblage-scale regional abundance estimation based on spatially explicit presence-absence data.

Species-energy relationships of indigenous and invasive species may arise in different ways - a demonstration using springtails.

Although the relationship between species richness and available energy is well established for a range of spatial scales, exploration of the plausible underlying explanations for this relationship is less common. Speciation, extinction, dispersal and environmental filters all play a role. Here we make use of replicated elevational transects and the insights offered by comparing indigenous and invasive species to test four proximal mechanisms that have been offered to explain relationships between energy availability, abundance and species richness: the sampling mechanism (a null expectation), and the more individuals, dynamic equilibrium and range limitation mechanisms. We also briefly consider the time for speciation mechanism. We do so for springtails on sub-Antarctic Marion Island. Relationships between energy availability and species richness are stronger for invasive than indigenous species, with geometric constraints and area variation playing minor roles. We reject the sampling and more individuals mechanisms, but show that dynamic equilibrium and range limitation are plausible mechanisms underlying these gradients, especially for invasive species. Time for speciation cannot be ruled out as contributing to richness variation in the indigenous species. Differences between the indigenous and invasive species highlight the ways in which deconstruction of richness gradients may usefully inform investigations of the mechanisms underlying them. They also point to the importance of population size-related mechanisms in accounting for such variation. In the context of the sub-Antarctic our findings suggest that warming climates may favour invasive over indigenous species in the context of changes to elevational distributions, a situation found for vascular plants, and predicted for springtails on the basis of smaller-scale manipulative field experiments.

The influence of life history characteristics on flea (Siphonaptera) species distribution models.

BACKGROUND: Ectoparasites exhibit pronounced variation in life history characteristics such as time spent on the host and host range. Since contemporary species distribution (SD) modelling does not account for differences in life history, the accuracy of predictions of current and future species' ranges could differ significantly between life history groups. RESULTS: SD model performance was compared between 21 flea species that differ in microhabitat preferences and level of host specificity. Distribution models generally performed well, with no significant differences in model performance based on either microhabitat preferences or host specificity. However, the relative importance of predictor variables was significantly related to host specificity, with the distribution of host-opportunistic fleas strongly limited by thermal conditions and host-specific fleas more associated with conditions that restrict their hosts' distribution. The importance of temperature was even more pronounced when considering microhabitat preference, with the distribution of fur fleas being strongly limited by thermal conditions and nest fleas more associated with variables that affect microclimatic conditions in the host nest. CONCLUSIONS: Contemporary SD modelling, that includes climate and landscape variables, is a valuable tool to study the biogeography and future distributions of fleas and other parasites taxa. However, consideration of life history characteristics is cautioned as species may be differentially sensitive to environmental conditions.

Thermal biology, population fluctuations and implications of temperature extremes for the management of two globally significant insect pests.

The link between environmental temperature, physiological processes and population fluctuations is a significant aspect of insect pest management. Here, we explore how thermal biology affects the population abundance of two globally significant pest fruit fly species, Ceratitis capitata (medfly) and C. rosa (Natal fruit fly), including irradiated individuals and those expressing a temperature sensitive lethal (tsl) mutation that are used in the sterile insect technique. Results show that upper and lower lethal temperatures are seldom encountered at the field sites, while critical minimum temperatures for activity and lower developmental thresholds are crossed more frequently. Estimates of abundance revealed that C. capitata are active year-round, but abundance declines markedly during winter. Temporal autocorrelation of average fortnightly trap captures and of development time, estimated from an integrated model to calculate available degree days, show similar seasonal lags suggesting that population increases in early spring occur after sufficient degree-days have accumulated. By contrast, population collapses coincide tightly with increasing frequency of low temperature events that fall below critical minimum temperatures for activity. Individuals of C. capitata expressing the tsl mutation show greater critical thermal maxima and greater longevity under field conditions than reference individuals. Taken together, this evidence suggests that low temperatures limit populations in the Western Cape, South Africa and likely do so elsewhere. Increasing temperature extremes and warming climates generally may extend the season over which these species are active, and could increase abundance. The sterile insect technique may prove profitable as climates change given that laboratory-reared tsl flies have an advantage under warmer conditions.

Spatial variation in plant interactions across a severity gradient in the sub-Antarctic.

The stress-gradient hypothesis predicts that the intensity of interspecific positive interactions increases along environmental severity (i.e. stress and disturbance) gradients faster than the intensity of negative interactions. This study is the first to test if the stress-gradient hypothesis is supported for a location in the climatically extreme and species-poor sub-Antarctic. To do so, we investigate the fine-scale spatial distribution of plant species across altitude- and aspect-related abiotic severity gradients on a scoria cone on Marion Island. A clear altitudinal severity gradient was observed across the scoria cone, with lower temperatures, stronger winds and greater soil movement at higher altitudes. The altitudinal severity gradient was matched by stronger interspecific spatial association between the four dominant species at higher altitudes and in areas of lower vegetation cover. This suggests that, relative to the intensity of competition, the intensity of facilitation is greater under more severe conditions, supporting the stress-gradient hypothesis at the community level (i.e. for multiple pairs of species) and corroborating its usefulness for predicting variation in plant interactions at high latitudes and altitudes. Furthermore, the directional intraspecific aggregation and interspecific association plant cover patterns found within the gradient suggest that protection from the prevailing wind and from burial by loose substrate are the dominant facilitative mechanisms. Thus, plants benefit from the presence of neighbours when they provide shelter and substrate stability, and the relative intensity of this positive interaction is greatest at higher altitudes, and varies between species pairs. This study, therefore, not only provides support for the stress-gradient hypothesis in the sub-Antarctic, but also demonstrates fine-scale directional spatial patterns between multiple species nested within the severity gradient.