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.

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.

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.

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.

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.

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.

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.

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.

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 .

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.

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.

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.

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.

CLIMATE CHANGE. Climate change impacts on bumblebees converge across continents.

For many species, geographical ranges are expanding toward the poles in response to climate change, while remaining stable along range edges nearest the equator. Using long-term observations across Europe and North America over 110 years, we tested for climate change-related range shifts in bumblebee species across the full extents of their latitudinal and thermal limits and movements along elevation gradients. We found cross-continentally consistent trends in failures to track warming through time at species' northern range limits, range losses from southern range limits, and shifts to higher elevations among southern species. These effects are independent of changing land uses or pesticide applications and underscore the need to test for climate impacts at both leading and trailing latitudinal and thermal limits for species.

Revision of the Neotropical subgenera Coelioxys (Platycoelioxys) Mitchell and C. (Rhinocoelioxys) Mitchell (Hymenoptera; Megachilidae) with the description of one new species.

Two Neotropical subgenera of Coelioxys Latreille are revised. The monotypic C. (Platycoelioxys) Mitchell and C. (Rhinocoelioxys) Mitchell has seven valid species; six of them (C. agilis Smith, C. barbata Schwarz & Michener, C. clypearis Friese, C. nasidens Friese, C. paraguayensis Schrottky and C. zapoteca Cresson) previously described, and one, C. platygnatha Rocha-Filho & Packer n. sp. is new from Amazonas State, Brazil. Coelioxys nasidens, previously considered a junior synonym of C. clypeata Smith, is resurrected. Coelioxys crassiceps Friese and C. excisa Friese are synonymized under C. agilis, and C. rostrata Friese is synonymized under C. paraguayensis. Nine names (C. clypeata, C. leucochrysea Cockerell, C. angustivalva Holmberg, C. doelloi Holmberg, C. blabera Holmberg, C. mesopotamica Holmberg, C. bilobata Friese, C. bilobata schenki Friese and C. bullaticeps Friese) are synonymized with C. zapoteca. In the latter species the shape of the apical margin of the clypeus of the female varies widely and, though intermediates occur among all forms, the DNA barcode sequences for the different forms are very similar. One species formerly considered as incertae sedis, C. clypearis, and one belonging to the subgenus C. (Cyrtocoelioxys) Mitchell, C. barbata, are both assigned to the subgenus C. (Rhinocoelioxys). Keys for both sexes of C. (Rhinocoelioxys), distribution maps, host and floral records, redescriptions of species and the description of a new species are provided. In the subgenus C. (Platycoelioxys), only one species, C. alatiformis Friese, is recognized with two junior synonyms: C. crassiceps Friese syn. nov. and C. spatuliventer Cockerell.

A new species of Samba s. str. (Hymenoptera: Melittidae) from the Turkana Basin, Kenya with observations on the function of the metatibial spur in females.

The third known species of the Afrotropical subgenus Samba s. str. is described based upon eight females. This is the northernmost and also the most arid habitat record for the genus. Images are provided of the habitus and diagnostic features for the species as well as the subgenus and some notes on floral hosts and habitat are provided. The species is added to a morphology-based phylogeny for the genus and results of barcoding of some species of the genus are presented. Some unusual morphological features of the subgenus are discussed, in particular, the function of the remarkable metatibial spurs of the female is recorded as assisting with removal of pollen from a floral host.

'Bee hotels' as tools for native pollinator conservation: a premature verdict?

Society is increasingly concerned with declining wild bee populations. Although most bees nest in the ground, considerable effort has centered on installing 'bee hotels'--also known as nest boxes or trap nests--which artificially aggregate nest sites of above ground nesting bees. Campaigns to 'save the bees' often promote these devices despite the absence of data indicating they have a positive effect. From a survey of almost 600 bee hotels set up over a period of three years in Toronto, Canada, introduced bees nested at 32.9% of sites and represented 24.6% of more than 27,000 total bees and wasps recorded (47.1% of all bees recorded). Native bees were parasitized more than introduced bees and females of introduced bee species provisioned nests with significantly more female larva each year. Native wasps were significantly more abundant than both native and introduced bees and occupied almost 3/4 of all bee hotels each year; further, introduced wasps were the only group to significantly increase in relative abundance year over year. More research is needed to elucidate the potential pitfalls and benefits of using bee hotels in the conservation and population dynamics of wild native bees.

Two new species of Geodiscelis Michener & Rozen (Hymenoptera: Apoidea: Colletidae) with a phylogenetic analysis and subgeneric classification of the genus.

Two new species of the genus Geodiscelis are described: Geodiscelis nazcalinea Packer & Dumesh, sp. nov. from Peru (the first record of the genus from that country) and G. phisquiri Packer & Dumesh, sp. nov. from northern Chile. The new species are most closely related to G. longiceps, but differ primarily in having somewhat less elongate heads and in details of the male terminalia. A key to the five known species of the genus is provided as are the results of a phylogenetic analysis based upon 68 characters, and the genus is formally divided into three subgenera: Geodiscelis s. str. Michener and Rozen, Geodiscelis (Nazcoediscelis) Packer and Dumesh, subgenus nov. and Geodiscelis (Thaumoediscelis) Packer and Dumesh, subgenus nov. The two new species described herein belong to subgenus Geodiscelis (Nazcoediscelis). Figures of the most important characters are provided. Tiquilia sp. (Boraginaceae) is the probable floral host of both new species and it is suggested that all species are ground-nesters. Sexual dimorphism in an unusual character is recorded for G. thaumaskelos Packer.