Changing densities of generalist species underlie apparent homogenization of UK bird communities

Generalist species are becoming increasingly dominant in European bird communities. This has been taken as evidence of biotic homogenization, whereby generalist ‘winners’ systematically replace specialist ‘losers’. We test this pattern by relating changes in the average specialization of UK bird communities to changes in the density of species with different degrees of habitat specialization. Although we find the expected decline in community specialization, this was driven by a combination of a strong increase in the density of the most generalist quartile of species and declines in the density of moderately generalist species. Contrary to expectation, specialist species increased slightly over the 18‐year study period but had little effect on the overall trend in community specialization. Our results indicate that the apparent homogenization of UK bird communities is not driven by the replacement of specialists by generalists, but instead by the changing fortunes of generalist species.     

Measuring community responses to large-scale disturbance in conservation biogeography

Aim   Which community metrics should be used to reflect community response to large-scale habitat alterations is unclear. Here, we assess what and how community changes should be measured to accurately track community responses to large-scale disturbance in space and/or time.
Location   France.
Method   We first developed a simulation model to examine temporal changes in the species composition of large-scale metacommunities. Using this model, we assessed how species richness, Shannon index, trends of particular subset of species or community indices of habitat specialization were influenced by different disturbance scenarios, and whether these indices were biased by imperfect detectability. We further used more than 1000 empirical bird communities from the French Breeding Bird Survey recently exposed to disturbances of various intensities as a case study.
Results   Our simulation and empirical results both demonstrate that species richness and diversity measures can show confusing trends and even provide misleading messages of communities' fate. In contrast, reflecting the composition of the community in terms of habitat specialist and generalist species was more robust and powerful to reflect disturbance effects.
Main conclusions   We highlight the weakness of using community metrics that fail to incorporate ecological difference among species when summarizing community-level trends in disturbed landscapes.     

Spatial predictability and resource specialization of bees (Hymenoptera: Apoidea) at a superabundant, widespread resource

For reciprocal specialization (coevolution) to occur among floral visitors and their host plants the interactions must be temporally and spatially persistent. However, studies repeatedly have shown that species composition and relative abundance of floral visitors vary dramatically at all spatial and temporal scales. We test the hypothesis that, on average, pollen specialist bee species occur more predictably at their floral hosts than pollen generalist bee species. Taxonomic floral specialization reaches its extreme among species of solitary, pollen-collecting bees, yet few studies have considered how pollen specialization by floral visitors influences their spatial constancy. We test this hypothesis using an unusually diverse bee guild that visits creosote bush (Larrea tridentatd), the most widespread, dominant plant of the warm deserts of North America. Twenty-two strict pollen specialist and 80 + generalist bee species visit Larrea for its floral resources. The sites we sampled were separated by 0.5 to > 1450 km, and spanned three distinct deserts and four vegetation zones. We found that species of Larrea pollen specialist bees occurred at more sites and tended to be more abundant than generalists. Surprisingly, spatial turnover was high for both pollen specialist and generalist bee species at all distances, and species composition of samples from sites 1–5 km apart varied as much as repeat samples made at single sites. Nevertheless, the pattern of bee species turnover was not haphazard. As distance among sites increased faunal similarity of sites decreased. Faunal similarities among sites within 250 km of each other were generally greater than if randomly distributed over all sites (the null model). No single ecological category of species (widespread, localized, Larrea pollen specialist, floral generalist) accounted for this spatial predictability. Evidently, concordant local distribution patterns of many ecologically diverse species contribute to the non-random spatial pattern. The ecological dominance of creosote bush does not confer obvious ecological advantages to its specialist floral visitors. Spatial turnover is comparable to that found for bee guilds from other biogeographic regions of the world and is not therefore limited to those bee species that inhabit highly seasonal climates, such as deserts. Philopatry and differences in bloom predictability among sites are probably more important causes for spatial turnover of bee species than are interspecific competition for nest sites or floral resources.     

Breakdown of the species-area relationship in exotic but not in native forest patches

We studied the pattern of bird species richness in native and exotic forest patches in Hungary. We hypothesized that species-area relationship will depend on forest naturalness, and on the habitat specialization of bird species. Therefore, we expected strong species-area relationship in native forest patches and forest bird species, and weaker relationship in exotic forest patches containing generalist species. We censused breeding passerine bird communities three times in 13 forest patches with only native tree species, and 14 with only exotic trees in Eastern Hungary in 2003. Although most bird species (92%) of the total of 41 species occurred in both exotic and native forests, the species-area relationship was significant for forest specialist, but not for generalist species in the native forests. No relationship between bird species and area was found for either species group in the forest with exotic tree species. The comparison of native versus exotic forest patches of similar sizes revealed that only large (>100 ha) native forests harbor higher bird species richness than exotic forests for the forest specialist bird species. There is no difference between small and medium forest patches and in richness of generalist species. Thus, the species-area relationship may diminish in archipelago of exotic habitat patches and/or for habitat generalist species; this result supports the warning that the extension of exotic habitats have been significantly contributing to the decline of natural community patterns.     

Differential hepatic gene expression of a dietary specialist (Neotoma stephensi) and generalist (Neotoma albigula) in response to juniper (Juniperus monosperma) ingestion

Dietary specialization is thought to be rare in mammalian herbivores because of limitations of their detoxification system in processing large doses of a single type of plant secondary compound (PSC). Therefore, in order to specialize on a single species of plant, mammalian herbivores must have a highly efficient detoxification system for the particular types of PSCs they ingest. Using microarray technology, we looked at the expression of hepatic genes of a dietary specialist, Neotoma stephensi, and a sympatric generalist, Neotoma albigula, in response to diets containing different levels of one-seeded juniper (Juniperus monosperma). We found large between species differences in gene expression, as well as large within species differences when specialists fed a low juniper diet (25% juniper) were compared to specialists fed their ecologically relevant level of juniper (70% juniper). We also tested the hypothesis that the specialist relies on less costly phase I detoxification enzymes more than phase II compared to the generalist. Although we found that the specialist had higher cumulative as well as average expression of phase I versus phase II enzymes, the generalist had a similar pattern of expression for phase I versus phase II enzymes.     

More and more generalists: two decades of changes in the European avifauna

Biotic homogenization (BH) is a process whereby some species (losers) are systematically replaced by others (winners). While this process has been related to the effects of anthropogenic activities, whether and how BH is occurring across regions and the role of native species as a driver of BH has hardly been investigated. Here, we examine the trend in the community specialization index (CSI) for 234 native species of breeding birds at 10 111 sites in six European countries from 1990 to 2008. Unlike many BH studies, CSI uses abundance information to estimate the balance between generalist and specialist species in local assemblages. We show that bird communities are more and more composed of native generalist species across regions, revealing a strong, ongoing BH process. Our result suggests a rapid and non-random change in community composition at a continental scale is occurring, most likely driven by anthropogenic activities.     

Contrasting effects of habitat reduction,conversion and alteration on neutral and non neutral biological communities

While habitat transformation driven by human activities is the main driver of current biodiversity changes, there is still no framework to explore and forecast the effects of different types of habitat changes on the richness and composition of biological communities. To tackle this issue, we modeled the dynamics of a regional meta‐community, composed either of ecologically equivalent species (neutral model) or of generalist and specialist species (specialization model), and explored the impact of the overall reduction, patch conversion or alteration of an original habitat into one or several other habitats of different total carrying capacity on the community metrics at equilibrium. Our simulations reveal strong interactions between the community model considered (neutral or specialization model) and the type of habitat change. Under neutrality, the impact of habitat changes on richness can be approached by a power law species–individual relationship (SIR), which may at constant density be simplified into the widely used power law species–area relationship (SAR), independent of the type of change. However, in the presence of generalist and specialist species, we found that 1) while habitat reduction in area also leads to approximately power law SIRs and SARs, 2) patch conversion and alteration have more complex effects on regional species richness, and 3) habitat alteration elicits the functional homogenization of communities, involving a decrease of their average level of specialization.     

Use of novel pollen species by specialist and generalist solitary bees (Hymenoptera: Megachilidae)

If trade-offs between flexibility to use a range of host species and efficiency on a limited set underlie the evolution of diet breadth, one resulting prediction is that specialists ought to be more restricted than generalists in their ability to use novel resource species. I used foraging tests and feeding trials to compare the ability of a generalist and a specialist solitary mason bee species to collect and develop on two pollen species that are not normally used in natural populations (novel pollens). Osmia lignaria (Hymenoptera: Megachilidae) is a generalist pollen feeder; O. californica, is more specialized. Adults of the specialist were more limited in use of novel hosts, but only in some contexts. Both bee species refused to collect one novel pollen. The specialist accepted a second novel pollen only when it was presented along with its normal pollen, whereas the generalist collected novel pollen whether presented alone or with normal pollen. Surprisingly, larvae of the specialist were more flexible than were generalists. The specialist grew well on mixtures of normal and novel pollen species, in some cases better than on its normal host alone. Larvae of the generalist grew more poorly on all diets containing novel pollens than on their normal host. Data on these two species of bees suggest that specialization by itself need not reduce flexibility on novel hosts. The findings also provide information about mechanisms of specialization in bees. Similar to some folivores, specific cues of the pollen host and the bee's interpretation of these contribute, along with foraging economics, to pollen choice by adults. The ability of the larvae to cope with specific components of one pollen species need not interfere with its ability to use others.     

Biodiversity and community structure of temperate butterfly species within a gradient of human disturbance: An analysis based on the concept of generalist vs. Specialist strategies

We monitored nine butterfly communities with varying degrees of human disturbance by conducting a census twice a month during 1980 by the line transect method in and around Tsukuba City, central Japan. We analyzed the biodiversity and community structures using the generalist/specialist concept. The site (community) order based on decreasing human disturbance was positively correlated with butterfly species diversity (H′), species richness (the total number of species), and the number of specialist species in a community, but not with the number of generalist species. The number of generalist species was rather constant, irrespective of the degree of human disturbance. Thus, both the butterfly species diversity and species richness were more dependent on the specialists than the generalists. Our analyses also showed that the generalist species were distributed widely over the communities, and they maintained high population densities, resulting in high rank status in abundance in a community, with more spatial variation in density per species. Specialist species showed the opposite trends. These results demonstrate that the generalist/specialist concept is a powerful tool applicable to analyse the biodiversity and structure of natural communities.