Native communities determine the identity of exotic invaders even at scales at which communities are unsaturated

Aim To determine why some communities are more invasible than others and how this depends on spatial scale. Our previous work in serpentine ecosystems showed that native and exotic diversity are negatively correlated at small scales, but became positively correlated at larger scales. We hypothesized that this pattern was the result of classic niche partitioning at small scales where the environment is homogeneous, and a shift to the dominance of coexistence mechanisms that depend on spatial heterogeneity in the environment at large scales. Location Serpentine ecosystem, Northern California. Methods We test the above hypotheses using the phylogenetic relatedness of natives and exotics. We hypothesized that (1) at small scales, native and exotic species should be more distantly related than expected from a random assemblage model because with biotic resistance, successful invaders should have niches that are different from those of the natives present and (2) at large scales, native and exotic species should not be more distantly related than expected. Result We find strong support for the first hypothesis providing further evidence of biotic resistance at small scales. However, at large scales, native and exotic species were also more distantly related than expected. Importantly, however, natives and exotics were more distantly related at small scales than they were at large scales, suggesting that in the transition from small to large scales, biotic resistance is relaxed but still present. Communities at large scales were not saturated in the sense that more species could enter the community, increasing species richness. However, species did not invade indiscriminately. Exotic species closely related to species already established the community were excluded. Main conclusions Native communities determine the identity of exotic invaders even at large spatial scales where communities are unsaturated. These results hold promise for predicting which species will invade a community given the species present.     

Will travel for food: spatial discounting in two new world monkeys

Nonhuman animals steeply discount the future, showing a preference for small, immediate over large, delayed rewards. Currently unclear is whether discounting functions depend on context. Here, we examine the effects of spatial context on discounting in cotton-top tamarins (Saguinus oedipus) and common marmosets (Callithrix jacchus), species known to differ in temporal discounting. We presented subjects with a choice between small, nearby rewards and large, distant rewards. Tamarins traveled farther for the large reward than marmosets, attending to the ratio of reward differences rather than their absolute values. This species difference contrasts with performance on a temporal task in which marmosets waited longer than tamarins for the large reward. These comparative data indicate that context influences choice behavior, with the strongest effect seen in marmosets who discounted more steeply over space than over time. These findings parallel details of each species' feeding ecology. Tamarins range over large distances and feed primarily on insects, which requires using quick, impulsive action. Marmosets range over shorter distances than tamarins and feed primarily on tree exudates, a clumped resource that requires patience to wait for sap to exude. These results show that discounting functions are context specific, shaped by a history of ecological pressures.     

Sampling design and the shape of species–area curves on the regional scale

Species–area relationships (SARs) are one of the fundamental patterns in ecology. However, how the way they were constructed influences resulting SAR shapes has gained astonishingly little attention. We use data of the distribution atlas of Polish butterflies to compare SARs constructed from four different designs: adding up species numbers of independent areas (species accumulation curves using contiguous and non-contiguous areas), using a nested design, and comparing species numbers of independent areas of different sizes. It appeared that the way of constructing SARs influences the outcome. We attribute this influence to the pronounced faunal dissimilarities of more distant areas (spatial species turnover). The nested design resulted in significantly higher slopes and lower intercepts of power function SARs than the other designs. SARs from all four sampling designs showed a pronounced downward curvature on small spatial scales. Only the nested design predicted species densities correctly. The implications of these results for the use of SARs in bioconservation are discussed.     

Strontium isotope analyses of large herbivore habitat use in the Cape Fynbos region of South Africa

The Cape Fynbos region of South Africa, a global biodiversity hotspot, hosted a diverse large mammal fauna till shortly after permanent European settlement (1652). How these animals survived in this exceptionally nutrient-poor environment is puzzling and it is generally believed that they restricted their movements to the more fertile shale areas. We tested the hypothesis that large herbivores avoid nutrient-poor limestone and sandstone fynbos shrublands in favour of shale-derived renosterveld vegetation using strontium (Sr) isotope analysis. If this technique could reconstruct the preferred feeding habitats of the contemporary fauna, it might also be useful for reconstructing the preferred feeding grounds of an extinct fauna. Using the assumption that small rodents have spatially restricted foraging activities, we determined the ⁸⁷Sr/⁸⁶Sr isotope ratios of rodent teeth to establish the isotopic signal characteristic of the different geological substrates in the area. We then analysed ⁸⁷Sr/⁸⁶Sr isotope ratios in the bones of a number of different large herbivores found in De Hoop Nature Reserve using laser ablation multi-collector inductively coupled plasma mass spectrometry. These values were compared to the bioavailable (rodent) values on the respective geological substrates. The technique identified differences in feeding substrate selection between different species and groups of the same species. The results also showed that shale renosterveld shrubland is not the exclusive source of nutrition for the large herbivores. Strikingly different isotope ratios among individuals in some populations pointed to significant dispersal events from distant sources. However, we were unable to pinpoint the exact feeding areas using Sr isotope analysis probably because some animals use a combination of substrates for feeding and because the geology of the study area is complex with graded isotope signals. We suggest that this technique is a valuable additional tool for exploring large mammal foraging behaviour on habitats associated with contrasting and less complex geology.     

Changes in plant species density in an experimentally fragmented forest landscape: Are the effects scale‐dependent?

The establishment of commercial tree plantations is a common cause of habitat fragmentation globally, yet the influence of this land use on plant species density in fragmented native forests requires further understanding. Theory predicts species density will be highest in large areas of habitat, and should decline as area is reduced, but whether these effects are scale‐dependent is largely unknown. We assessed plant species density (total, common and uncommon species) in experimentally fragmented eucalypt forest remnants (0.25, 0.88, 3.06 ha) surrounded by a pine plantation, at three spatial scales using nested quadrats. Specifically, we consider how plant species density varies across three different sized fragments, and whether the response of species density is magnified in common or uncommon species. Species density in small fragments was higher than continuous forest for all species groupings at the smallest spatial scale (1 m²), and for total and common species at the next smallest spatial scale (16 m²). No species groupings responded to reductions in habitat area at the largest spatial scale (144 m²). We suggest that pine plantations may cause higher species density in small fragments via two mechanisms, either by allowing species to infill unoccupied areas within small fragments, or by buffering small fragments from species losses during a severe and prolonged drought. In both cases we suspect reduced moisture stress (e.g. increased soil moisture, higher shading and reduced temperature) in small fragments has led to the observed changes in species density.     

Assessing the ecological impacts of an introduced seastar: the importance of multiple methods

Introduced species are having major impacts in terrestrial, freshwater, and marine ecosystems worldwide. Given that resources for management are limited and that only a small percentage of invaders are likely to cause large ecological change, management priorities should be based on the severity of immediate and anticipated impacts on native assemblages and commercial species. This paper synthesizes work on the current and predicted impacts of an introduced predatory seastar (Asterias amurensis) on soft sediment assemblages, including native species subject to commercial fishing, in the Derwent Estuary and other areas of southeast Tasmania. Due to the absence of baseline data prior to the arrival of the seastar and the presence of other anthropogenic stressors in the estuary, estimating the impact of the seastar is difficult. To help overcome the weaknesses of any single method, our assessment of impact rests on 'weight of evidence' from multiple approaches. Results from experimental manipulations at small scales, detailed observations of feeding, and field surveys over a range of spatial scales in areas with and without the seastar provide strong evidence that predation by the seastar is likely to be responsible for the decline and subsequent rarity of bivalve species that live just below or on the sediment surface in the Derwent Estuary. The data suggest that should seastar densities in other areas on the Tasmanian coast attain the current levels in the Derwent Estuary, there are likely to be large direct effects on native assemblages, particularly on populations of large surface dwelling bivalves, including several commercial species. Given the seastar's ability to exploit a broad range of food resources other than bivalves, and the functional importance of bivalves in native systems, we predict broader direct and indirect effects on native assemblages. We would be unable to reach these same conclusions from a single approach to assessing impacts. The overall picture from the combination of methods at different scales provides more information than the sum of the results of the separate lines of investigation.     

Investigating microbial associations from sequencing survey data with co‐correspondence analysis

Microbial communities, which drive major ecosystem functions, consist of a wide range of interacting species. Understanding how microbial communities are structured and the processes underlying this is crucial to interpreting ecosystem responses to global change but is challenging as microbial interactions cannot usually be directly observed. Multiple efforts are currently focused to combine next‐generation sequencing (NGS) techniques with refined statistical analysis (e.g., network analysis, multivariate analysis) to characterize the structures of microbial communities. However, most of these approaches consider a single table of sequencing data measured for several samples. Technological advances now make it possible to collect NGS data on different taxonomic groups simultaneously for the same samples, allowing us to analyse a pair of tables. Here, an analytical framework based on co‐correspondence analysis (CoCA) is proposed to study the distributions, assemblages and interactions between two microbial communities. We show the ability of this approach to highlight the relationships between two microbial communities, using two data sets exhibiting various types of interactions. CoCA identified strong association patterns between autotrophic and heterotrophic microbial eukaryote assemblages, on the one hand, and between microalgae and viruses, on the other. We demonstrate also how CoCA can be used, complementary to network analysis, to reorder co‐occurrence networks and thus investigate the presence of patterns in ecological networks.     

The influence of predation on horizontal distribution of zooplankton species

SUMMARY. 1. The horizontal distributions of Daphnia longispina and Bosmina tongispina in Lake Kvernavatn (Norway) were investigated twice in 1982. In late spring, when populations were small, the two species inhabited the same areas, and they were evenly distributed from the littoral to the pelagic. At high population densities, during midsummer, the species were spatially segregated, D. longispina being pelagic and B. longispina littoral in distribution.
2. The distribution and feeding of three-spined sticklebacks (Gasteros-teus aculeatus) were also studied. The sticklebacks were apparently forced into littoral areas by larger piscivorous predators in the pelagic and they were consequently restricted to foraging primarily on B. longispina, which formed dense swarms during daytime in summer.
3. We suggest that predation and competition influence the spatial distribution of zooplankton species. The feeding efficiency of fish foraging on high-density zooplankton populations can be reduced by spatial segregation of zooplankton species. Where high local densities occur, due to swarm formation, predation is changed from size-selective feeding to consumption of spatially isolated individuals.     

Large‐scale patterns of seed removal by small mammals differ between areas of low‐ versus high‐wolf occupancy

Because most tree species recruit from seeds, seed predation by small‐mammal granivores may be important for determining plant distribution and regeneration in forests. Despite the importance of seed predation, large‐scale patterns of small‐mammal granivory are often highly variable and thus difficult to predict. We hypothesize distributions of apex predators can create large‐scale variation in the distribution and abundance of mesopredators that consume small mammals, creating predictable areas of high and low granivory. For example, because gray wolf (Canis lupus) territories are characterized by relatively less use by coyotes (C. latrans) and greater use by foxes (Vulpes vulpes, Urocyon cinereoargentus) that consume a greater proportion of small mammals, wolf territories may be areas of reduced small‐mammal granivory. Using large‐scale, multiyear field trials at 22 sites with high‐ and low‐wolf occupancy in northern Wisconsin, we evaluated whether removal of seeds of four tree species was lower in wolf territories. Consistent with the hypothesized consequences of wolf occupancy, seed removal of three species was more than 25% lower in high‐wolf‐occupancy areas across 2 years and small‐mammal abundance was more than 40% lower in high‐wolf areas during one of two study years. These significant results, in conjunction with evidence of seed consumption in situ and the absence of significant habitat differences between high‐ and low‐wolf areas, suggest that top‐down effects of wolves on small‐mammal granivory and seed survival may occur. Understanding how interactions among carnivores create spatial patterns in interactions among lower trophic levels may allow for more accurate predictions of large‐scale patterns in seed survival and forest composition.     

Ontogeny of feeding in two native and one alien fish species from the Murray-Darling Basin, Australia

Investigations into the feeding of the early stages of fishes can provide insights into processes influencing recruitment. In this study, we examined ontogenetic changes in morphology and feeding behaviour of two native Australian freshwater species, Murray cod, Maccullochella peelii peelii, and golden perch, Macquaria ambigua, and the alien species, common carp, Cyprinus carpio. Murray cod free embryos are large and well developed at the onset of feeding, whereas the other two species begin exogenous feeding much younger and are smaller and less-developed. Carp commence exogenous feeding 3 days earlier than golden perch, and show more advanced development of the eyes and ingestive apparatus. We conducted feeding experiments, presenting larvae of the three species with a standardised prey mix (comprising equal numbers of small calanoid copepods, large calanoid copepods, small Daphnia, and large Daphnia). Larvae of most tested ages and species showed a preference for mid-sized prey (300–500 μm wide). This was true even when their gapes substantially exceeded the largest prey offered. Daphnia were consumed more than similar-sized copepods. The results of this study suggest that survival through their larval period will be threatened in all three species if catchable prey <500 μm in width are not available throughout such time. They also suggest that interspecific competition for prey may occur, especially when larvae are very young. The precocious development of structures involved in feeding and the extended transition from endogenous to exogenous feeding of early carp larvae are likely to have contributed to the success of this species since its introduction to Australia.     

Spatial Scaling of Non-Native Fish Richness across the United States

A major goal and challenge of invasion ecology is to describe and interpret spatial and temporal patterns of species invasions. Here, we examined fish invasion patterns at four spatially structured and hierarchically nested scales across the contiguous United States (i.e., from large to small: region, basin, watershed, and sub-watershed). All spatial relationships in both richness and fraction between species groups (e.g., natives vs. exotics) were positive at large scales. However, contrary to predictions using null/neutral models, the patterns at small scales were hump-shaped (unimodal), not simply negative. The fractions of both domestic (introduced among watersheds within the USA) and foreign (introduced from abroad) exotics increased with area across scales but decreased within each scale. The foreign exotics exhibited the highest dominance (lowest evenness) and spatial variation in distribution, followed by domestic exotics and natives, although on average natives still occupy larger areas than domestic and foreign exotics. The results provide new insight into patterns and mechanisms of fish species invasions at multiple spatial scales in the United States.     

红松存在/不存在数据的多尺度空间分布格局

红松属小兴安岭地区地带性植被优势种,该地区也是其分布的北缘。在景观尺度上开展红松的分布格局研究有利于进一步了解红松分布机理、未来迁移过程等问题,对其经营和保护有重要意义。将景观指数法与点格局分析法结合,设定8个空间尺度,利用红松存在/不存在数据,通过计算各空间尺度上红松聚集程度和景观指数,分析小兴安岭地区红松种群在多尺度上的分布格局。研究结果表明,小尺度上红松聚集分布明显,随机分布区多处于其聚集分布区的边缘,均匀分布区则散布在其聚集分布区内。景观指数研究表明,通过景观指数可判断红松聚集分布格局趋势,而不能判断均匀分布、随机分布格局趋势,因为它们在多尺度下景观指数波动大,不能用景观指数来描述分布格局。研究得出如下结论:1)红松主要分布在其分布区的核心区域内,在分布区边缘和过渡带上呈随机分布,2)存在/不存在数据能够用来分析种群的多尺度空间分布格局,3)空间尺度的变化会引起树种分布格局的变化,随机分布随尺度增加,边缘化程度加强,4)单一尺度上,景观格局指数不能完全描述种群分布格局;而在多尺度上,变化趋势稳定的景观指数表明聚集分布存在,而波动剧烈的景观指数常与随机分布和均匀分布联系在一起,5)地形因子中,红松对坡度和海拔两个因子变化敏感。     

Evolution of termite functional diversity: analysis and synthesis of local ecological and regional influences on local species richness

Aim To (1) describe termite functional diversity patterns across five tropical regions using local species richness sampling of standardized areas of habitat; (2) assess the relative importance of environmental factors operating at different spatial and temporal scales in influencing variation in species representation within feeding groups and functional taxonomic groups across the tropics; (3) achieve a synthesis to explain the observed patterns of convergence and divergence in termite functional diversity that draws on termite ecological and biogeographical evidence to‐date, as well as the latest evidence for the evolutionary and distributional history of tropical rain forests. Location Pantropical. Methods A pantropical termite species richness data set was obtained through sampling of eighty‐seven standardized local termite diversity transects from twenty‐nine locations across five tropical regions. Local‐scale, intermediate‐scale and large‐scale environmental data were collected for each transect. Standardized termite assemblage and environmental data were analysed at the levels of whole assemblages and feeding groups (using components of variance analysis) and at the level of functional taxonomic groups (using correspondence analysis and canonical correspondence analysis). Results Overall species richness of local assemblages showed a greater component of variation attributable to local habitat disturbance level than to region. However, an analysis accounting for species richness across termite feeding groups indicated a much larger component of variation attributable to region. Mean local assemblage body size also showed the greater overall significance of region compared with habitat type in influencing variation. Ordination of functional taxonomic group data revealed a primary gradient of variation corresponding to rank order of species richness within sites and to mean local species richness within regions. The latter was in the order: Africa > south America > south‐east Asia > Madagascar > Australia. This primary gradient of species richness decrease can be explained by a decrease in species richness of less dispersive functional taxonomic groups feeding on more humified food substrates such as soil. Hence, the transects from more depauperate sites/regions were dominated by more dispersive functional taxonomic groups feeding on less humified food substrates such as dead wood. Direct gradient analysis indicated that ‘region’ and other large‐scale factors were the most important in explaining patterns of local termite functional diversity followed by intermediate‐scale geographical and site variables and, finally, local‐scale ecological variables. Synthesis and main conclusions Within regions, centres of termite functional diversity lie in lowland equatorial closed canopy tropical forests. Soil feeding termite evolution further down food substrate humification gradients is therefore more likely to have depended on the long‐term presence of this habitat. Known ecological and energetic constraints upon contemporary soil feeders lend support for this hypothesis. We propose further that the anomalous distribution of termite soil feeder species richness is partly explained by their generally very poor dispersal abilities across oceans. Evolution, radiation and dispersal of soil feeder diversity appears to have been largely restricted to what are now the African and south American regions. The inter‐regional differences in contemporary local patterns of termite species richness revealed by the global data set point to the possibility of large differences in consequent ecosystem processes in apparently similar habitats on different continents.     

Foraging and territory utilization of blackbirds (Turdus merula) and song thrushes (Turdus philomelos)

Territory utilization of blackbirds and song thrushes is examined in relation to spatial and temporal changes in the distribution of prey. Pairs within a territory showed similar preferences for particular areas. There was considerable diurnal variation in foraging localities and overall the two species tended to concentrate their search effort in areas of high prey density. Rates of feeding attempts and feeding techniques also varied with time of day. Changes were similar in the two species. At high rates of feeding during the middle of the day the foraging range was more restricted than at other times. Over short time periods individuals searched non-randomly by avoiding recently exploited areas.     

Comparative organ differentiation during early life stages of marine fish

The basic developmental mechanisms of teleosts are similar, but there are differences with respect to the timing of developmental events. These events are controlled by genetic and environmental factors. Direct comparisons of organogenesis are complicated due to large variations in egg sizes and incubation temperatures between species. But in general, cultivated small marine pelagic fish larvae originating from rather small eggs (like gadoids, flatfishes, sparids) hatch with a relatively large yolk sac, a larval finfold and subdermal space and under-developed organs. Developmental status at hatch differs between species and the duration of the yolk sac period varies. Main organs and organ systems become functional by first feeding and differentiate during the larval stage and metamorphosis. Species developing directly via large yolk-rich eggs and a long incubation period have a juvenile like morphology and organ functionality at first feeding, sometimes immediately after hatch (like wolffishes). Histomorphological and cell- or organ functional studies of developing embryos and larvae of cultivated species constitute basic information for understanding species-specific events, of utmost importance for improving production protocols. Information is still lacking on early functionality of endocrine and immunocompetent tissues and organs, areas that deserve future focus.     

Untangling the relationships among regional occupancy,species traits,and niche characteristics in stream invertebrates

The regional occupancy and local abundance of species are affected by various species traits, but their relative effects are poorly understood. We studied the relationships between species traits and occupancy (i.e., proportion of sites occupied) or abundance (i.e., mean local abundance at occupied sites) of stream invertebrates using small‐grained data (i.e., local stream sites) across a large spatial extent (i.e., three drainage basins). We found a significant, yet rather weak, linear relationship between occupancy and abundance. However, occupancy was strongly related to niche position (NP), but it showed a weaker relationship with niche breadth (NB). Abundance was at best weakly related to these explanatory niche‐based variables. Biological traits, including feeding modes, habit traits, dispersal modes and body size classes, were generally less important in accounting for variation in occupancy and abundance. Our findings showed that the regional occupancy of stream invertebrate species is mostly related to niche characteristics, in particular, NP. However, the effects of NB on occupancy were affected by the measure itself. We conclude that niche characteristics determine the regional occupancy of species at relatively large spatial extents, suggesting that species distributions are determined by environmental variation among sites.     

Quantitative analysis of bottlenose dolphin movement patterns and their relationship with foraging

1. Broad-scale telemetry studies have greatly improved our understanding of the ranging patterns and habitat-use of many large vertebrates. However, there often remains considerable uncertainty over the function of different areas or the factors influencing habitat selection. Further insights into these processes can be obtained through analyses of finer scale movement patterns. For example, search behaviour may be modified in response to prey distribution and abundance. 2. In this study, quantitative analysis techniques are applied to the movements of bottlenose dolphins, recorded from land using a theodolite, to increase our understanding of their foraging strategies. Movements were modelled as a correlated random walk (CRW) and a biased random walk (BRW) to identify movement types and using a first-passage time (FPT) approach, which quantifies the time allocated to different areas and identifies the location and spatial scale of intensive search effort. 3. Only a quarter of the tracks were classed as CRW movement. Turning angle and directionality appeared to be key factors in determining the type of movement adopted. A high degree of overlap in search effort between separate movement paths indicated that there were small key sites (0.3 km radius) within the study area (4 km(2)). Foraging behaviour occurred mainly within these intensive search areas, indicating that they were feeding sites. 4. This approach provides a quantitative method of identifying important foraging areas and their spatial scale. Such techniques could be applied to movement paths for a variety of species derived from telemetry studies and increase our understanding of their foraging strategies.     

Phenotypic integration in feliform carnivores: Covariation patterns and disparity in hypercarnivores versus generalists

The skeleton is a complex arrangement of anatomical structures that covary to various degrees depending on both intrinsic and extrinsic factors. Among the Feliformia, many species are characterized by predator lifestyles providing a unique opportunity to investigate the impact of highly specialized hypercarnivorous diet on phenotypic integration and shape diversity. To do so, we compared the shape of the skull, mandible, humerus, and femur of species in relation to their feeding strategies (hypercarnivorous vs. generalist species) and prey preference (predators of small vs. large prey) using three-dimensional geometric morphometric techniques. Our results highlight different degrees of morphological integration in the Feliformia depending on the functional implication of the anatomical structure, with an overall higher covariation of structures in hypercarnivorous species. The skull and the forelimb are not integrated in generalist species, whereas they are integrated in hypercarnivores. These results can potentially be explained by the different feeding strategies of these species. Contrary to our expectations, hypercarnivores display a higher disparity for the skull than generalist species. This is probably due to the fact that a specialization toward high-meat diet could be achieved through various phenotypes. Finally, humeri and femora display shape variations depending on relative prey size preference. Large species feeding on large prey tend to have robust long bones due to higher biomechanical constraints.     

From ratites to rats: the size of fleshy fruits shapes species' distributions and continental rainforest assembly

Seed dispersal is a key process in plant spatial dynamics. However, consistently applicable generalizations about dispersal across scales are mostly absent because of the constraints on measuring propagule dispersal distances for many species. Here, we focus on fleshy-fruited taxa, specifically taxa with large fleshy fruits and their dispersers across an entire continental rainforest biome. We compare species-level results of whole-chloroplast DNA analyses in sister taxa with large and small fruits, to regional plot-based samples (310 plots), and whole-continent patterns for the distribution of woody species with either large (more than 30 mm) or smaller fleshy fruits (1093 taxa). The pairwise genomic comparison found higher genetic distances between populations and between regions in the large-fruited species (Endiandra globosa), but higher overall diversity within the small-fruited species (Endiandra discolor). Floristic comparisons among plots confirmed lower numbers of large-fruited species in areas where more extreme rainforest contraction occurred, and re-colonization by small-fruited species readily dispersed by the available fauna. Species'' distribution patterns showed that larger-fruited species had smaller geographical ranges than smaller-fruited species and locations with stable refugia (and high endemism) aligned with concentrations of large fleshy-fruited taxa, making them a potentially valuable conservation-planning indicator.