Conspecific and Heterospecific Plant Densities at Small-Scale Can Drive Plant-Pollinator Interactions

Generalist pollinators are important in many habitats, but little research has been done on small-scale spatial variation in interactions between them and the plants that they visit. Here, using a spatially explicit approach, we examined whether multiple species of flowering plants occurring within a single meadow showed spatial structure in their generalist pollinator assemblages.We report the results for eight plant species for which at least 200 individual visits were recorded. We found that for all of these species, the proportions of their general pollinator assemblages accounted for by particular functional groups showed spatial heterogeneity at the scale of tens of metres. This heterogeneity was connected either with no or only subtle changes of vegetation and flowering species composition. In five of these species, differences in conspecific plant density influenced the pollinator communities (with greater dominance of main pollinators at low-conspecific plant densities). The density of heterospecific plant individuals influenced the pollinator spectrum in one case.Our results indicate that the picture of plant-pollinator interactions provided by averaging data within large plots may be misleading and that within-site spatial heterogeneity should be accounted for in terms of sampling effort allocation and analysis. Moreover, spatially structured plant-pollinator interactions may have important ecological and evolutionary consequences, especially for plant population biology.     

Abundance-range size relationships of breeding and wintering birds in Britain: a comparative analysis

Both breeding and wintering assemblages of birds in Britain exhibit positive interspecific relationships between population size and geographic range size, such that the average density of species is greater if they are more widely distributed Species in common to both assemblages, that is resident species, had greater population sizes, geographic range sizes, and densities in winter In contrast, whilst winter migrants had higher abundances than summer migrants, the range sizes of the former were disproportionately larger still, resulting in a lower density for species that only winter in Britain than for those that only breed Such differences aside, the overall form of the abundance-range size relationship is remarkably similar between the two assemblages and their constituent subsets of species     

Field tests of interspecific competition in ant assemblages: revisiting the dominant red wood ants

1. There has been considerable debate on the importance of competition in ecological communities, but its importance in structuring ant assemblages has often been uncritically accepted. Here, we briefly review field experiments examining competition in ant assemblages and use a removal experiment to test the effect of the classical territorial dominant ant, Formica aquilonia. Ants of this species group are thought to structure communities through a dominance hierarchy. 2. First, we used pitfall traps to compare the abundance of other ants in replicated sites with low and high densities of F. aquilonia. We found differences in community composition, in particular, Camponotus herculeanus was more common in low-density sites, in accordance with predictions. Differences in ant assemblages were not owing to differences in measured habitat variables. 3. We removed F. aquilonia from a set of high-density sites, using physical and chemical methods, and repeated these procedures at procedural control sites. One year after removal, abundances of F. aquilonia at removal sites were similar to those at low-density sites. However, the composition of other species did not change in response to F. aquilonia removal. Replication rates were identical in the mensurative and experimental components of this study, so this is unlikely to be owing to the analysis being insufficiently powerful. 4. We suggest three possibilities for the lack of difference. First, the study may have been too short term or small scale to detect differences. However, previous studies have shown effects on smaller spatial- and temporal-scales. Second, priority effects may be important in the successful colonisation by F. aquilonia. Thirdly, boreal ant assemblages may be too depauperate for redundancy in ecological roles and for competition to play an important structuring role. 5. We thus recommend that long-term large-scale experiments be considered essential if we are to distinguish between competing hypotheses in community ecology.     

Biogeography and the structure of coral reef fish communities on isolated islands

Aim To determine the applicability of biogeographical and ecological theory to marine species at two remote island locations. This study examines how biogeography, isolation and species geographic range size influence patterns of species richness, endemism, species composition and the abundance of coral reef fishes. Location Christmas Island and the Cocos (Keeling) Islands in the tropical eastern Indian Ocean. Methods Published species lists and underwater visual surveys were used to determine species richness, endemism, species composition and abundance of reef fishes at the islands. These data were statistically compared with patterns of species composition and abundance from the neighbouring ‘mainland’ Indonesian region. Results The two isolated reef fish communities were species‐poor and contained a distinct taxonomic composition with an overrepresentation of species with high dispersal potential. Despite low species richness, we found no evidence of density compensation, with population densities on the islands similar to those of species‐rich mainland assemblages. The mix of Indian and Pacific Ocean species and the proportional representations of the various regional faunas in the assemblages were not influenced by the relative proximity of the islands to different biogeographical provinces. Moreover, species at the edge of their range did not have a lower abundance than species at the centre of their range, and endemic species had substantially higher abundances than widespread species. At both locations, endemism was low (less than 1.2% of the community); this may be because the locations are not sufficiently isolated or old enough to promote the evolution of endemic species. Main conclusions The patterns observed generally conform to terrestrial biogeographical theory, suggesting that similar processes may be influencing species richness and community composition in reef fish communities at these remote islands. However, species abundances differed from typical terrestrial patterns, and this may be because of the life history of reef fishes and the processes maintaining isolated populations.     

The ecological characteristics of idiosyncratic and nested diatoms

Nestedness represents the degree to which species assemblages are proper subsets of larger assemblages. Highly nested assemblages are characterized by a high number of species conforming to nested subset pattern and a low number of species which depart from nested pattern, that is, idiosyncratic species. The main aims of this paper were to (i) examine whether local stream habitat factors and spatial variables govern the number of nested stream diatom species at sites, (ii) to examine whether nested and idiosyncratic species differ in ecological characteristics, and (iii) to study if these distinctions have implications for turnover in stream diatom communities. Stream diatom communities showed a highly significant nested pattern, with observed matrix temperature of 28.4 degrees C. However, number of idiosyncratic species was high, constituting 44% of total species number of the survey. The number of nested species at sites was not related to any of the measured habitat factors or geographical location of the sampling sites. Idiosyncratic species were significantly (ANOVA: P=0.002) more widely distributed than nested species. Partial mantel tests showed that idiosyncratic species exhibited faster turnover along geographical and environmental distance than nested species. These data showed that although the degree of nestedness was highly significant, stream diatom communities were nevertheless characterized by a number of idiosyncratic species departing from the nested subset pattern. It seems that the compositional turnover is faster in communities that are dominated by idiosyncratic species suggesting that turnover diversity in diatoms may be governed more by the distribution of idiosyncratic species.     

Nestedness, anti-nestedness, and the relationship between prevalence and intensity in ectoparasite assemblages of marine fish: a spatial model of species coexistence

Nested species subset patterns consist in a hierarchical structure of species composition in related assemblages, with the species found in depauperate assemblages representing non-random subsets of progressively richer ones. This pattern has been found at the infracommunity level in about a third of the fish ectoparasite assemblages studied to date. Here we present evidence for another non-random structural pattern in assemblages of fish ectoparasites, anti-nestedness, which corresponds to situations in which parasite species are always absent from infracommunities richer than the most depauperate one in which they occur. We show that this pattern is exactly as common as nestedness, and that anti-nested assemblages are characterised by significantly lower prevalence and mean intensities of parasites than nested assemblages. In addition, we found a positive relationship between the prevalence and the mean intensity of parasites across the different assemblages. We propose a link between the nestedness/anti-nestedness continuum and the prevalence-intensity relationship that may involve colonisation-extinction processes. The results presented here suggest that, although nestedness may not be common in parasite communities, other departures from random species assembly are possible, and that some form of structure may be present in many communities. The continuum between nestedness and anti-nestedness also has implications for recent models of species coexistence in communities.     

Toward ecologically explicit null models of nestedness

A community is "nested" when species assemblages in less rich sites form nonrandom subsets of those at richer sites. Conventional null models used to test for statistically nonrandom nestedness are under- or over-restrictive because they do not sufficiently isolate ecological processes of interest, which hinders ecological inference. We propose a class of null models that are ecologically explicit and interpretable. Expected values of species richness and incidence, rather than observed values, are used to create random presence-absence matrices for hypothesis testing. In our examples, based on six datasets, expected values were derived either by using an individually based random placement model or by fitting empirical models to richness data as a function of environmental covariates. We describe an algorithm for constructing unbiased null matrices, which permitted valid testing of our null models. Our approach avoids the problem of building too much structure into the null model, and enabled us to explicitly test whether observed communities were more nested than would be expected for a system structured solely by species-abundance and species-area or similar relationships. We argue that this test or similar tests are better determinants of whether a system is truly nested; a nested system should contain unique pattern not already predicted by more fundamental ecological principles such as species-area relationships. Most species assemblages we studied were not nested under these null models. Our results suggest that nestedness, beyond that which is explained by passive sampling processes, may not be as widespread as currently believed. These findings may help to improve the utility of nestedness as an ecological concept and conservation tool.     

The Colonisation of Exotic Species Does Not Have to Trigger Faunal Homogenisation: Lessons from the Assembly Patterns of Arthropods on Oceanic Islands

Human-caused disturbances can lead to the extinction of indigenous (endemic and native) species, while facilitating and increasing the colonisation of exotic species; this increase can, in turn, promote the similarity of species compositions between sites if human-disturbed sites are consistently invaded by a regionally species-poor pool of exotic species. In this study, we analysed the extent to which epigean arthropod assemblages of four islands of the Azorean archipelago are characterised by nestedness according to a habitat-altered gradient. The degree of nestedness represents the extent to which less ubiquitous species occur in subsets of sites occupied by the more widespread species, resulting in an ordered loss/gain of species across environmental or ecological gradients. A predictable loss of species across communities while maintaining others may lead to more similar communities (i.e. lower beta-diversity). In contrast, anti-nestedness occurs when different species tend to occupy distinct sites, thus characterising a replacement of species across such gradients. Our results showed that an increase in exotic species does not promote assemblage homogenisation at the habitat level. On the contrary, exotic species were revealed as habitat specialists that constitute new and well-differentiated assemblages, even increasing the species compositional heterogeneity within human-altered landscapes. Therefore, contrary to expectations, our results show that both indigenous and exotic species established idiosyncratic assemblages within habitats and islands. We suggest that both the historical extinction of indigenous species in disturbed habitats and the habitat-specialised character of some exotic invasions have contributed to the construction of current assemblages.     

Structure of communities of ground-dwelling animals at the junction of two phytogeographic zones

Changes in the structure of tenebrionid beetle, lizard and rodent communities along the aridity gradient that determines the position of a phytogeographic province boundary were studied to test: (1) whether there are two different species assemblages on either side of the boundary (2) the extent of their differentiation and (3) if they are composed of different or the same faunal elements. Several ordination methods were tested for their capacity to reflect community structure, and discriminant function analysis was selected as the primary method of ordination. Each of the ordination axes reflected a complex environmental gradient. The latter was different for each taxon. The gradient of productivity cross-cuts ecological space for each animal group in an individual manner. Revealed spatial assemblages of species were related to the level of productivity. There was one assemblage of species at high productivity and one or two assemblages occurred at low productivity in each taxon. The α–diversity curve of rodents changed weakly along the productivity gradient, whereas those of lizard and beetle communities were distinctly unimodal and had maxima near the middle of the gradient. β–diversity curves of rodents had a concave shape, but those of lizard and beetle communities increased from low productivity values to the middle of the gradient and after that weakly declined to the upper (productive) end of the gradient. Seven types of ranges or areographic groups of species were distinguished. Each species assemblage was composed of different faunal elements. Comparison of the results of ecological and areographic analyses demonstrated that two assemblages of rodents and lizards represent different guilds within the same community rather than different communities. The phytogeographic border for these taxa is an ecological rather than a geographical boundary. The difference between the two tenebrionid assemblages appears to be zoogeographical rather than ecological, and coincides with the phytogeographic boundary.     

Ghosts of yellowstone: multi-decadal histories of wildlife populations captured by bones on a modern landscape

Natural accumulations of skeletal material (death assemblages) have the potential to provide historical data on species diversity and population structure for regions lacking decades of wildlife monitoring, thereby contributing valuable baseline data for conservation and management strategies. Previous studies of the ecological and temporal resolutions of death assemblages from terrestrial large-mammal communities, however, have largely focused on broad patterns of community composition in tropical settings. Here, I expand the environmental sampling of large-mammal death assemblages into a temperate biome and explore more demanding assessments of ecological fidelity by testing their capacity to record past population fluctuations of individual species in the well-studied ungulate community of Yellowstone National Park (Yellowstone). Despite dramatic ecological changes following the 1988 wildfires and 1995 wolf re-introduction, the Yellowstone death assemblage is highly faithful to the living community in species richness and community structure. These results agree with studies of tropical death assemblages and establish the broad capability of vertebrate remains to provide high-quality ecological data from disparate ecosystems and biomes. Importantly, the Yellowstone death assemblage also correctly identifies species that changed significantly in abundance over the last 20 to ～80 years and the directions of those shifts (including local invasions and extinctions). The relative frequency of fresh versus weathered bones for individual species is also consistent with documented trends in living population sizes. Radiocarbon dating verifies the historical source of bones from Equus caballus (horse): a functionally extinct species. Bone surveys are a broadly valuable tool for obtaining population trends and baseline shifts over decadal-to-centennial timescales.     

Massive structural redundancies in species composition patterns of floodplain forest moths

Terrestrial arthropod communities usually consist of very large species numbers. Data from experiments or long time‐series would be required to ascertain the functional significance of individual species. Both are largely unavailable for species‐rich natural communities. Recognising structural redundancies in species composition allows for an alternative approach to address how strong functional redundancy might be in natural assemblages, if structural and functional redundancies are related to each other. Determining structural redundancies is a regular topic in aquatic ecology, but has rarely been applied to terrestrial communities. We explored the extent of structural redundancy in species‐rich terrestrial insect assemblages and whether structural redundancies are contingent to species abundances or functional group affiliations. We used the BVSTEP algorithm to determine structural redundancies in a large data set of moth species (32 181 individuals; 448 species) that had been sampled with light‐traps in three different floodplain forests in eastern Austria. We partitioned the moth species into 12 functional types based on larval host‐plant affiliations to test if moth species included in reduced subsets represent functional groups in the same proportions as they occur in the entire fauna. We observed far more massive structural redundancies in moth assemblages than previously found in aquatic communities. Subsets containing only 8–15 species (1.8–3.3% of all recorded species) were still highly representative for the overall data. Subsets selected by the BVSTEP procedure performed better than equally small subsets that were defined solely by species abundances or by functional group affiliations. Effective ‘surrogate’ subsets contained only 6–9 of the 12 functional moth types. High abundance only loosely corresponded with the frequency at which a moth species was included in the subsets. Thus, certain uncommon species contribute importantly to species composition patterns. Our results show unexpectedly extensive structural redundancies in complex floodplain forest moth communities, which may also indicate strong functional redundancies.     

Linking community assembly and structure across scales in a wild mouse parasite community

Understanding what processes drive community structure is fundamental to ecology. Many wild animals are simultaneously infected by multiple parasite species, so host–parasite communities can be valuable tools for investigating connections between community structures at multiple scales, as each host can be considered a replicate parasite community. Like free‐living communities, within‐host–parasite communities are hierarchical; ecological interactions between hosts and parasites can occur at multiple scales (e.g., host community, host population, parasite community within the host), therefore, both extrinsic and intrinsic processes can determine parasite community structure. We combine analyses of community structure and assembly at both the host population and individual scales using extensive datasets on wild wood mice (Apodemus sylvaticus) and their parasite community. An analysis of parasite community nestedness at the host population scale provided predictions about the order of infection at the individual scale, which were then tested using parasite community assembly data from individual hosts from the same populations. Nestedness analyses revealed parasite communities were significantly more structured than random. However, observed nestedness did not differ from null models in which parasite species abundance was kept constant. We did not find consistency between observed community structure at the host population scale and within‐host order of infection. Multi‐state Markov models of parasite community assembly showed that a host's likelihood of infection with one parasite did not consistently follow previous infection by a different parasite species, suggesting there is not a deterministic order of infection among the species we investigated in wild wood mice. Our results demonstrate that patterns at one scale (i.e., host population) do not reliably predict processes at another scale (i.e., individual host), and that neutral or stochastic processes may be driving the patterns of nestedness observed in these communities. We suggest that experimental approaches that manipulate parasite communities are needed to better link processes at multiple ecological scales.     

Endozoochory varies with ecological scale and context

Several studies on endozoochory have established large herbivores as important for seed dispersal, yet no studies have evaluated how endozoochory is dependent on ecological scale and context. Here we address effects of reindeer density on endozoochory in a hierarchical, multi-scale study, encompassing several ecological contexts.
We found reindeer density effects on endozoochory to vary with spatial scale. Higher reindeer densities at the level of landscape areas, as indexed through faeces abundance, were related to both less species and lower abundance of emerging plants from faeces. In contrast, there was no effect of higher reindeer densities at the level of herding districts (i.e. large scale assemblages of landscapes). Lack of consistency between scales reflects ecological hierarchy, indicating that reindeer density effects on endozoochory best matches at the scale of landscapes.
Pasture seed plant composition was only partly an important ecological factor. That is, ericoid species, the dominating plants in the pastures, were also the most abundant seed plants found to emerge from the faeces. However, most herbaceous seed plant species in the pastures were not emerging from the faeces and the few that emerged were positively related to the site fertility and altitude of the pasture.
Studies addressing endozoochory of ruminants are typically concerned with seed plants, whereas in this study we also found indications of that diaspores of ferns are viable after passing the digestive tract of large herbivores. Vascular spore plants were even more abundant in the faeces than were vascular seed plants.
Results from this study demonstrate that reindeer may counteract a potentially negative impact on seed limitation from their grazing by returning viable seeds in their faeces. However, in Finnmark, northern Norway, this effect is only marginal, relates only to a very few species and individuals and shows ecological scale and context dependence.     

Niche conservatism constrains Australian honeyeater assemblages in stressful environments

The hypothesis of phylogenetic niche conservatism proposes that most extant members of a clade remain in ancestral environments because expansion into new ecological space imposes a selectional load on a population. A prediction that follows is that local assemblages contain increasingly phylogenetically clustered subsets of species with increasing difference from the ancestral environment of a clade. We test this in Australian Meliphagidae, a continental radiation of birds that originated in wet, subtropical environments, but subsequently spread to drier environments as Australia became more arid during the late Cenozoic. We find local assemblages are increasingly phylogenetically clustered along a gradient of decreasing precipitation. The pattern is less clear along a temperature gradient. We develop a novel phyloclimatespace to visualise the expansion of some lineages into drier habitats. Although few species extend into arid regions, those that do occupy larger ranges and thus local species richness does not decline predictably with precipitation.     

Community structure in ichneumonid parasitoids at different spatial scales

The processes underlying parasitoid community structure are little known. Stochastic niche-apportionment models provide one route to underlying assemblage rules in this and other groups. Previous work has applied this approach to parasitoids found on single host species in single populations. However, parasitoid communities are known to extend across multiple hosts and scales. The patterns of relative abundances generated by five niche-apportionment models were compared to those observed in assemblages of two sub-families of the Ichneumonidae, the Diplazontinae and Pimplinae, at landscape and patch scales, Yorkshire, UK. Three of the five models produced patterns that were significantly different to the observed pattern for all taxonomic levels at both spatial scales. The Diplazontinae fit the random fraction (RF) model at the landscape scale in broadleaved woods. This suggests that hierarchical structuring and biotic interactions may play a role in the structuring of Diplazontinae assemblages at this scale. In contrast the Pimplinae fit the RF model only at the patch scale and only at one site. However, the Pimplini tribe (all chiefly parasitoids of Lepidoptera) fit the random assortment (RA) model at both the landscape and the patch scales, whilst the Ephialtini tribe (wide range of hosts) fit no model at either scale. The ecological interpretation of the RA model suggests that the Pimplini tribe is an unsaturated assemblage, where some of the total available resources are unused. Our results show, through the fit of mechanistic niche-apportionment models, that the processes that may structure ichneumonid parasitoid assemblages are not consistent across taxa and spatial scales.     

Nitrogen addition does not reduce the role of spatial asynchrony in stabilising grassland communities

While nitrogen (N) amendment is known to affect the stability of ecological communities, whether this effect is scale‐dependent remains an open question. By conducting a field experiment in a temperate grassland, we found that both plant richness and temporal stability of community biomass increased with spatial scale, but N enrichment reduced richness and stability at the two scales considered. Reduced local‐scale stability under N enrichment arose from N‐induced reduction in population stability, which was partly attributable to the decline in local species richness, as well as reduction in asynchronous local population dynamics across species. Importantly, N enrichment did not alter spatial asynchrony among local communities, which provided similar spatial insurance effects at the larger scale, regardless of N enrichment levels. These results suggest that spatial variability among local communities, in addition to local diversity, may help stabilise ecosystems at larger spatial scales even in the face of anthropogenic environmental changes.     

The impact of ecological differentiation and dispersal limitation on species turnover and phylogenetic structure of inselberg's plant communities

Phylogenetic structure analysis is a novel way to address the relative importance of stochastic and deterministic processes governing species assemblages. Here we investigate the phylogenetic structure of the vegetation of inselbergs located in the African rain forest. Inselbergs combine strong ecological gradients at the local scale due to soil depth variation and insular properties at the regional scale. They are therefore ideal models to assess the influence of ecological sorting and dispersal limitation on the phylogenetic structure of plant communities. On 21 inselbergs separated by up to 200 km where five microhabitat-types were recognized, 311 vegetation plots were inventoried. We found that floristic similarity between plots depended on both microhabitat differentiation and spatial distance, while phylogenetic clustering (i.e. excess of phylogenetic similarity between species from a same plot) only appeared between plots from differentiated microhabitats and increased with ecological distance. Within a microhabitat-type, the absence of phylogenetic structure between inselbergs indicates that species turnover is probably due to dispersal limitation rather than to regional-scale variations in environmental factors. Hence, phylogenetic structure analysis can help disentangle the effects of ecological sorting and dispersal limitation on species assemblages. To estimate the time-scale of the processes generating the phylogenetic structure, we investigated how lineage similarity changes with increasing age in the phylogenetic tree. High lineage similarity levels between ecologically very differentiated plots were only reached at the proximity of the root of the phylogenetic tree. This was observed even when considering only plots sharing no species and indicates that phylogenetic niche conservatism has been important for generating the observed phylogenetic structure. Hence, ancient diversification exerts an impact on the assembly of current plant communities.     

尺度与密度:测定不同尺度下的种群密度

群落中的种群密度由于空间尺度的变化而存在着一定差异,那么,某一种群的密度随着空间尺度的变化会发生怎样的变化?抑或某一物种相对于另一物种而言,随着空间尺度的变化其密度会怎样变化?这是与尺度有关的种群密度问题,当属生态学的基本问题.该文提出这样的问题,并把不同尺度下的种群密度称之为尺度密度(scale density).O-Ring函数的实质是计算不同尺度下的种群密度.因此,在研究实例中,应用O-Ring函数计算了典型草原处于不同恢复阶段的羊草(Leymuschinensis)种群、米氏冰草(Agropyron michnoi)种群,以及米氏冰草种群相对于羊草种群在不同尺度下的种群密度,结果发现:羊草和米氏冰草2个种群的尺度密度,在小尺度范围内严重退化群落均高于恢复演替群落,这一结果验证了“胁迫梯度假说”,同时表明该结果是放牧胁迫下正相互作用所致;通过比较羊草种群与米氏冰草相对于羊草的尺度密度发现,在严重退化的群落中,羊草与米氏冰草的种间关联为负联结,这种负联结是由正相互作用引起的,而在恢复8年和恢复21年群落中,二者之间是正联结,当为竞争所致.该实例说明分析种群密度随尺度变化的规律对于深入认识生态学问题可能会有很大帮助.     

Competitive dominance in the organisation of Mediterranean ant communities

1. Competition by dominant species is thought to be key to structuring ant communities. However, recent findings suggest that the effect of dominant species on communities is less pronounced than previously assumed. 2. The aim of the present study was to identify the role of dominant ants in the organisation of Mediterranean communities, particularly the role of competition in invaded and uninvaded communities. The effects on ant assemblages of two dominant ants, the invasive Argentine ant and the native ant, Tapinoma nigerrimum Nylander, were assessed. 3. The abundances of both dominant ants were significantly correlated with a decrease in native ant richness at traps. However, only the invasive ant was associated with a reduction in diversity and abundance of other ant species at site scale. In the presence of T. nigerrimum, species co‐occurrence patterns were segregated or random. Community structure in both the dominant‐free and the Argentine ant sites showed random patterns of species co‐occurrence. 4. The present findings indicate that dominant ants regulate small‐scale diversity by competition. However, at the broader scale of the assemblage, T. nigerrimum may only affect species distribution, having no apparent effect on community composition. Moreover, we find no evidence that inter‐specific competition shapes species distribution in coastal Mediterranean communities free of dominant ants. 5. These results show that dominant species may affect ant assemblages but that the nature and the intensity of such effects are species and scale dependent. This confirms the hypothesis that competitive dominance may be only one of a range of factors that structure ant communities.