Impacts of extreme climatic events on competition during grassland invasions

Phytometers of five C3 and five C4 species were transplanted into three different grasslands to study the effects of extreme climatic events on community invasibility and competition. Single extreme heating (eight hours at 52.5 °C) and rainfall (the equivalent of 100 mm) events in factorial combinations were superimposed on the grassland communities. A novel technique involving portable computer‐controlled chambers was used to create the heating events. In order to generate predictions of response to the extreme climatic events, the 10 phytometer species were categorized on the basis of 12 key plant functional traits. Using principal component analysis, two functional types (FTs) were identified as most likely to be advantaged (FT1, fast‐growing C4 annuals) and disadvantaged (FT2, slower‐growing C3 perennials) by an extreme climatic event. Competition between the resident vegetation and FT1 plus other C4 phytometers was consistently more intense within the exclusively C3 community compared to the dry C3/C4 community or moist C3/C4 community. The single extreme heating event had the greatest impact on competition, lowering the intensity of competition between the phytometers and resident vegetation. Our results indicate that competition is highly important in limiting the invasion of C3 grasslands by C4 species. The FT1 and FT2 responses confirmed predictions based on plant functional traits, whether growing as phytometers or as part of the resident vegetation. Future increases in climatic variability and the incidence of extreme climatic events are expected to suppress C3 competitive dominance and promote invasion of C4 species, in particular, the FT1 species.     

Climate disruption and parasite-host dynamics: patterns and processes associated with warming and the frequency of extreme climatic events

Levels of parasitism and the dynamics of helminth systems is subject to the impact of environmental conditions such that we may expect long term increases in temperature will increase the force of infection and the parasite's basic reproduction number, R0. We postulate that an increase in the force of infection will only lead to an increase in mean intensity of adults when adult parasite mortality is not determined by acquired immunity. Preliminary examination of long term trends of parasites of rabbits and grouse confirm these predictions. Parasite development rate increases with temperature and while laboratory studies indicate this is linear some recent studies indicate that this may be non-linear and would have an important impact on R0. Warming would also reduce the selective pressure for the development of arrestment and this would increase R0 so that in systems like the grouse and Trichostrongylus tenuis this would increase the instability and lead to larger disease outbreaks. Extreme climatic events that act across populations appear important in synchronizing transmission and disease outbreaks, so it is speculated that climate disruption will lead to increased frequency and intensity of disease outbreaks in parasite populations not regulated by acquired immunity.     

To breed or not to breed: a seabird's response to extreme climatic events

Intermittent breeding is an important life-history strategy that has rarely been quantified in the wild and for which drivers remain unclear. It may be the result of a trade-off between survival and reproduction, with individuals skipping breeding when breeding conditions are below a certain threshold. Heterogeneity in individual quality can also lead to heterogeneity in intermittent breeding. We modelled survival, recruitment and breeding probability of the red-footed booby (Sula sula), using a 19 year mark–recapture dataset involving more than 11 000 birds. We showed that skipping breeding was more likely in El-Niño years, correlated with an increase in the local sea surface temperature, supporting the hypothesis that it may be partly an adaptive strategy of birds to face the trade-off between survival and reproduction owing to environmental constraints. We also showed that the age-specific probability of first breeding attempt was synchronized among different age-classes and higher in El-Niño years. This result suggested that pre-breeders may benefit from lowered competition with experienced breeders in years of high skipping probabilities.     

Continuous and discrete extreme climatic events affecting the dynamics of a high-arctic reindeer population

Climate at northern latitudes are currently changing both with regard to the mean and the temporal variability at any given site, increasing the frequency of extreme events such as cold and warm spells. Here we use a conceptually new modelling approach with two different dynamic terms of the climatic effects on a Svalbard reindeer population (the Brøggerhalvøya population) which underwent an extreme icing event (“locked pastures”) with 80% reduction in population size during one winter (1993/94). One term captures the continuous and linear effect depending upon the Arctic Oscillation and another the discrete (rare) “event” process. The introduction of an “event” parameter describing the discrete extreme winter resulted in a more parsimonious model. Such an approach may be useful in strongly age-structured ungulate populations, with young and very old individuals being particularly prone to mortality factors during adverse conditions (resulting in a population structure that differs before and after extreme climatic events). A simulation study demonstrates that our approach is able to properly detect the ecological effects of such extreme climate events.     

Stochastic approach to determine spatial patterns of lizard community on a desert island

One of the principal sources of error in identifying spatial arrangements is autocorrelation, since nearby points in space tend to have more similar values than would be expected by random change. When a Markovian approach is used, spatial arrangements can be measured as a transition probability between occupied and empty spaces in samples that are spatially dependent. We applied a model that incorporates first-order Markov chains to analyse spatial arrangement of numerical dominance, richness, and abundance on a lizard community at different spatial and temporal scales. We hypothesized that if a spatial dependence on abundance and richness exists in a diurnal desert community, then the Markov chains can predict the spatial arrangement. We found that each pair of values was dependent only on its immediate predecessor segment. In this sense, we found intergeneric differences at temporal and spatial scales of recurrence estimates. Also, in desert scrub, species show higher spatial aggregation and had lower species richness than at the island level; the inverse pattern occurred on rocky hillsides. At the species level, Uta stansburiana is the most abundant species in desert scrub, while Sauromalus slevini is the most abundant species on rocky hillsides. This report attempts to understand, using Markovian spatial models, the effect of nearby samples on local abundance and richness on different scales and over several seasons.     

Effects of climatic variation on field metabolism and water relations of desert tortoises

We used the doubly labeled water method to measure the field metabolic rates (FMRs, in kJ kg^?1?day^?1) and water flux rates (WIRs, in ml H₂O?kg^?1?day^?1) of adult desert tortoises (Gopherus agassizii) in three parts of the Mojave Desert in California over a 3.5-year period, in order to develop insights into the physiological responses of this threatened species to climate variation among sites and years. FMR, WIR, and the water economy index (WEI, in ml H₂O?kJ^?1, an indicator of drinking of free water) differed extensively among seasons, among study sites, between sexes, and among years. In high-rainfall years, males had higher FMRs than females. Average daily rates of energy and water use by desert tortoises were extraordinarily variable: 28-fold differences in FMR and 237-fold differences in WIR were measured. Some of this variation was due to seasonal conditions, with rates being low during cold winter months and higher in the warm seasons. However, much of the variation was due to responses to year-to-year variation in rainfall. Annual spring peaks in FMR and WIR were higher in wet years than in drought years. Site differences in seasonal patterns were apparently due to geographic differences in rainfall patterns (more summer rain at eastern Mojave sites). In spring 1992, during an El Niño (ENSO) event, the WEI was greater than the maximal value obtainable from consuming succulent vegetation, indicating copious drinking of rainwater at that time. The physiological and behavioral flexibility of desert tortoises, evident in individuals living at all three study sites, appears central to their ability to survive droughts and benefit from periods of resource abundance. The strong effects of the El Niño (ENSO) weather pattern on tortoise physiology, reproduction, and survival elucidated in this and other studies suggest that local manifestations of global climate events could have a long-term influence on the tortoise populations in the Mojave Desert.     

Effects of spatial patterns on the pollination success of a less attractive species

Plant individuals rely on pollinator services for their reproduction and often have to share these services with co‐occurring neighbours, creating complex indirect plant–plant interactions. Many current theoretical models focus on the effect of floral resources’ density on the net outcome of these indirect plant–plant interactions, often neglecting the identity of plant species in the communities and especially the species’ spatial pattern. To fill this gap, we created a spatially explicit model whose goal was to study the interplay between relative densities and spatial distribution patterns of two plant species differing in their attractiveness for pollinators. Since theory predicts that pollinator behaviour strongly governs the outcome of pollination, we allowed the pollinators to systematically change their plant preferences based on their foraging experience. Thus the interplay between density and spatial patterns of plants was tested over a continuum of behaviours from specialists to generalists. Our most striking finding was that reproductive success of the less attractive species was affected in an opposite way by spatial patterns depending on whether the species had relatively low or high densities. Namely, when the less attractive species was highly abundant, its survival was higher when aggregated in large monospecific patches than when uniformly distributed. On the other hand, when the attractive species was more abundant, the less attractive species survived better when uniformly distributed. These results were consistent as long as the scale of the plant spatial aggregation was similar to or larger than the pollinators’ detection range. Our results suggest that aggregated plant spatial patterns manipulate pollinator behaviour by trapping them within monospecific patches. This effect was sufficiently strong to enhance the survival of a competitively inferior species and hence to act in a way similar to the more familiar niche or temporal separation among plant species.     

The influence of food supply on foraging behaviour in a desert spider

We tested the alternative hypotheses that foraging effort will increase (energy maximizer model) or decrease (due to increased costs or risks) when food supply increased, using a Namib desert burrowing spider, Seothyra henscheli (Eresidae), which feeds mainly on ants. The web of S. henscheli has a simple geometrical configuration, comprising a horizontal mat on the sand surface, with a variable number of lobes lined with sticky silk. The sticky silk is renewed daily after being covered by wind-blown sand. In a field experiment, we supplemented the spiders' natural prey with one ant on each day that spiders had active webs and determined the response to an increase in prey. We compared the foraging activity and web geometry of prey-supplemented spiders to non-supplemented controls. We compared the same parameters in fooddeprived and supplemented spiders in captivity. The results support the costs of foraging hypothesis. Supplemented spiders reduced their foraging activity and web dimensions. They moulted at least once and grew rapidly, more than doubling their mass in 6 weeks. By contrast, food-deprived spiders increased foraging effort by enlarging the diameter of the capture web. We suggest that digestive constraints prevented supplemented spiders from fully utilizing the available prey. By reducing foraging activities on the surface, spiders in a prey-rich habitat can reduce the risk of predation. However, early maturation resulting from a higher growth rate provides no advantage to S. henscheli owing to the fact that the timing of mating and dispersal are fixed by climatic factors (wind and temperature). Instead, large female body size will increase fitness by increasing the investiment in young during the period of extended maternal care.     

Modeling the climatic drivers of spatial patterns in vegetation composition since the Last Glacial Maximum

Projecting the future composition and function of communities is a major challenge, and there is an urgent need to develop, improve, and test the predictive capacity of ecological models under different climate states. We tested the effect of climate on spatial patterns of plant community composition over the past 21 000 yr, focusing on whether the spatial relationships between environmental distance and compositional dissimilarity are stable over time. We used a network of fossil‐pollen sites in eastern North America, combined with paleoclimate simulations from the Last Glacial Maximum (LGM; 21 000 calibrated years before present, 21 kyr BP) to the present. We modeled relationships between climate, geography, and compositional dissimilarity at 1 kyr periods using generalized dissimilarity modeling (GDM) and determined the strongest predictors of compositional dissimilarity. We assessed the performance of models calibrated for one time period (e.g. 14 kyr BP) in predicting patterns in the same period as well as at other times (e.g. 12 kyr BP), and tested whether predictive performance was related to the magnitude of climate change between the calibration and prediction time periods. Finally, we examined whether pooling data from multiple time periods improved predictive performance. Models explained 32 to 51% of compositional dissimilarity between locations within any single time period. The best set of predictors changed across time, with summer temperature and geographic distance the strongest predictors of compositional dissimilarity for most time periods. Models built for one time period explained turnover during nearby time periods relatively well, but performance decayed across time and with increasing climate change. Results were similar regardless of whether models were projected forward or backward through time, and did not improve when data were pooled across time. GDM predicts well the spatial patterns of past compositional dissimilarity and holds promise for modeling the drivers of compositional dissimilarity across space and time. However, the modeled relationships between compositional turnover and environmental distance are non‐stationary, so caution is needed when predicting across periods of significant climatic change.     

Local population success in heterogeneous habitats: reciprocal transplant experiments completed on a desert spider

Reciprocal transplant experiments were completed to test for selection against the mixing of behavioural phenotypes in a desert spider. Most Agelenopsis aperta populations experience low prey abundances and competition for web‐sites that provide shelter from thermal extremes. These conditions favour aggressiveness towards both prey and conspecifics (an ‘arid‐land behavioural phenotype’). The spider also occupies narrow stretches of riparian habitat bordering spring‐fed streams and rivers. Here it is released from competition for prey and foraging sites, but is subject to predation by birds. A less aggressive/more fearful behaviour is selected for in these riparian habitats (a ‘riparian behavioural phenotype’). Previous work with this spider indicates that there is genetic differentiation between arid‐land and riparian populations. However, the degree to which genetic differentiation is achieved may be limited by gene flow. Reciprocal sets of enclosures were established in: (1) a dry evergreen woodland site (arid‐land phenotype) and (2) a neighbouring riparian site (riparian phenotype) in south‐eastern Arizona. Equal numbers of field collected, early instar A. aperta were introduced into native and transplant enclosures in each habitat. After 6 months of site‐imposed selection, survivorship was determined and growth estimates and behavioural trials completed on spiders remaining in the different enclosures. The same behavioural test was subsequently applied to lab‐reared offspring of the spiders surviving the respective selection regimes. Riparian transplants showed both poor survival and retarded growth in the dry woodland habitat when compared with both arid‐land and riparian natives. Arid‐land transplants that survived, however, grew equally well in riparian habitat as did dry woodland and riparian natives. Behavioural assays conducted on test subjects after selection and on their offspring reared in a controlled laboratory environment indicate that phenotypes that were inappropriate to the respective habitats were selected against in the transplant experiments. The frequency distribution of transplant spider behaviour on a continuum from fearful to aggressive was intermediate between that exhibited by respective native riparian and dry woodland spiders. It is concluded that while arid‐land and riparian behavioural ecotypes do exist, directional gene flow of arid‐land phenotypes into riparian habitat limits population subdivision.     

Effects of heavy rainfall on polychaetes: Differential spatial patterns generated by a large-scale disturbance

Heavy rainfall caused a severe disturbance on the Thalassia testudinum meadows at Morrocoy National Park in December 1999, affecting its associated infauna and inducing responses to stress in opportunistic polychaetes belonging to the Spionidae Family. The changes were evaluated at four seagrass beds, examining the structure and species composition of the Spionid macrofauna over eight samplings: four prior and four after the disturbance. Immediately after the heavy rainfall the Spionids showed transient changes, expressed as large increases of species density and richness, reaching values of 875 individuals m^− 2 (against the usual 20 individuals m^− 2) and 7 species (against the previous 1–3 species) during February 2000. These abundance pulses were represented by selected species at each location. Opportunistic species, having limited dispersion (Streblospio gynobranchiata and polydorids) were commonly found in continentally-influenced locations whereas species having dispersal mechanisms were commonly found in stations under oceanic influence (species of the Genus Prionospio). Likewise, the observed recovery patterns indicate different rhythms: while the oceanic communities showed a fast recovery, the continentally ones returned to their original states slowly. Hence, the patterns observed in the benthic polychaete communities associated with the T. testudinum beds from Morrocoy N.P. clearly indicate that the recolonization processes, succession dynamics and recovery time depend on the spatial scale of disturbance, the life history of species involved and the particular conditions of the habitat.     

Characterizing a zooplankton neighbourhood: small-scale patterns of association and abundance

• 1 This study compares small-scale (i.e. neighbourhood) associations among distributions of phytoplankton, zooplankton, and zooplankton grazing rates using a combination of correlations and analyses of spatial patchiness.
• 2 All zooplankton and phytoplankton populations had patchy distributions. On average, individuals experienced members of their same taxon at a density that was 42% greater than the mean sample density.
• 3 The various zooplankton taxa experienced different average neighbourhoods, even within the same stratum of the lake. For example, Daphnia galeata mendotae (Birge) experienced phytoplankton concentrations that were nearly 50% greater than those experienced by Daphnia pulicaria (Forbes) at the same depth.
• 4 The distributions of the various phytoplankton taxa were positively correlated with each other, but the distributions of zooplankton were negatively correlated, or more often, unrelated to each other.
• 5 Phytoplankton abundance was negatively correlated with zooplankton grazing rates.
• 6 We hypothesize that while phytoplankton distributions are driven by common external factors, the factors that determine horizontal distributions of zooplankton are species specific. In addition, zooplankton grazing appears at least partially responsible for the generation of patchiness in the distribution of phytoplankton.

     

Organic carbon and biogenic silica in marine sediments in the vicinities of the Antarctic Peninsula: spatial patterns across a climatic gradient

To assess whether sea floor sediment reflects the characteristics of the upper water column, organic carbon (OC) and biogenic silica (bSi) were measured in seventeen 5-cm-long sediment cores recovered within a climatic gradient from the northwestern Weddell Sea (WS) to the Drake Passage (DP) across the Bransfield Strait (BS). Climate settings in the study area vary from dry and cold (polar) conditions with seasonal sea ice coverage in the WS to a more humid and warm (oceanic) environment where no seasonal sea ice develops in the DP, with the BS as transitional zone undergoing seasonal sea ice coverage. OC varied between 0.2 and 1.7 % and represented more than 90 % of the total carbon, and bSi varied between 2 and 13 %. The profiles of both variables along the sediment cores suggested that the surface mixed layer is at least 5 cm thick. The inventories of the upper 5 cm of the sediment column were calculated for both variables. Regional averages were significantly lower for OC in DP samples and higher for bSi in the BS. These results suggested relatively high bSi export to the seabed in the BS, higher degradation for OC in the DP and lower bSi export from the euphotic zone in the WS. The observations made evident that the biogenic matter contents in the sediment not necessarily replicate their production characteristics at the upper ocean even across strong climatic gradients. The results may provide a useful baseline for paleo-reconstructions in a rapidly changing environment.