Does specialization explain rarity and decline among British bumblebees? A response to Goulson et al.

Recent studies of British bumblebees have proposed a seemingly simple explanation for the decline in some species: that greater dietary specialization among the rarer species has put them at greater risk. However, comparisons of dietary specialization require: (1) that bees have access to the same dietary options among which to make their choices; (2) that the differing relative breadths of dietary choices made are not obscured by the differing sample sizes among bee species. Using one of the few suitable data sets, I find no evidence for a relationship between, on the one hand, rarity or declines in British bumblebees and, on the other, their dietary breadths, the strengths of their dietary preferences, or their proboscis lengths (which influence dietary choices). In contrast, there is support for a relationship between rarity or declines within Britain and the sizes of species’ European ranges, particularly when these measures are adjusted to represent their ranges near sea level. Adjusted range sizes may reflect overall niche breadth and perhaps climatic and habitat specialization. This is not to say that climate change is the driving factor for declines or that changing food-plant availability is unimportant, but that climatic and habitat specialization may be a better indicator of risk of decline, which deserves further study.     

Declines in forage availability for bumblebees at a national scale

We assessed national scale changes in the forage plants of bumblebees in Britain, as a means of providing quantitative evidence for the likely principal cause of declines in bumblebee species. We quantified the relative value of native and long-established plant species as forage (nectar and pollen) resources for bumblebees by collating visitation data from 14 field sites across Britain. Twentieth Century changes in range and frequency of these forage plants were assessed using data from the New Atlas of the British and Irish Flora (1930-1969 to 1987-1999) and the Countryside Surveys of Britain (1978-1998). Forage plants declined in both large-scale range and local-scale frequency between the two survey periods. These changes were of greater magnitude than changes in other native plant species, reflecting serious reductions in quality of foraging habitats for bees as well as a general decline in insect-pollinated plants. Seventy-six percent of forage plants declined in frequency within 1-km squares, including those (e.g. Trifolium pratense) of particular value for threatened bumblebee species. We consider how our findings relate to other recorded changes in the British flora, how they may help to explain declines in bumblebees and how they could contribute to a conservation strategy.     

The use of historical collections to estimate population trends: A case study using Swedish longhorn beetles (Coleoptera: Cerambycidae)

Long term data to estimate population trends among species are generally lacking. However, Natural History Collections (NHCs) can provide such information, but may suffer from biases due to varying sampling effort. To analyze population trends and range-abundance dynamics of Swedish longhorn beetles (Coleoptera: Cerambycidae), we used collections of 108 species stretching over 100 years. We controlled for varying sampling effort by using the total number of database records as a reference for non-red-listed species. Because the general frequency of red-listed species increased over time, a separate estimate of sampling effort was used for that group. We observed large interspecific variation in population changes, from declines of 60% to several hundred percent increases. Most species showed stable or increasing ranges, whereas few seemed to decline in range. Among increasing species, rare species seemed to expand their range more than common species did, but this pattern was not observed in declining species. Historically, rare species did not seem to be at larger risk of local extinction, and population declines were mostly due to lower population density and not loss of sub-populations. We also evaluated the species’ declines under IUCN red-list criterion A, and four currently not red-listed species meet the suggested threshold for Near Threatened (NT). The results also suggested that species’ declines may be overlooked if estimated only from changes in species range.     

Major declines in the abundance of vultures and other scavenging raptors in and around the Masai Mara ecosystem, Kenya

Vulture population declines have been noted in West and Southern Africa, but have not been assessed in East Africa. Roadside transects conducted in 1976 and 1988 were compared with surveys done from 2003–2005 in and around Masai Mara National Reserve, Kenya. Staggering declines in abundance were found for seven of eight scavenging raptors surveyed. No Egyptian vultures were seen during recent transects. We compared trends between the ungulate migration and non-migration season among three land use types (reserve, buffer, and grazed) and among the species surveyed to establish the causes of declines in scavenging raptors. Large declines during the ungulate migratory period suggest that most scavenging raptor species are declining well beyond the area of study. For all species, except Hooded vultures, substantial declines outside of the reserve indicate an important role of land use change in causing observed declines. In addition, significant declines of populations of Gyps species in the reserve itself, especially during the migration season, provide evidence that human activities occurring in other parts of the species’ range such as poisoning of carcasses may be causing their decline. Declines found in this study suggest that at a minimum African white-backed, Rüppell’s, and Hooded vultures should be relisted as Vulnerable. Management actions that limit land use change around the reserve combined with a countrywide ban on carbamate pesticides will be important for conserving keystone members of the scavenging guild. Future research should further examine possible causes of these declines and quantify the effect of reduced scavenging raptor abundance on scavenging efficiency.     

Effects of non-cropped landscape diversity on spatial dynamics of farmland birds in intensive farming systems

Increasing habitats diversity in agricultural landscapes has been proposed as a key measure for reversing the decline of farmland biodiversity in Europe. However, indicators used for assessing such a potential compensation effect usually only rely on species diversity and abundance while ignoring variations in species-specific vulnerability. The extent to which habitat diversity may reverse the decline of specialist species in Europe to farming systems is thus still unclear. In this study, we investigate whether the effect of non-cropped habitat diversity on farmland birds’ occurrences was dependent on species’ specialization for habitats. In particular, we focused on the relative effects of non-cropped habitat diversity on species’ abilities to persist or to colonize new vacant areas. We used a capture-recapture statistical framework to study the spatial dynamics of 20 farmland bird species in France monitored from 2001 to 2007. We found that non-cropped landscape diversity reduces both the probabilities that a species becomes extinct locally and that a species colonizes new vacant areas, and the occupancy rate. Although this suggests a possible stabilizing effect of the surrounding habitat diversity on species occurrence in farming systems, the occupancy was only weakly affected. Moreover, we found that the most specialist species were the more negatively affected by this landscape diversity in terms of colonization abilities. We argue that accounting for the differences in habitat specialization among farmland species can improve conservation policies dedicated to the management of landscape diversity.     

Crofting and bumblebee conservation: The impact of land management practices on bumblebee populations in northwest Scotland

The northwest of Scotland is a stronghold for two of the UK’s rarest bumblebee species, Bombus distinguendus and Bombus muscorum. The predominant form of agricultural land management in this region is crofting, a system specific to Scotland in which small agricultural units (crofts) operate rotational cropping and grazing regimes. Crofting is considered to be beneficial to a wide range of flora and fauna. However, currently there is a lack of quantitative evidence to support this view with regard to bumblebee populations. In this study we assessed the effect of land management on the abundance of foraging bumblebees and the availability of bumblebee forage plants across crofts in northwest Scotland. The results of our study show that current crofting practices do not support high densities of foraging bumblebees. Traditional crofting practice was to move livestock to uplands in the summer, but this has been largely abandoned. Summer sheep grazing of lowland pasture had a strong negative impact on bumblebee abundance and forage plant availability throughout the survey period. The use of specific ‘bird and bee’ conservation seed mixes appears to improve forage availability within the crofted landscape, although the number of bees observed remained low. Of the forage plants available, the three most frequently visited species were from the Fabaceae. We therefore conclude that the creation of agri-environment schemes which promote the use of Fabaceae-rich seed mixes and encourage the removal of sheep grazing on lowland areas throughout the summer are essential in order to conserve bumblebee populations within crofted areas.     

Long-term population declines in Afro-Palearctic migrant birds

We present the first continent-wide analysis of the population trends of European breeding birds to show that populations of Afro-Palearctic migrant birds have shown a pattern of sustained, often severe, decline. The mean trend of inter-continental migrants was significantly negative between 1970 and 1990 and non-significantly so between 1990 and 2000. Mean population trends were positively correlated between periods, suggesting little change in the trajectory of most migrant species’ populations over this 30-year period. In both periods, trends of inter-continental migrants were significantly more negative than those of short-distance migrants or residents. This negative trend appeared to be largely, although not entirely, due to declines in species wintering in dry, open habitats in Africa. Analyses of trends of 30 closely related pairs of species, one a long-distance migrant and the other not, indicated significantly more negative trends in the former, irrespective of breeding habitat. Conservation action to address these declines is required under the Convention on Migratory Species and the Pan-European Biological and Landscape Diversity Strategy, to which most European countries are signatories and which aim, respectively, to conserve migratory species and to halt the loss of biodiversity by 2010. Our results indicate that more conservation action may be required outside Europe to achieve these targets. Further research is needed to assess whether the declines are caused by factors operating on the birds’ wintering grounds, breeding grounds or on migration routes, and to identify ways to reverse them.     

The response of foraging bumblebees to successional change in newly created arable field margins

Agricultural intensification is likely to have been a major factor leading to serious declines in bumblebee abundance and diversity in the UK and elsewhere over recent decades. Opportunities to restore habitat for bumblebees on uncropped arable field margins are now available, although the methods by which this restoration can be achieved have not been fully investigated. We present the results of a three year study undertaken to investigate the response of foraging bumblebees to five different arable field margin treatments (sown and unsown), as part of a replicated field experiment on arable farmland in northern England (UK). Bumblebee abundance was closely linked to successional changes in availability of suitable forage plant species. Field margin treatments sown with a `grass and wildflower' mixture had the highest bumblebee abundance, and provided a consistent supply of forage species, with different components of the seed mixture flowering in each year. The unsown natural regeneration treatment attracted foraging bumblebees in only the second year due to the local abundance of thistles, so we consider this option to be both inconsistent in terms of forage provision and agronomically unacceptable. Our results are discussed in terms of developing suitable measures to achieve the restoration of habitats for bumblebees on arable farmland.     

Climate change and translocations: The potential to re-establish two regionally-extinct butterfly species in Britain

Climate change is causing many organisms to migrate to track climatically-suitable habitat. In many cases, this will happen naturally, but in others, human intervention may be necessary in the form of ‘assisted colonisation’. Species re-establishments in suitable parts of their historic ranges provide an opportunity to conserve some species and to test ideas about assisted colonisation. Here, bioclimatic models of the distributions of two extinct British butterflies, Aporia crataegi and Polyommatus semiargus, were used to investigate the potential for re-establishment in Britain. Generalised additive models and generalised linear models were created to describe the species’ European distributions for the period 1961–1990. All models projected the British climate during this period to be suitable for both species. Thirty-year climate projections for the periods 1991–2020 and 2021–2050, and for three climate change scenarios, were then put into the models to generate projections of climatic suitability throughout the 21st century. British climate was projected to remain highly suitable for A. crataegi, but to decline somewhat for P. semiargus. Southern and eastern Britain were found to be the areas most likely to support suitable climate. This difference between the species appeared to be due in part to decreasing summer rainfall in climate change projections, as this should only benefit A. crataegi. It is concluded that, with further study of habitat requirements, both species could be reintroduced to Britain as part of a long-term European conservation strategy.     

Commercialized European bumblebee can cause pollination disturbance: An experiment on seven native plant species in Japan

Non-native pollinator species are now widely utilized to facilitate pollination of agricultural crops. Evaluation of the ecological risk of alien pollinators is necessary because they could have a large impact on native ecosystems through disturbing native plant-pollinator interactions. We conducted a greenhouse experiment to examine the impact of the non-native commercialized European bumblebees, Bombus terrestris, on the pollination success of seven Japanese bumblebee-pollinated plant species. Plants were exposed to three groups of bumblebees: native bumblebee(s) only (NATIVE treatment); the alien bee only (ALIEN) and a mix of the two (MIX). ALIEN treatment had negative effects on fruitset and/or fruit quality of five plants, including self-incompatible and compatible, herb and woody, and queen- and worker-pollinated species. The negative effects were caused by a decrease in legitimate flower visitation due to (1) physical inaccessibility to nectary in deep-corolla flowers by the alien bee with insufficient tongue length and, (2) biased flower preference between short-corolla flowers. Fruitset tended to decrease drastically for the self-incompatible species while fruit quality decreased moderately for the self-compatible species. Effects of MIX were not intermediate between NATIVE and ALIEN in most plant species, and caused pollination success to vary in an unpredictable manner amongst plant species, probably due to interaction between native and alien bees. This non-linear relationship between plants’ pollination success and the relative density of the alien suggests that the alien bee can disturb pollination of a plant species even when only representing a small fraction of the total pollinator community.     

New perspectives on Holocene flooding in Ireland using meta-analysis of fluvial radiocarbon dates

The last decade has seen advances in models of Holocene river flooding founded on meta-analysis of fluvial ¹⁴C databases allied to new multivariate environmental correlatives. This investigation presents application of these generic meta-analysis techniques to 33 stratigraphically significant ‘change after’ dates in a new fluvial ¹⁴C database for Ireland. Despite the relatively small number of ¹⁴C dates, the emerging pattern of Holocene flooding in Ireland corresponds closely to palaeoclimate proxies for regional temperature and precipitation, and to recently published results from a much larger British database. An underlying climate forcing of fluvial activity is proposed, although Irish ‘flooding episodes’ appear to lag those in Britain by ca. 100 to 300 years. This may be caused by ¹⁴C date precision and the embryonic nature of the Irish database, but could also reflect the respective effects of peatland cover and glacial inheritance on hydrological connectivity and sediment delivery resulting in a slower response of Irish rivers to climate events. These considerations, together with an increasing focus on regional variations in fluvial activity across the Holocene, will only be properly addressed with a more concerted and expanded programme of research in Ireland. Renewed research focus should seek to broaden the geographical coverage of ¹⁴C dated fluvial sites, with particular emphasis given to collaborative research of Irish lowland river catchments, especially where suitable palaeochannel and flood basin depositional contexts are present.     

The effect of arable field margin composition on invertebrate biodiversity

A replicated field experiment designed to compare five types of field margin in terms of their invertebrate biodiversity was established in North Yorkshire, UK. Four replicate margins on arable land each contained five treatment plots 72 m long by 6 m wide. The treatments were: (1) cropped to the edge; (2) sown ‘tussocky’ grass mix; (3) sown ‘grass and wildflower’ mix; (4) Split margin (3 m ‘tussocky’ grass adjacent to hedge and 3 m ‘grass and wildflower’ next to crop); and (5) natural regeneration. Invertebrates were sampled by pitfall trapping, sweep netting, and butterfly and bumblebee transects, and identified to species level. The use of different flower species by foraging bumblebees was also examined.Despite all treatments containing a flush of annual weeds early in the establishment year, the five margin types were distinct in their vegetation composition by their first mid-summer. Where statistically significant results were obtained for invertebrates, avoidance of the ‘cropped’ treatment was by far the commonest observed response. This tendency could be clearly demonstrated amongst the carabids, spiders, butterflies, bumblebees, millipedes and harvestmen, with margins often containing double or more the number of invertebrates of similar areas cropped to the edge. Having avoided the crop, preferences for other margin types were mixed, but there was a marked tendency by many nectar and/or pollen-feeding, flying insects towards greater abundance on those margins containing sown ‘wildflowers’ (e.g. butterflies Meadow Brown and Ringlet, Bumblebees, Pollen Beetle Meligethes sp.) or flowers either sown or unsown (Soldier Beetle Rhagonycha fulva). There was a less strong tendency for predatory species occurring in the vegetation canopy and depending on small, especially flying insects for food, to also prefer the flowery treatments (e.g. total spiders caught by sweeping, seven-spot ladybird). Harvestmen in autumn reject natural regeneration in favour of any sown treatment. Only one species, the carabid Nebria brevicollis, was trapped in higher numbers in the cropped treatment than on any sown margin, and then only in autumn. Of the six common bumblebee species, the two longest-tongued species showed different patterns of flower visitation from the remaining four.These results clearly demonstrate that sown field margins can rapidly produce substantial biodiversity benefits on arable land, with the resulting fauna influenced by the type of field margin created.     

The “Crex crex” lament: Estimating landowners willingness to pay for corncrake conservation on Irish farmland

This paper considers farmers’ willingness to pay (WTP) to conserve an endangered Irish farmland bird, the corncrake (Crex crex). An Irish National Farm Survey is used to produce individual farm-level WTP estimates for the year 2006. These figures are then aggregated to obtain a total value figure for the farming community of corncrake conservation in Ireland. We focus on the willingness to pay of farmers rather than the WTP of the general Irish population, as farmers will ultimately be the ones that will have to take responsibility if targets set out in the All Ireland Action Plan for corncrake conservation are to be achieved. Quantifying willingness to pay on the part of farmers can help inform the design of agri-environment schemes aimed at improving conservation of many bird species on farmland. Results indicate that the non-market benefit of corncrake conservation in Ireland may significantly outweigh the costs of existing conservation schemes.     

Regional population viability of grassland songbirds: Effects of agricultural management

Although population declines of grassland songbirds in North America and Europe are well-documented, the effect of local processes on regional population persistence is unclear. To assess population viability of grassland songbirds at a regional scale (∼150,000 ha), we quantified Savannah Sparrow Passerculus sandwichensis and Bobolink Dolichonyx oryzivorus annual productivity, adult apparent survival, habitat selection, and density in the four most (regionally) common grassland treatments. We applied these data to a female-based, stochastic, pre-breeding population model to examine whether current grassland management practices can sustain viable populations of breeding songbirds. Additionally, we evaluated six conservation strategies to determine which would most effectively increase population trends. Given baseline conditions, over 10 years, simulations showed a slightly declining or stable Savannah Sparrow population (mean bootstrap λ = 0.99; 95% CI = 1.00-0.989) and severely declining Bobolink population (mean bootstrap λ = 0.75; 95% CI = 0.753-0.747). Savannah Sparrow populations were sensitive to increases in all demographic parameters, particularly adult survival. However for Bobolinks, increasing adult apparent survival, juvenile apparent survival, or preference by changing habitat selection cues for late-hayed fields (highest quality) only slightly decreased the rate of decline. For both species, increasing the amount of high-quality habitat (late- and middle-hayed) marginally slowed population declines; increasing the amount of low-quality habitat (early-hayed and grazed) marginally increased population declines. Both species were most sensitive to low productivity and survival on early-hayed fields, despite the fact that this habitat comprised only 18% of the landscape. Management plans for all agricultural regions should increase quality on both low- and high-quality fields by balancing habitat needs, nesting phenology, and species’ response to management.     

Habitat use and conservation of bumblebees (Bombus spp.) under different grassland management regimes

Declines in the natural populations of several bumblebee species across Britain and Europe are an increasing cause for concern. In this study the habitat use of bumblebees was investigated on Salisbury Plain Training Area, the largest remaining area of unimproved chalk grassland in north-west Europe. Habitat characteristics influencing the overall abundance, species richness and foraging activity of bumblebees included the diversity and abundance of flowering plant species (particularly of favoured forage plants such as Trifolium pratense), vegetation structure and height. It is suggested that different Bombus species respond to these habitat characteristics depending on their specific foraging and nesting requirements, the case of Bombus humilis being especially relevant. The effects of several grassland management practices were considered in terms of their suitability for the conservation of bumblebee habitats. Cattle grazing was shown to be preferable to both sheep grazing and the absence of any management, although the timing and intensity of such grazing was important. Small-scale disturbances caused by vehicle activity were also of value in producing locally abundant forage resources in less intensively managed grasslands.     

Extinction debt or habitat change? - Ongoing losses of woodland birds in north-eastern New South Wales, Australia

The loss, fragmentation and degradation of native vegetation are major causes of loss of biodiversity globally. Extinction debt is the term used to describe the ongoing loss of species from fragmented landscapes long after the original loss and fragmentation of habitat. However, losses may also result from habitat changes that are unrelated to fragmentation, which reduce breeding success and recruitment. Many woodland birds have declined in fragmented landscapes in Australia, probably due to loss of small, isolated populations, though the ecological processes are poorly understood. We record the progressive regional loss of two ground-foraging, woodland birds, the Brown Treecreeper Climacteris picumnus and Hooded Robin Melanodryas cucullata, in northern New South Wales, over 30 years. This has happened despite most habitat loss occurring over 100 years ago, suggesting the payment of an extinction debt. Our observations suggest that several ecological processes, caused by habitat loss, fragmentation or degradation, and operating over different time scales, have led to both species’ declines. Female Brown Treecreepers disperse poorly among vegetation remnants, leaving only males in isolated populations, which then go extinct. In contrast, Hooded Robins suffer high nest predation in fragmented landscapes, producing too few recruits to replace adult mortality. Foraging by both species may also be affected by regrowth of ground vegetation and shrubs. We found little support for a major role played by drought, climate change or aggressive Noisy Miners Manorina melanocephala. We propose that both extinction debt in the classical sense and ongoing habitat change frequently contribute to species’ decline in modified landscapes. Management to arrest and reverse such declines needs to consider these multiple causes of decline. For instance, reconnecting isolated populations may be inadequate alone, and activities such as appropriate grazing, fires and the addition of woody debris may also be required.     

Dragonfly crisis in Japan: A likely consequence of recent agricultural habitat degradation

Many Japanese dragonfly species depend on habitat complexes maintained in rice paddy systems. We postulated that recent alterations to habitat complexes in paddy systems have had adverse effects on dragonfly populations, especially those ‘once common species’ that have come to depend primarily on paddy systems following losses of natural floodplain habitats. A high proportion of Japanese lentic dragonfly species depends on paddy fields or agricultural ponds that have been extensively degraded, while lotic species can often use both paddies and natural river systems. Thus we also postulated that lentic species are more susceptible to changes in agricultural habitats and are subject to higher extinction risks than lotic species. We aimed to extend previous work on estimating dragonfly extinction risk by developing mechanistic insights into the processes involved. Postulates were tested by analyzing relationships between (1) previous quantitative extinction risk assessments for dragonfly species and (2) species’ ecological characteristics (i.e., distribution range and habitat type [lentic or lotic]). Lentic species were disproportionately represented among those with elevated extinction risk. Species with large distribution ranges were also subject to higher extinction risks than those with narrower ranges, reflecting a driving force acting at a national scale (i.e., intensive degradation of paddy systems).     

Using integrated population modelling to quantify the implications of multiple threatening processes for a rapidly declining population

Many species of conservation concern are in decline due to threats from multiple sources. To quantify the conservation requirements of these species we need robust estimates of the impact of each threat on the rate of population decline. However, for the vast majority of species this information is lacking. Here we demonstrate the application of integrated population modelling as a means of deriving robust estimates of the impact of multiple threats for a rapidly declining koala (Phascolarctos cinereus) population in South-east Queensland, Australia. Integrated population modelling provides a basis for reducing uncertainty and bias by formally integrating information from multiple data sources into a single model. We quantify mortality rates due to threats from dog attacks, vehicle collisions and disease and the extent to which each of these mortality rates would need to be reduced, or how much habitat would need to be restored, to stop the population declining. We show that the integrated population modelling approach substantially reduces uncertainty. We also show that recovery actions that only address single threats would need to reduce those threats to implausibly low levels to recover the population. This indicates that strategies for simultaneously tackling multiple threats are necessary; a situation that is likely to be true for many of the world’s threatened species. This study provides an important framework for quantifying the conservation requirements of species undergoing declines due to multiple threats.