PK {< refs.MYD ? + Cameron, S. A.
Hines, H. M.
Williams, P. H. 20075 A comprehensive phylogeny of the bumble bees (Bombus) 161-188) Biological Journal of the Linnean Society 91 Bumble bees (Bombus Latreille) occupy a wide diversity of habitats, from alpine meadows to lowland tropical forest, yet they appear similar in morphology throughout their range, suggesting that behavioral adaptations play a more important role in colonizing diverse habitats. Notwithstanding their structural homogeneity, bumble bees exhibit striking inter- and intraspecific variation in color pattern, purportedly the outcome of mimetic evolution. A robust phylogeny of Bombus would provide the framework for elucidating the history of their wide biogeographic distribution and the evolution of behavioral and morphological adaptations, including color pattern. However, morphological studies of bumble bees have discovered too few phylogenetically informative characters to reconstruct a robust phylogeny. Using DNA sequence data we report the first nearly complete species phylogeny of bumble bees, including most of the 250 known species from the 38 currently recognized subgenera. Bayesian analysis of nuclear (opsin, EF-1a, arginine kinase, PEPCK) and mitochondrial (16S) sequences results in a highly resolved and strongly supported phylogeny from base to tips, with clear-cut support for monophyly of most of the conventional morphology-based subgenera. Most subgenera fall into two distinct clades (short-faced and long-faced) associated broadly with differences in head morphology. Within the short-faced clade is a diverse New World clade, which includes nearly one-quarter of the currently recognized subgenera, many of which are restricted to higher elevations of Central and South America. The comprehensive phylogeny provides a firm foundation for reclassification and evaluating character evolution in the bumble bees.$ doi:10.1111/j.1095-8312.2007.00784.x ? + Hines, H. M.
Cameron, S. A.
Williams, P. H. 2006 Molecular phylogeny of the bumble bee subgenus Pyrobombus (Hymenoptera: Apidae: Bombus) with insights into gene utility for lower-level analysis 289-303 Invertebrate Systematics 20 Species of the largest bumble bee subgenus, Pyrobombus, possess too few known morphological synapomorphies to reveal a clear pattern of their relationships. To obtain a robust phylogeny of the group, we use sequence data from four genes (mitochondrial 16S rRNA, and three nuclear genes: elongation factor-1 alpha (EF-1a), long wavelength rhodopsin (LW Rh or opsin), and arginine kinase (ArgK)) for 36 of the 43 Pyrobombus species. The resulting phylogeny is well resolved and indicates a monophyletic Pyrobombus when assessed against representatives of 20 additional subgenera. The sister group to Pyrobombus comprises the Alpinobombus+Bombus sensu stricto clade. We also assess the utility of these genes for resolving species-level relationships. The more conserved nuclear genes, especially EF-1a and ArgK, performed better than 16S in nearly all measures of gene utility, with 16S providing resolution mostly at the tips of the tree. The exon regions of ArgK were the most conserved and may be promising for higher-level phylogenetics. We discuss species relationships relative to previous taxonomic studies and their implications for commercial rearing of Bombus. *? Goulson, D.
Williams, P. 2001? Bombus hypnorum (Hymenoptera: Apidae), a new British bumblebee? 129-1311 British Journal of Entomology and Natural History 14 A male of the bumblebee Bombus hypnorum (Linnaeus) has been collected in Wiltshire. This species has not been recorded previously from Britain and its identification and status are discussed. ? j Williams, P. H. 1982B The distribution and decline of British bumble bees (Bombus Latr.) 236-245 Journal of Apicultural Research 21 The suggested recent decrease in the distributional ranges of certain species of bumble bees was investigated in the data collected by the Bumblebee Distribution Maps Scheme, using a numerical classificatory approach. Three major biogeographic elements and four biogeographic regions are defined, and changes in their composition and distribution described. Large reductions in the distributional ranges recorded after 1960 were found for two of the biogeographic elements (especially for the Southern Local Species: Bombus subterraneus (L.), B. sylvarum (L.), B. ruderatus (F.) and B. humilis (Ill.), which have resulted in the emergence of a new Central Impoverished Region covering 23 vice-counties in Central England.'? = Williams, P. H. 1985b A preliminary cladistic investigation of relationships among the bumble bees (Hymenoptera, Apidae) 239- 255 Systematic Entomology 10 b/ The cladistic method is used to classify sixty species of bumble bees by characters of the male genitalia selected for their functional importance. A system of three genera is recommended, preserving the holophyletic groups Mendacibombus Skorikov, Psithyrus Lepeletier and Bombus Latreille, with the retention of the named subgroups of Bombus in subgeneric status. A new subgenus Brachycephalibombus is recognized in order to avoid paraphyletic groupings. The phylogenetic interpretation of the cladogram is considered and biogeographic problems discussed. ? Williams, P. H. 1985t On the distribution of bumble bees (Hymenoptera, Apidae) with particular regard to patterns within the British Isles 180 Department of Applied Biology Cambridge University of Cambridge Ph. D.
If ecology can be considered as the study of the distribution and abundance of organisms, then the mechanisms governing the distributions of bumble bees are not only of practical importance to pollination and conservation interests, but also of primary theoretical concern. In this dissertation the distributions of bumble bees are examined at successively finer scales and patterns described from analyses of surveys.
The cladistic method is used to classify sixty species by characters of the male genitalia selected for their functional significance. The phylogenetic interpretation of the composite character state tree is considered in conjunction with present patterns in distribution between continents for the evaluation of possible histories of spread and speciation.
Regional patterns of distribution within England, Wales and Scotland in the Bumblebee Distribution Maps Scheme data are described using a classificatory approach. Three major biogeogarphic elements and four biogeographic regions are defined. Large reductions in the distributional ranges recorded after 1960 are found for two of the elements, which have resulted in the emergence of a new Central Impoverished Region encompassing twenty-three vice-counties in central England.
Local patterns of distribution are described from Kent within the Southern Region, the sole region wherein all three elements retain a strong representation after 1960. Those species that occur only in particular regions of Britain after 1960 are also very locally restricted within this region and less abundant where present. They are further associated with certain open habitats, characterised by coastal and old meadow vegetation. This may be indicative of a relative lack of disturbance. A general mechanism is proposed, founded here on considerations of the economics of energy, to explain a species' pattern of distribution, in which fragmentation at the margins of the distribution occurs in response to a mosaic of effectively only two types of habitat presenting differing fundamental levels of limiting resources.
Coexistence is examined within a small area of Dungeness in Kent, where there is a particularly large assemblage of species. Flower choice by worker bumble bees and partitioning of nectar resources is compared with that at Shoreham, which is a more typical, species-poor site in north Kent. The possible causes and effects of differences in the flora utilised by bumble bees are discussed. The marginal mosaic model is used to predict an alternative form of resource-dependent 'structure' in local species assemblages that is amenable to testing.
Finally, the apparent decline of the British bumble bees nearer the edges of their distributions is discussed with regard to factors including climate and changes in land use.& Ph.D. thesis, University of Cambridge. ? j Williams, P. H. 1986P Environmental change and the distributions of British bumble bees (Bombus Latr.) 50-61 Bee World 67 [None] ? j Williams, P. H. 1988( Habitat use by bumble bees (Bombus spp.) 223-237 Ecological Entomology 13 1. Analysis of surveys of bumble bee distribution among 2 km grid-squares in Kent revealed that some species are nearly ubquitous among localities and abundant within each. For these ubiquitous species, Kent is near the middle of their latitudinal ranges.
2. The other species have very restricted distributions among localities and are less abundant where they occur. For each of these local species, Kent is near the margin of its latitudinal range.
3. The areas to which the local species are restricted are characterized by only a few of the many kinds of vegetation represented in Kent (shingle, sand dune, saltmarsh or old meadow). The ubquitous species are also more abundant in areas with these kinds of vegetation.
4. A random model (Gause, 1936) and the core/satellite hypothesis (Hanski, 1982a) were proposed to explain similar patterns of distribution. I conclude that their assumptions are not well supported by the results of the present analysis.
5. A marginal mosaic model (after Andrewartha & Birch, 1954; Hengeveld & Haeck, 1981; Brown, 1984) is used to account for these patterns. This model depends on environmental factors (including resource levels and climate) affecting the economics of energy and, consequently, local persistence.
6. It follows from the marginal mosaic model that if the level of limiting resources in a habitat declines, then it is the species closest to their distribution limits that are most likely to face local extinction. These selective extinctions would result in a form of 'community structure'. Մ? Williams, P. H. 1989* Bumble bees - and their decline in Britain 15 Ilford" Central Association of Bee-Keepers [None]C http://www.nhm.ac.uk/research-curation/projects/bombus/decline.html ?
Williams, P. H. 1989: Why are there so many species of bumble bees at Dungeness? 31-44( Botanical Journal of the Linnean Society 101 Dungeness is unique in the British Isles in that it has more species of bumble bees than any other locality. Three ideas about what governs the numbers of species at a locality are examined by looking at patterns of flower visits at Dungeness in comparison with those at Shoreham, a species-poor locality also in Kent. The species of bumble bees that are present at Dungeness but absent from Shoreham show no association in their distributions among 2 km grid-squares in Kent with the species of food-plants that they prefer at Dungeness, nor is there any correlation between the diversity of bees and diversity of food-plants at Dungeness and Shoreham. From the information available, Dungeness is most likely to have more species of bumble bees because it has a particularly high density of the more nectar-rich flowers that bumble bees can use. Bumble bees feed most profitably from deep flowers because these contain more nectar than shallow flowers, although direct access to deeper flowers is ultimately limited by the length of each bee's proboscis. The distribution of worker proboscis lengths among species in the species-pool in Kent is clumped about a median of 7.9 mm. The best foraging conditions for the maximum number of species should be provided when flowers of similar depths are present in sufficiently large numbers for all foragers to make near-optimal flower choices. Although there is no difference in median between the distribution of the bees' proboscis lengths and the depths of the flowers they use at Dungeness, at Shoreham the flower depths used are shorter than the proboscis lengths. Among the food-plants at Dungeness, high densities of Teucrium scorodonia and Echium vulgare are likely to be especially important.? o Williams, P. H. 1991F The bumble bees of the Kashmir Himalaya (Hymenoptera: Apidae, Bombini) 1-204= Bulletin of the British Museum (Natural History) (Entomology) 60 Bumble bees are extraordinarily variable in the colour patterns of their pubescence. This has contributed to the lack of agreement among authors concerning the discrimination of species. Material from Kashmir has been listed previously under 80 names, but for the broadest interpretations of these species, more than 500 names have been published for differing individuals from throughout their distributions. In this review, all of the available voucher material from Kashmir (6312 museum specimens from 78 sites) is assigned to just 29 species. The reduction in the number of nominal taxa that are inferred to represent separate species depends on two factors. First, variation is examined among the larger samples of new material. Second, current species concepts require a re-assessment of this evidence for the limits of reproductive cohesion in space and time. A particular study is made of the variation in male mate-searching behaviour and of its relationship to the different kinds of habitat. For Bombus asiaticus, patterns of colour variation across the Great Himalaya are compared with those expected from simple genetic models. Type material is examined for 103 of the nominal specific and subspecific taxa. Seventeen lectotypes are designated and 80 new synonyms and provisional new synonyms are established. Keys to the species are accompanied by diagrams of the colour patterns and by distribution maps. ? Williams, P. H. 1993[ Measuring more of biodiversity for choosing conservation areas, using taxonomic relatedness 194-227> International Symposium on Biodiversity and Conservation (KEI)
T. Y. Moon Seoul One of the major goals of conservation is the maintenance with only limited resources of as much as possible of the variety of life. If we are to choose among areas in order to protect the greatest overall amount of biodiversity, then we shall need to be able to measure and compare biodiversity among areas. This has usually been measured only in terms of species richness. Diversity also includes a concept of difference, and the degree of difference between organisms can be represented in biodiversity measures using readily available information on group membership from taxonomic classifications. Furthermore, by using the complementarity in species composition between faunas, stepwise procedures can identify optimal sequences of priority areas for biodiversity protection, taking existing protected areas into account or not, as required. In some circumstances it may be possible to apply this approach to higher taxa, rather than to species. This could greatly reduce survey costs, allowing survey effort to be re-deployed to cover much more of overall biodiversity. These methods are illustrated by their application to the bumble bees and milkweed butterflies. ?
Williams, P. H. 1995d Phylogenetic relationships among bumble bees (Bombus Latr.): a reappraisal of morphological evidence 327-344 Systematic Entomology 19 Cladistic analyses of morphological characters of adult bumble bees show significantly higher congruence among characters of the male genitalia than among other characters of males and females. Phylogenetic interpretations of groups supported by the combined morphological evidence (even with minimal assumptions) include (1) that the earliest diverging species belong to the paraphyletic subgenus Mendacibombus, together with Bombus nevadensis and B. confusus; (2) that, in contrast, species of the subgenus Bombus (sensu stricto) share a relatively recent common ancestor with B. pressus and B. rufipes; and (3) that Psithyrus (the social parasites) is a monophyletic group, and the sister-group of B. persicus. For genus-group nomenclature a single genus Bombus (sensu lato) is recommended, to include Psithyrus as another subgenus. Names for sections and for groups of subgenera are synonymized with subgeneric names. B? Williams, P. H. 1996g Mapping variations in the strength and breadth of biogeographic transition zones using species turnover 579-588. Proceedings of the Royal Society of London (B) 263 bl Biogeographic regions are widely regarded as real entities, or at least as useful summaries of the complex patterns of spatial concordance amongst species. The problem is that, whereas some parts of the transition zones between regions may be strong and abrupt boundaries, other parts of the same zones may be weak or broad, so that the corresponding parts of border lines drawn on maps, although convenient, are arbitrary constructs. One approach to investigating transition zones ascribes values to the area units themselves, by quantifying the spatial turnover among species within the surrounding neighbourhoods of areas on maps. Using data for bumble bee distributions world-wide, I show that quantitative measures of neighbourhood turnover can discover many of the transition zones that are found by classification techniques when applied to the same data. But unlike classification techniques, turnover measures, particularly when used in combination, can show how a transition zone varies along its length, not only in its strength (the proportion of species contributing to the zone) but also in its breadth (the degree of spatial overlap or the degree of coincidence among species replacements across it). For bumble bees at least, these transition zones are also negatively associated with areas that have a combination of both high species richness and high species nestedness ? Williams, P. H. 1998p An annotated checklist of bumble bees with an analysis of patterns of description (Hymenoptera: Apidae, Bombini)C 79-152 (updated at www.nhm.ac.uk/research-curation/projects/bombus)3 Bulletin of The Natural History Museum (Entomology) 67 Bumble bees are among the minority of groups of organisms for which there is some evidence that most species have already been described. Nonetheless, a synoptic revision of the group has been delayed, in part by the difficulties imposed by an unusually high ratio of names to species (averaging more than 11). To explore some of the factors contributing to this phenomenon, historical and geographical trends in the naming of bumble bees are summarised. This shows that most taxa were named by European authors, beginning with the most widespread European species, moving later to not only the more narrowly distributed species and to species from other parts of the world, but also to taxa at progressively lower nomenclatural ranks, particularly within the more widespread European species. Nearly half of all of these names have been published since the last world-wide checklist in 1922. In attempting to bring this up to date, the present checklist adopts broad interpretations of species and recognises a total of 239 recent species (including the social parasites but excluding fossil taxa), with 24 new synonyms and 29 provisional synonyms. The list also includes notes on alternative interpretations of taxonomic status and on nomenclatural problems, drawing attention to those cases where further research is most urgently needed. In particular, suggestions are presented for an application to the International Commission on Zoological Nomenclature to use its Plenary Power in order to conserve current usage of the commonly used names atratus, balteatus, distinguendus, flavifrons, humilis, hyperboreus, mesomelas, mixtus, norvegicus, polaris, pyrenaeus, soroeensis and variabilis.H http://www.nhm.ac.uk/research-curation/projects/bombus/introduction.htmld ? Williams, P. H. 2000/ Are Bombus lucorum and magnus separate species? 15-17 BWARS Newsletter 2000 [None]B http://www.nhm.ac.uk/research-curation/projects/bombus/magnus.html, ? Williams, P. H. 2005e Does specialization explain rarity and decline among British bumblebees? A response to Goulson et al. 33-43 Biological Conservation 122R 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; and (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. doi:10.1016/j.biocon.2004.06.019? Williams, P. H. 2007 The distribution of bumblebee colour patterns world-wide: possible significance for thermoregulation, crypsis, and warning mimicry 87-118) Biological Journal of the Linnean Society 92 Bumblebee colour patterns can be highly variable within species, but are often closely similar among species. This paper takes a quantitative approach to survey bumblebee colour patterns in order to address some of the most basic questions concerning resemblances: do colour-pattern groups exist? are species within colour-pattern groups geographically clumped? are some colour-pattern groups associated with particular kinds of habitat? The results using data for 632 worker patterns from all of the world's bumblebee species show that: (1) there are many repeating colour patterns, forming relatively few groups of species with similar patterns; (2) colour-pattern groups can be recognised using simple rules; (3) species within the 24 largest colour-pattern groups are significantly aggregated in particular areas of the world. Three principal divisions of colour-pattern groups are associated with three likely functions: (1) the darkest bumblebees are associated primarily with the tropics, where a thermoregulatory function is suggested; (2) the palest bumblebees are associated with intermediate northern latitudes, where a cryptic function in drying grasslands is suggested; and (3) the intermediate, strongly banded bumblebees are widespread, but these patterns predominate where the other two groups are less numerous and where banding may have advantages as collective warning signals to predators (Müllerian mimicry). Further studies are needed to test these explanations.$ doi:10.1111/j.1095-8312.2007.00878.x ? j Williams, P. H.
Cameron, S. A. 1993K Bumble bee (Bombus Latr.) records from the Valley of Flowers, Uttar Pradesh 125-127 Bulletin of Entomology 31 [None] k ?
Williams, P.H.
Hernández, L.M. 2000 Distinguishing females of the bumble bees Bombus ruderatus (F.) from Bombus hortorum (L.) in Britain: a preliminary application of quantitative techniques 25B Report to the UK Biodiversity Action Plan Bumblebee Working Group. London The Natural History Museum" This study examined the British bumble bees Bombus ruderatus and B. hortorum to assess which characters might be used to give the best discrimination between females of the two taxa. The approach taken was to apply quantitative methods, in order to provide explicit and accountable comparisons. The first stage of the analysis examined queens to identify which of 30 morphological and pubescence characters might be most promising. The second stage then sought to assess which of the subset of 13 promising characters worked best with a larger sample of the workers, which are more difficult to distinguish. Although these characters showed statistically significant differences between taxa, many of them showed poor discriminatory power. However, a combination of two morphological characters, the breadth of the head and the length of the malar area, was found to distinguish most queens of the two taxa. This combination has not been used before, and for our sample showed an improved discrimination (90% individuals non-overlapping) compared to some of the most popular traditional characters (39% non-overlapping).
1. We recommend that a combination of morphological characters and pubescence characters is needed for separation of B. hortorum from B. ruderatus. No single character is guaranteed to separate these taxa in every case.
2. We recommend that the ratio between the breadth of the head and the length of the malar area provides the best separation for queens of B. hortorum from B. ruderatus.
3. We recommend that further study is required to examine quantitatively variation in character 24 (YHAGII), the longitudinal breadth of the band of yellow hairs on gastral segment II in dorsal view.
4. We recommend that further study is required to examine quantitatively variation among a larger sample of female specimens and variation in characters of males.
5. We recommend that further study is required to examine quantitatively variation within colonies of known parentage for comparison of within-colony, between-colony, and between-locality variation. ? Williams, P. H.
Seddon, A 1993 Mongolian bumble bee survey 6_ Raleigh International Mongolia 92D expedition: June-August 1992 preliminary scientific reports. S. Belbin London [None] ? Williams, P.H.
Thomas, J.C. 2005L A bumblebee new to the New World: Bombus distinguendus (Hymenoptera: Apidae) 158-162 Canadian Entomologist 137 2J Bombus (Subterraneobombus) distinguendus Morawitz, 1869, is recorded from Attu Island, in the western Aleutian archipelago, and is recognised as a species new to both the Alaskan and the New World faunal lists. Colour variation and food plants are described and a key to females of Nearctic Subterraneobombus species is provided. ? Edwards, M.
Williams, P. H. 2004? Where have all the bumblebees gone, and could they ever return? 305-312 British Wildlife 5 15 [None]h ? Cameron, S. A.
Williams, P.H. 2003 Phylogeny of bumble bees in the New World subgenus Fervidobombus (Hymenoptera: Apidae): congruence of molecular and morphological data 552-563% Molecular Phylogenetics and Evolution 28 We present new DNA sequence data (12S, 16S and opsin gene fragments) and morphological characters of the male genitalia for a phylogenetic analysis of the bumble bee subgenus Fervidobombus. There is no significant incongruence between the three molecular data sets, and little incongruence between the DNA and morphology. Simultaneous analysis of all the data partitions resulted in a tree that was entirely congruent with the All-DNA tree. Optimization of the geographic locations of the taxa onto the tree topology using dispersal/vicariance analysis suggests a complex picture of spread and diversification of Fervidobombus from the Old World into the southern New World. There is a phylogenetic component to their spread into tropical rain forest, as the two primary rain forest species (B. transversalis and B. pullatus) comprise a monophyletic clade, along with a third species, B. atratus, which is widely distributed in South America, including lowland subtropical habitats.S? Williams, P. H. 2000@ Some properties of rarity scores used in site quality assessment 73-861 British Journal of Entomology and Natural History 13 Species lists for sites are often compared for rarities using an index of the average or mean national range size of the species ('species quality score' or SQS). This paper describes some properties of SQS that need to be kept in mind when interpreting the results, illustrated using atlas data for bumble bees on a 10 × 10 km grid. Results show that SQSs may be correlated (1) with recorded species richness; and (2) with recording effort. With these data, national SQSs are capable of identifying concentrations of species with narrow national distributions even within northern Britain, so that a separate regional treatment is not always necessary. However, the most important observation is that despite these correlations, the most extreme high and low SQS values can only occur when recorded species richness is low, which when due to low recording effort, could be particularly misleading. Similar measures of range-size rarity from the conservation literature are discussed, as well as other approaches for looking at how available data may be used to find combinations of sites (some of them species-poor but with rare species) that represent a greater diversity of wildlife. D ? 0 5 Plowright, C. M. S.
Plowright, R. C.
Williams, P. H. 1997G Replacement of Bombus muscorum by Bombus pascuorum in northern Britain? 985-990 Canadian Entomologist 129R Records of bumble bees from four collecting trips (1977, 1978, 1988, and 1994) in northern Britain show apparent changes in the incidence of two species. Until 1988, Bombus muscorum (Linnaeus) and Bombus pascuorum (Scopoli) were found in the counties of Cumbria, Durham, and North Yorkshire. Only B. muscorum was found in the Orkney Islands in 1978. In 1994, however, B. pascuorum was found abundantly in all of these areas, and in the Orkney Islands we could find only a single B. muscorum. From these data, B. pascuorum appears to have been replacing B. muscorum in some parts of Britain.c? 1 Vane-Wright, R.I.
Humphries, C.J.
Williams, P. H. 19916 What to protect? - Systematics and the agony of choice 235-254 Biological Conservation 55 Politicians and scientists alike now agree that a priority list of global centres for preservation of biological diversity is required. Diversity has generally been measured only in terms of species richness, or in the form of indices combining richness and abundance. Such measures are considered inadequate for the task in hand. A novel index, based on the information content of cladistic classifications and giving a measure of taxonomic distinctness, is introduced. This taxic diversity measure, when coupled with detailed knowledge of distribution, can be used in modified analyses of the type previously developed as 'critical faunas analysis' or 'network analysis'. Central to all such analyses is the concept of complementarity of floras or faunas. By employing complementarity, step-wise procedures can identify optimally efficient, single-site sequences of priority areas for a group, taking existing reserves into account or not, as required. For practical planning it is concluded that two basic rounds of analysis are required: first, recognition of global priority areas by taxic diversity techniques; secondly within any such area, analysis without taxic weighting (as being developed by Margules and his co-workers) to identify a network of reserves to contain all local taxa and ecosystems. The paper concludes with a brief discussion of some immediate prospects for development of a systematic approach to global conservation evaluation. ? i Williams, P.H. 2004 Genus Bombus Latreille Web page providing a list of bumblebees of India and adjacent countries with common synonyms and a summary of their distributions within the region> An annotated catalogue of the bee species of the Indian Region Gupta, R.K. Jodhpur Jai Narain Vyas University 2004, http://www.geocities.com/beesind2/bombus.htm? ) Williams, P.H.
Araújo, M. B.
Rasmont, P. 2007f Can vulnerability among British bumblebee (Bombus) species be explained by niche position and breadth? 493-505 Biological Conservation 138 Comparison of the two flagship species of British bumblebee conservation (Bombus distinguendus and B. sylvarum) with a widespread, common, and more stable species (B. pascuorum) shows (1) that the two rarer and range-declining species in Britain had narrower (more specialized) climatic niches in western Europe even before their most severe declines, and (2) that the areas where they persist in Britain from 2000 onwards are closer climatically to the centres of their pre-decline west-European climatic niches than the areas from which they have been lost. Although data are available for few bumblebee species at present and further tests are needed, the first result supports earlier suggestions that it is bumblebee species with narrower climatic niches that are most vulnerable to decline. The second result supports the suggestion that it is in areas nearer the edges of their climatic niches where these species are most vulnerable to decline, although this can be ameliorated locally by higher food-plant densities. This is not to say that all patterns of bumblebee decline have been influenced by climatic niche, particularly in North America. Nonetheless, in Britain we find that even without climatic change, an interaction between climatic niche and food-plant reductions from land-use change retains the potential to explain at least some of the broader patterns of which species have declined, where they have declined, and how they have declined. doi:10.1016/j.biocon.2007.06.001 ? Williams, P. 2007# Places to see bumblebees: Dungeness 4-78 Buzzword, Newsletter of the Bumblebee Conservation Trust 4/? j E Williams, P. H.
Cameron, S. A.
Hines, H. M.
Cederberg, B.
Rasmont, P. 2008G A simplified subgeneric classification of the bumblebees (genus Bombus) 1-29
Apidologie 39 A system of subgenera has been widely used for nearly a century to communicate ideas of relationships among bumblebee species. However, with 38 subgenera in recent lists for about 250 species, this system has come to be seen as too complicated. In this paper we suggest four criteria to guide the process of simplifying the subgeneric system, suggesting that ideally subgenera should become: (1) monophyletic; (2) fewer; (3) diagnosable from morphology; and (4) names for important behavioural and ecological groups. Using a new strongly-supported estimate of phylogeny for almost all bumblebee species, we apply these criteria to reduce the system to 15 subgenera, and we assess the consequences. Ten new subgeneric synonyms are recognised. doi: 10.1051/apido:2007052 ?! N Monfared, A.
Talebi, A.A.
Tahmasbi, G.
Williams, P.H.
Ebrahimi, E.
Taghavi, A. 2008b A survey of the localities and food-plants of the bumblebees of Iran (Hymenoptera: Apidae: Bombus) 283-299 Entomologia Generalis 30(2007) 3 Bumblebees are common in the mountains of Iran and are known to be important pollinators of both native and greenhouse plants. Over the last seventy years, 32 bumblebee species (28 true bumblebees and 4 cuckoo bumblebees) have been reported from Iran. However, most previous collections of bumblebees from Iran have concentrated on the Alborz mountains of the north, although bumblebees are also known to inhabit the mountains in the northwest (Sabalan), west (Zagros), and some central parts. Between 2004-2007, bumblebee specimens were collected from all of these mountain regions, and 26 species were recorded. Bombus (Psithyrus) maxillosus Klug 1817 and Bombus (Psithyrus) sylvestris (Lepeletier 1832) are reported from Iran for the first time. A combined list of species with revised interpretations of species and nomenclature is presented, together with localities, elevation ranges, and lists of food plants.1?" Williams, P.H. 2007* Bumble bees - and their decline in Britain 161-176 Aspects of sociality in insects Carreck, N.L.
Johnson, T.W. Ilford" Central Association of Bee-KeepersC http://www.nhm.ac.uk/research-curation/projects/bombus/decline.html reprinted from Williams (1989) ?# Williams, P. H. 2008L Do the parasitic Psithyrus resemble their host bumblebees in colour pattern? 637-649
Apidologie 39 It has been claimed for the parasitic Psithyrus bumblebees that each parasite species resembles closely its particular narrow range of bumblebee host species in colour pattern. The generality of colour-pattern resemblance is assessed by applying quantitative tests at three levels of resolution in the detail of the colour patterns. The results show that at all three levels the parasites and hosts are significantly more likely to share similar colour patterns than would be expected by chance in Europe, but not in North America. Parasites do not appear to be significantly more specialised in host choice in Europe than in North America, although there is more evidence of parasite-host co-speciation in Europe than in North America. Parasite-host resemblance appears most likely to be explained by Müllerian mimicry. For the host-specific Psithyrus, the host species might serve as the most influential model because necessarily the host species must co-occur with the parasite and must be common. doi: 10.1051/apido:2008048 10.1051/apido:2008048?$ B , Williams, P. H.
Tang, Y.
Yao, J.
Cameron, S. 20098 The bumblebees of Sichuan (Hymenoptera: Apidae, Bombini) 101-190 Systematics & Biodiversity 7 2 Bumblebees are important pollinators in mountainous regions. The highland region of Sichuan and Chongqing (together, Sichuan in the former broad sense) includes part of the greatest hotspot of bumblebee diversity world-wide, with half of the species of China and more than a fifth of the world's species. In this paper we present the first review of this diverse but particularly poorly known fauna, drawing on 6705 bees from selective samples made by the authors during field work and from museum collections, together with 1123 literature records (an overlapping set). Among this material we recognise 56 species that have 847 names (including infrasubspecific names). One new species, Bombus (Pyrobombus) wangae, is described from Sichuan and Gansu. B. laesus is found to have a colour pattern in this part of its range that was previously undescribed. Six species are recorded from the Sichuan-Chongqing region for the first time (B. avanus, B. branickii, B. humilis, B. norvegicus, B. tibetanus, B. difficillimus), of which B. avanus is only the second published record worldwide. One species, B. braccatus, is endemic to Sichuan and is confirmed as extant in 2005. We provide diagnoses and keys to species for both sexes. Colour variation is described, distributions within the Sichuan-Chongqing region are mapped, altitudinal and seasonal activity are plotted, and the authors' records of food plants are listed.
Syst. Biodiv. 10.1017/S1477200008002843 ?% Xie, Z.
Williams, P. H.
Tang, Y. 2008d The effect of grazing on bumblebees in the high rangelands of the eastern Tibetan Plateau of Sichuan 695-703 Journal of Insect Conservation 12x Many Tibetans who were formerly nomadic yak herders are settling near towns. However, yak remain a traditional symbol of wealth, which can lead to increased local stocking densities and severe over-grazing. We used belt-transect counts to study the area around Hongyuan in Sichuan. We find that intensive summer grazing significantly reduces the vegetation height and is associated with significant reductions in bumblebee-food-plant abundance and bumblebee diversity. For the significantly reduced bumblebee species, we identify the most-frequently used and preferred bumblebee-food plants. For the food plants, we identify changes in absolute flower availability and changes in bumblebee visits. In particular, reductions in the bumblebees Bombus supremus, B. filchnerae, B. humilis and B. impetuosus are associated with reductions in flowers of the food plants Hedysarum and Saussurea. DOI: 10.1007/s10841-008-9180-3 J. Insect Conserv. 10.1007/s10841-008-9180-38' D?& k Williams, P.
Colla, S.
Xie, Z. 2009X Bumblebee vulnerability: common correlates of winners and losers across three continents 931-940 Conservation Biology 23 4 It is widely agreed that in many parts of the world some bumblebee (Bombus) species have declined, and that this has often been driven by land-use changes that cause reductions in the abundance of food plants. There is much less agreement about how changes in food plants affect some bumblebee species more than others. We sought to identify which species’ characteristics are generally associated with the relative winners and losers by comparing the 3 independent bumblebee faunas from parts of Britain, Canada, and China. Using available survey data, we assessed species characteristics, including competition with congeners, climatic specialization, proximity to climatic range edge, food specialization, phenology, body size, and range size. Results of our meta-analysis of correlations showed support for the hypotheses that decline susceptibility is generally greater for species that have greater climatic specialization, for species in areas where they occur closest to the edges of their climatic ranges, and for species that have queens that become active later in the year. The latter characteristic may render a species at a particular disadvantage when they have long colony cycles if there are losses of food plants in mid to late colony development. 10.1017/nc 0?' m Williams, P. H.
Osborne, J. L. 20093 Bumblebee vulnerability and conservation world-wide 367-387
Apidologie 40^ We review evidence from around the world for bumblebee declines and review management
to mitigate threats. We find that there is evidence that some bumblebee species are declining in Europe,
North America, and Asia. People believe that land-use changes may be having a negative effect through
reductions in food plants in many parts of the world, but that other factors such as pathogens may be
having a stronger effect for a few species in some regions (especially for Bombus s. str. in North America).
Evidence so far is that greater susceptibility to land-use change is associated world-wide with small climatic
ranges, range edges, and late-starting colony-development cycles. More evidence is needed on the roles of
pollen specialization, nest sites, hibernation sites, and pesticides. It is still too early to assess the success
of schemes aimed at improving forage in agricultural and conservation areas. However, schemes aimed at
raising public awareness have been very successful. Until proven safe, we recommend that live bumblebees
should not be moved across continents or oceans for commercial pollination. 10.1051/ap ido/2009025 S1477200008002843 ?( ( Williams, P. H.
An, J.
Huang, J.
Yao, J. 2010O BBCI: A new initiative to document Chinese bumble bees for pollination research 221-222 Journal of Apicultural Research 49 2 { Ǩ?* Packer, L.
Sheffield, C. S.
Gibbs, J.
de Silva, N.
Best, L. R.
Ascher, J.
Ayala, R.
Martins, D.
Roberts, S. V P. M.
Tadauchi, O.
Kuhlmann, M.
Willliams, P. H.
Eardley, C.
Droege, S.
Levchenko, T. V. 2009L The campaign to barcode the bees of the world: progress, problems, prognosis 178-1809 Memorias: VI Congresso Mesoamericano Sobre Abejas Nativas Yurrita, C. L.$ Centro de Estudios Conservacionistas& Universidad de San Carlos de Guatemala PK mb:I/* * refs.FRM 0 B < ! // ! H PRIMARYyearIndex
6 By P /) id reference_type text_styles author year title
pages secondary_title volume
number number_of_volumes secondary_author place_published
publisher subsidiary_author edition keywords
type_of_work date 2) abstract label url tertiary_title tertiary_author notes
isbn custom_1 custom_2
custom_3 custom_4 alternate_title accession_number call_number short_title custom_5 custom_6 section original_publication H) reprint_edition reviewed_item author_address image caption custom_7
electronic_resource_number link_to_pdf translated_author
translated_title name_of_database database_provider research_notes language access_date last_modified_date
! ! H ! H ! H ! ( H !
3 H ! > H ! I H !
T H ! _ H ! j H ! u H !
H ! H ! H ! H !
H ! H ! H ! H ! H ! H ! H ! H !
H ! H ! H !
% H ! 0 H ! ; H ! F H ! Q H ! \ H ! g H ! r H ! } H ! H ! H ! H ! H ! H ! H ! H !
H ! H ! H !
H ! H ! H ! H ! " H ! - H ! 8 H ! idreference_typetext_stylesauthoryeartitlepagessecondary_titlevolumenumbernumber_of_volumessecondary_authorplace_publishedpublishersubsidiary_authoreditionkeywordstype_of_workdateabstractlabelurltertiary_titletertiary_authornotesisbncustom_1custom_2custom_3custom_4alternate_titleaccession_numbercall_numbershort_titlecustom_5custom_6sectionoriginal_publicationreprint_editionreviewed_itemauthor_addressimagecaptioncustom_7electronic_resource_numberlink_to_pdftranslated_authortranslated_titlename_of_databasedatabase_providerresearch_noteslanguageaccess_datelast_modified_date PK {< refs.MYDPK mb:I/* * * refs.FRMPK l (