The evolutionary history of the cellophane bee genus Colletes Latreille (Hymenoptera: Colletidae): Molecular phylogeny, biogeography and implications for a global infrageneric classification.

Colletes Latreille (Hymenoptera: Colletidae) is a diverse genus with 518 valid species distributed in all biogeographic realms, except Australasia and Antarctica. Here we provide a comprehensive dated phylogeny for Colletes based on Bayesian and maximum likelihood-based analyses of DNA sequence data of six loci: 28S rDNA, cytochrome c oxidase subunit 1, elongation factor-1α copy F2, long-wavelength rhodopsin, RNA polymerase II and wingless. In total, our multilocus matrix consists of 4824 aligned base pairs for 143 species, including 112 Colletes species plus 31 outgroups (one stenotritid and a diverse array of colletids representing all subfamilies). Overall, analyses of each of the six single-locus datasets resulted in poorly resolved consensus trees with conflicting phylogenetic signal. However, our analyses of the multilocus matrix provided strong support for the monophyly of Colletes and show that it can be subdivided into five major clades. The implications of our phylogenetic results for future attempts at infrageneric classification for the Colletes of the world are discussed. We propose species groups for the Neotropical species of Colletes, the only major biogeographic realm for which no species groups have been proposed to date. Our dating analysis indicated that Colletes diverged from its sister taxon, Hemicotelles Toro and Cabezas, in the early Oligocene and that its extant lineages began diversifying only in the late Oligocene. According to our biogeographic reconstruction, Colletes originated in the Neotropics (most likely within South America) and then spread to the Nearctic very early in its evolutionary history. Geodispersal to the Old World occurred soon after colonization of the Northern Hemisphere. Lastly, the historical biogeography of Colletes is analyzed in light of available geological and palaeoenvironmental data.

Phylogeny and biogeography of the cleptoparasitic bee genus Epeolus (Hymenoptera: Apidae) and cophylogenetic analysis with its host bee genus Colletes (Hymenoptera: Colletidae).

The bee genus Epeolus Latreille (Hymenoptera: Apidae) consists of 109 species, which are known to be exclusively cleptoparasites of polyester (or cellophane) bees of the genus Colletes Latreille (Hymenoptera: Colletidae). Both genera have a nearly cosmopolitan distribution and are represented on all continents except Antarctica and Australia. We present the most comprehensive phylogeny for Epeolus to date, based on combined molecular and morphological data. In total, 59 ingroup taxa (species of Epeolus) and 7 outgroup taxa (other Epeolini) were scored for 99 morphological characters, and sequence data were obtained for seven genes (one mitochondrial and six nuclear, 5399 bp in total). Epeolus was found to be monophyletic, with a crown age estimated to be 25.0-13.4 Ma (95% HPD) and its origins traced to the Nearctic region. Epeolus was found to contain six major clades, five of which were well supported. The evolutionary history of Epeolus is explored in the context of earth history events and the evolutionary history of its host genus Colletes, for which a molecular phylogeny was constructed based on the same seven genes. A comparison of Epeolus and Colletes phylogenies limited to taxa for which there is evidence of an association suggests there was some cospeciation. However, more cladogenetic events in Epeolus were linked to instances of dispersal/vicariance. It is not yet clear the extent to which allopatric speciation contributed to diversification in Colletes, but the genus' success in having colonized and diversified across much of the globe made it possible for Epeolus to do the same.

Forecasting pollination declines through DNA barcoding: the potential contributions of macroecological and macroevolutionary scales of inquiry.

While pollinators are widely acknowledged as important contributors to seed production in plant communities, we do not yet have a good understanding of the importance of pollinator specialists for this ecosystem service. Determination of the prevalence of pollinator specialists is often hindered by the occurrence of cryptic species and the limitations of observational data on pollinator visitation rates, two areas where DNA barcoding of pollinators and pollen can be useful. Further, the demonstrated adequacy of pollen DNA barcoding from historical records offers opportunities to observe the effects of pollinator loss over longer timescales, and phylogenetic approaches can elucidate the historical rates of extinction of specialist lineages. In this Viewpoint article, we review how advances in DNA barcoding and metabarcoding of plants and pollinators have brought important developments to our understanding of specialization in plant-pollinator interactions. We then put forth several lines of inquiry that we feel are especially promising for providing insight on changes in plant-pollinator interactions over space and time. Obtaining estimates of the effects of reductions in specialists will contribute to forecasting the loss of ecosystem services that will accompany the erosion of plant and pollinator diversity.

DNA barcoding the bees (Hymenoptera: Apoidea) of Chile: species discovery in a reasonably well known bee fauna with the description of a new species of Lonchopria (Colletidae).

We compare the diversity of bees in the Chilean fauna as understood from traditional taxonomy-based catalogues with that currently known from DNA barcodes using the BIN system informed by ongoing morphology-based taxonomic research. While DNA barcode surveys of the Chilean bee fauna remain incomplete, it is clear that new species can readily be distinguished using this method and that morphological differentiation of distinct barcode clusters is sometimes very easy. We assess the situation in two genera in some detail. In Lonchopria Vachal one "species" is readily separable into two BINs that are easily differentiated based upon male mandibular and genitalic morphology (characters generally used in this group) as well as female hair patterns. Consequently, we describe Lonchopria (Lonchopria) heberti Packer and Ruz, new species. For Liphanthus Reed, a large number of new species has been detected using DNA barcoding and considerable additional traditional morphological work will be required to describe them. When we add the number of BINs (whether identified to named species or not) to the number of Chilean bee species that we know have not been barcoded (both described and new species under study in our laboratories) we conclude that the bee fauna of Chile is substantially greater than the 436 species currently known. Spanish language abstract available as supplementary data 1 .

DNA barcoding as a useful tool in the systematic study of wild bees of the tribe Augochlorini (Hymenoptera: Halictidae).

Special care is needed in the delimitation and identification of halictid bee species, which are renowned for being morphologically monotonous. Corynura Spinola and Halictillus Moure (Halictidae: Augochlorini) contain species that are key elements in southern South American ecosystems. These bees are very difficult to identify due to close morphological similarity among species and high sexual dimorphism. We analyzed 170 barcode-compliant COI sequences from 19 species. DNA barcodes were useful to confirm gender associations and to detect two new cryptic species. Interspecific distances were significantly higher than those reported for other bees. Maximum intraspecific divergence was less than 1% in 14 species. Barcode index numbers (BINs) were useful to identify putative species that need further study. More than one BIN was assigned to five species. The name Corynura patagonica (Cockerell) probably refers to two cryptic species. The results suggest that Corynura and Halictillus species can be identified using DNA barcodes. The sequences of the species included in this study can be used as a reference to assess the identification of unknown specimens. This study provides additional support for the use of DNA barcodes in bee taxonomy and the identification of specimens, which is particularly relevant in insects of ecological importance such as pollinators.

Phylogenetic position of the bee genera Ancyla and Tarsalia (Hymenoptera: Apidae): a remarkable base compositional bias and an early Paleogene geodispersal from North America to the Old World.

We address the phylogenetic position of the bee genera Tarsalia and Ancyla (currently forming the tribe Ancylaini) on the basis of morphological, molecular and combined data. We assembled a matrix of 309 morphological characters and 5246 aligned nucleotide positions from six nuclear genes (28S, EF-1a, wingless, POL2, LW-Rhodopsin, NAK). In addition to both constituent genera of Ancylaini, we include all three subtribes of the Eucerini as well as a large number of other tribes from the "eucerine line". The morphological data suggest Ancyla to be sister to Tarsalia+Eucerini and analyses of the entire molecular dataset suggest Tarsalia to be sister to Ancyla+Eucerini. However, analyses of the combined dataset suggests the Ancylaini to be monophyletic. We address possible bias within the molecular data and show that the base composition of two markers (EF-1a and NAK) is significantly heterogeneous among taxa and that this heterogeneity is strong enough to overcome the phylogenetic signal from the other markers. Analyses of a molecular matrix where the heterogeneous partitions have been RY-recoded yield trees that are better resolved and have higher nodal support values than those recovered in analyses of the non-recoded matrix, and strongly suggest the Ancylaini to be a monophyletic sister group to the Eucerini. A dated phylogeny and ancestral range reconstructions suggest that the common ancestor of the Ancylaini reached the Old World from the New World most probably via the Thulean Land Bridge in a time window between 69 and 47 mya, a period that includes the Early Eocene Climatic Optimum. No further exchanges between the New World and the Old World are implied by our data until the period between 22 mya and 13.9 mya. These more recent faunal exchanges probably involved geodispersal over the Bering Land Bridge by less thermophilic lineages.

Three New Species of Microsphecodes s.str. (Hymenoptera: Halictidae) with an updated key to species of the subgenus.

We diagnose, describe and illustrate three new species of Microsphecodes Eickwort and Stage, all from the nominate subgenus: M. peckorum Graham and Packer n. sp. from Venezuela, M. fernandoi Graham and Packer n. sp. from French Guiana, and M. septentrionalis Graham and Packer n. sp. from Guatemala. The latter extends the known range of the subgenus further north by approximately 500 km. We provide an updated, illustrated, key to the species of Microsphecodes s.str.

Three new species of Neofidelia (Hymenoptera: Apoidea: Megachilidae) from Northern Chile.

Neofidelia apacheta Dumesh and Packer, sp. nov., N. camanchaca Dumesh and Packer, sp. nov., and N. submersa Dumesh and Packer, sp. nov. are described from northern Chile. Neofidelia apacheta is known from large numbers of specimens and localities at moderately high elevation (1600-3200m). Neofidelia camanchaca is only known from the ho-lotype male. Neofidelia submersa is known from the holotype male and a pair of paratypes in poor condition, all from a coastal fog oasis. Comparisons are made with the two previously described species of the genus. The collection of N. apa-cheta from so far north in Chile suggests that Neofidelia is likely to be found in southern Peru. A key for the five species of the genus is provided.

Revision and reclassification of Lasioglossum (Evylaeus), L. (Hemihalictus) and L. (Sphecodogastra) in eastern North America (Hymenoptera: Apoidea: Halictidae).

The black species of weak-veined Lasioglossum (or Hemihalictus series) in eastern North America are revised to clarify their taxonomy and nomenclature and to facilitate identification. A subgeneric classification based upon available phylogenetic data is applied. Lasioglossum (Sphecodogastra) is applied more broadly than any previous usage to include many species typically classified as L. (Evylaeus). The subgenus L. (Evylaeus) is retained but applied narrowly in agreement with phylogenetic results. Lasioglossum (Hemihalictus) has historically been considered monotypic but is here applied to many species of L. (Dialictus) sensu lato (equivalent to the carinaless L. (Evylaeus) of some authors). Usage of L. (Dialictus) is restricted primarily to species with metallic integument. Additional subgeneric synonymies for extralimital taxa are formalized and discussed. Descriptions are provided for each species with a synonymic list, diagnosis, and notes on taxonomy and biology. The recently revised Onagraceae-specialist species of L. (Sphecodogastra) are given abbreviated treatments. Notes on available DNA barcode data are given, with diagnostic characters supplied for closely related spe-cies. One new species is described: L. (Sphecodogastra) seillean Gibbs and Packer and the males of L. fedorense (Crawford) and L. pectinatum (Robertson) are described for the first time. The following three new synonymies are proposed: Lasioglossum (Hemihalictus) sopinci (Crawford), senior subjective synonym of Evylaeus bradleyi Mitchell; Lasioglosum (Hemihalictus) macoupinense (Robertson), senior subjective synonym of Halictus divergens Lovell; and Lasioglossum (Hemihalictus) inconditum (Cockerell), senior subjective synonym of Halictus tracyi Cockerell. Lasioglossum inconditum is here considered to be distinct from the Palaearctic species L. rufitarse (Zetterstedt). A lectotype is designat-ed for Halictus quebecensis Crawford. We present the first record of L. lustrans (Cockerell) and L. swenki (Crawford) in Canada and the first record of L. lusorium (Cresson) east of the Mississippi River. Updated keys to species are provided for the fauna of eastern North America.

The evolutionary history of bees in time and space.

Bees are the most significant pollinators of flowering plants. This partnership began ca. 120 million years ago, but the uncertainty of how and when bees spread across the planet has greatly obscured investigations of this key mutualism. We present a novel analysis of bee biogeography using extensive new genomic and fossil data to demonstrate that bees originated in Western Gondwana (Africa and South America). Bees likely originated in the Early Cretaceous, shortly before the breakup of Western Gondwana, and the early evolution of any major bee lineage is associated with either the South American or African land masses. Subsequently, bees colonized northern continents via a complex history of vicariance and dispersal. The notable early absences from large landmasses, particularly in Australia and India, have important implications for understanding the assembly of local floras and diverse modes of pollination. How bees spread around the world from their hypothesized Southern Hemisphere origin parallels the histories of numerous flowering plant clades, providing an essential step to studying the evolution of angiosperm pollination syndromes in space and time.

Dual origins of social parasitism in North American Dialictus (Hymenoptera: Halictidae) confirmed using a phylogenetic approach.

The bee subgenus Dialictus (Halictidae: Lasioglossum) displays a large array of behaviours including solitary behaviour, eusociality, and social parasitism. Socially parasitic Dialictus share a suite of morphological traits; these could result from shared ancestry, but given their functional significance, could also have resulted from adaptive convergence. A combined morphological and molecular phylogenetic approach was used to test for monophyly of North American socially parasitic Dialictus. Two data sets were used in the phylogenetic analyses. First, short mitochondrial DNA sequences from previous taxonomic studies of North American Dialictus, including six social parasites, were used because of the broad taxon sampling they provide. These data were analysed in combination with a set of 40 morphological characters, including a large proportion of characters associated with social parasitism. Phylogenetic analysis of the combined DNA barcode and morphology data set resolves two distinct lineages of social parasite. The second data set was based on three genes (cytochrome c oxidase subunit 1, elongation factor 1α, and long-wavelength rhodopsin), but with sparser taxon sampling, including one representative from each putative social parasite-lineage. This also supported dual origins of social parasitism among North American Dialictus. The evolution of social parasitism is discussed. © The Willi Hennig Society 2011.

Fifteen new species of Chilicola (Oroediscelis) (Hymenoptera: Colletidae: Xeromelissinae) with illustrated keys to the males and females of the subgenus.

Fifteen new species of Chilicola subgenus Oroediscelis are described and illustrated. The following 11 species are from Argentina recording a remarkable diversity and the first records of the subgenus in the country: Chilicola capillitas Packer and Dumesh, new species, C. abrebotellas, Packer and Dumesh, new species; C. goloboffi Packer and Dumesh, new species; C. fritzi, Packer and Dumesh, new species; C. rozeni Packer and Dumesh, new species; C. roigi Packer and Dumesh, new species; C. carpenteri Packer Dumesh, new species; C. calchaqui Packer and Dumesh, new species; C. jaguense Packer and Dumesh, new species; C. cuyense Dumesh Packer, new species; and C. huarpe Packer and Dumesh, new species. Chilicola pustulata Packer and Dumesh, new species, and Chilicola luna Dumesh, and Packer new species, are described from Peru; C. boharti Packer and Dumesh, new species, and C. ashei Dumesh and Packer, new species, are described from Bolivia, where C. carpenteri also occurs. An unusual feature of the male antennae is described: long setae on F2 and F3 (less commonly F4) that are appressed to, and occasionally longer than, the succeeding flagellomere. Additional records for previously described species are listed and a revised key to males is provided as well as a provisional complete key for all females.

Fifteen new species of Liphanthus Reed (Hymenoptera: Andrenidae) with two submarginal cells.

Hitherto, the panurgine genus Liphanthus Reed 1894 has been thought to have only a single species with two, as opposed to three, submarginal cells. Here we describe an additional fifteen species with two submarginal cells. These new species are: L. jenamro Mir Sharifi Packer, L. sapos Mir Sharifi Packer, L. domeykoi Packer, L. discolor Mir Sharifi Packer, L. centralis Mir Sharifi Packer, L. molavi Mir Sharifi Packer (all of the above are from Chile), L. abotorabi Mir Sharifi Packer, L. cochabambensis Mir Sharifi Packer (both from Bolivia), L. fritzi Mir Sharifi Packer, L. amblayensis Mir Sharifi Packer (both from Argentina), L. ancashensis Mir Sharifi Packer (from Peru), L. tregualemensis Packer (from Chile), L. yrigoyeni Packer, L. sparsipunctus Packer (both from Argentina) and L. aliavenus Packer (from Chile). Only L. tregualemensis readily fits within any of the previously described subgenera-Liphanthus (Leptophanthus) Ruz and Toro 1983. Liphanthus aliavenus is known from two specimens, one with three and one with two submarginal cells whereas L. molavi has one individual with two submarginal cells on one forewing and three on the other while all other specimens have two submarginal cells on each forewing. We verified that none of these new species are merely two submarginal celled variants of species with three submarginal cells (such intraspecific variation arises also in some other bees) by i) comparing each of the new species with all keys, figures and descriptions of all Liphanthus species, ii) comparisons with holotypes and/or paratypes of most of the described species and iii) surveys of the specimens of undescribed species with three submarginal cells in our collection. None of the new species seem closely related to L. (Neoliphanthis) bicellularis Ruz and Toro 1983, the only previously described Liphanthus species with two submarginal cells. It is the second submarginal crossvein that is lost in all species except L. aliavenus in which the first submarginal cross vein is lost. DNA barcode data are presented for some of the species. Some interesting morphological features associated with the penis valves are described and discussed. The genus is recorded from Bolivia for the first time.

A new species of Lepidotrigona (Hymenoptera: Apidae) from Thailand with the description of males of L. flavibasis and L. doipaensis and comments on asymmetrical genitalia in bees.

We describe Lepidotrigona satun Attasopa and Bänziger new species from southern Thailand based upon associated males and females (workers). The new species is a member of the L. ventralis species group, which is otherwise represented in Thailand only by L. flavibasis and L. doipaensis. We also describe the males of the latter two species, associated with nests from close to their type localities in northern Thailand. Lepidotrigona doipaensis Schwarz and L. flavibasis (Cockerell) had previously often been misidentified as L. ventralis (Smith), a species confirmed only from Borneo. Based upon differences in male morphology, especially of the metasomal sterna, we conclude that the male described from Vietnam by Sakagami (1975) as belonging to L. flavibasis represents an undescribed species. Our findings support previous taxonomic studies that highlight the importance of including males in the differentiation of closely related species of meliponines and their association with workers. The three species whose males we describe have asymmetric penis valves with the asymmetry differentially developed among the three. We compare this genitalic asymmetry with that known from a different apid genus, Tarsalia.

Pollination and other ecosystem services produced by mobile organisms: a conceptual framework for the effects of land-use change.

Many ecosystem services are delivered by organisms that depend on habitats that are segregated spatially or temporally from the location where services are provided. Management of mobile organisms contributing to ecosystem services requires consideration not only of the local scale where services are delivered, but also the distribution of resources at the landscape scale, and the foraging ranges and dispersal movements of the mobile agents. We develop a conceptual model for exploring how one such mobile-agent-based ecosystem service (MABES), pollination, is affected by land-use change, and then generalize the model to other MABES. The model includes interactions and feedbacks among policies affecting land use, market forces and the biology of the organisms involved. Animal-mediated pollination contributes to the production of goods of value to humans such as crops; it also bolsters reproduction of wild plants on which other services or service-providing organisms depend. About one-third of crop production depends on animal pollinators, while 60-90% of plant species require an animal pollinator. The sensitivity of mobile organisms to ecological factors that operate across spatial scales makes the services provided by a given community of mobile agents highly contextual. Services vary, depending on the spatial and temporal distribution of resources surrounding the site, and on biotic interactions occurring locally, such as competition among pollinators for resources, and among plants for pollinators. The value of the resulting goods or services may feed back via market-based forces to influence land-use policies, which in turn influence land management practices that alter local habitat conditions and landscape structure. Developing conceptual models for MABES aids in identifying knowledge gaps, determining research priorities, and targeting interventions that can be applied in an adaptive management context.

Phylogenetic analysis of the corbiculate Apinae based on morphology of the sting apparatus (Hymenoptera: Apidae).

This study aimed to test the various competing hypotheses regarding the relationships among the four tribes of corbiculate apine bees (Euglossini "orchid bees", Bombini "bumble bees", Meliponini "stingless bees", and Apini "honey bees") with a completely new set of previously unstudied morphological characters derived from the sting apparatus. The result was one most parsimonious tree of 49 steps, CI = 89, RI = 93 that is perfectly congruent with most studies based on morphological and combined morphological/molecular data, i.e., Euglossini + (Bombini + (Meliponini + Apini)), supporting a well accepted scenario of social evolution for these bees. This data matrix was then combined with other published matrices for this group in order to perform simultaneous analyses. The problem of how to best combine the multiple matrices that did not use the same exemplars was investigated.

Queens and Workers Contribute Differently to Adaptive Evolution in Bumble Bees and Honey Bees.

Eusociality represents a major transition in evolution and is typified by cooperative brood care and reproductive division of labor between generations. In bees, this division of labor allows queens and workers to phenotypically specialize. Worker traits associated with helping are thought to be crucial to the fitness of a eusocial lineage, and recent studies of honey bees (genus Apis) have found that adaptively evolving genes often have worker-biased expression patterns. It is unclear however if worker-biased genes are disproportionately acted on by strong positive selection in all eusocial insects. We undertook a comparative population genomics study of bumble bees (Bombus) and honey bees to quantify natural selection on queen- and worker-biased genes across two levels of social complexity. Despite sharing a common eusocial ancestor, genes, and gene groups with the highest levels of positive selection were often unique within each genus, indicating that life history and the environment, but not sociality per se, drives patterns of adaptive molecular evolution. We uncovered differences in the contribution of queen- and worker-biased genes to adaptive evolution in bumble bees versus honey bees. Unlike honey bees, where worker-biased genes are enriched for signs of adaptive evolution, genes experiencing positive selection in bumble bees were predominately expressed by reproductive foundresses during the initial solitary-founding stage of colonies. Our study suggests that solitary founding is a major selective pressure and that the loss of queen totipotency may cause a change in the architecture of selective pressures upon the social insect genome.

The Bees among Us: Modelling Occupancy of Solitary Bees.

Occupancy modelling has received increasing attention as a tool for differentiating between true absence and non-detection in biodiversity data. This is thought to be particularly useful when a species of interest is spread out over a large area and sampling is constrained. We used occupancy modelling to estimate the probability of three phylogenetically independent pairs of native-introduced species [Megachile campanulae (Robertson)-Megachile rotundata (Fab.), Megachile pugnata Say-Megachile centuncularis (L.), Osmia pumila Cresson-Osmia caerulescens (L.)] (Apoidea: Megachilidae) being present when repeated sampling did not always find them. Our study occurred along a gradient of urbanization and used nest boxes (bee hotels) set up over three consecutive years. Occupancy modelling discovered different patterns to those obtained by species detection and abundance-based data alone. For example, it predicted that the species that was ranked 4th in terms of detection actually had the greatest occupancy among all six species. The native M. pugnata had decreased occupancy with increasing building footprint and a similar but not significant pattern was found for the native O. pumila. Two introduced bees (M. rotundata and M. centuncularis), and one native (M. campanulae) had modelled occupancy values that increased with increasing urbanization. Occupancy probability differed among urban green space types for three of six bee species, with values for two native species (M. campanulae and O. pumila) being highest in home gardens and that for the exotic O. caerulescens being highest in community gardens. The combination of occupancy modelling with analysis of habitat variables as an augmentation to detection and abundance-based sampling is suggested to be the best way to ensure that urban habitat management results in the desired outcomes.

Long-Chain Omega-3 Polyunsaturated Fatty Acids Have Developmental Effects on the Crop Pest, the Cabbage White Butterfly Pieris rapae.

Nutritional enhancement of crops using genetic engineering can potentially affect herbivorous pests. Recently, oilseed crops have been genetically engineered to produce the long-chain omega-3 polyunsaturated fatty acids, eicosapentaenoic acid (EPA) and docosahexaenoic acid (DHA) at levels similar to that found in fish oil; to provide a more sustainable source of these compounds than is currently available from wild fish capture. We examined some of the growth and development impacts of adding EPA and DHA to an artificial diet of Pieris rapae, a common pest of Brassicaceae plants. We replaced 1% canola oil with EPA: DHA (11:7 ratio) in larval diets, and examined morphological traits and growth of larvae and ensuing adults across 5 dietary treatments. Diets containing increasing amounts of EPA and DHA did not affect developmental phenology, larval or pupal weight, food consumption, nor larval mortality. However, the addition of EPA and DHA in larval diets resulted in progressively heavier adults (F 4, 108 = 6.78; p = 0.011), with smaller wings (p < 0.05) and a higher frequency of wing deformities (R = 0.988; p = 0.001). We conclude that the presence of EPA and DHA in diets of larval P. rapae may alter adult mass and wing morphology; therefore, further research on the environmental impacts of EPA and DHA production on terrestrial biota is advisable.