Thibaut De Meulemeester

An overview of the Bombus terrestris (L. 1758) subspecies (Hymenoptera: Apidae)

Abstract Bombus terrestris is one of the most abundant bumblebee species in the West-Palaearctic. Its widespread domestication results in the movement of many colonies. The aim of this paper is to describe the 9 subspecies currently recognised and to list some of their most obvious characteristics. Bombus terrestris is not declining anywhere, on the contrary, its synanthropic spread through domestication may be expected. However, its autumn and winter populations rely on a restricted choice of flowers so they may be threatened locally by scrub clearance and the destruction of their favourite autumnal flowers.

Geometric morphometric analysis of a new Miocene bumble bee from the Randeck Maar of southwestern Germany (Hymenoptera: Apidae)

The first fossil bumble bee (Apinae: Bombini) from the Miocene Randeck Maar of southwestern Germany is described and illustrated. The specimen is subjected to a geometric morphometric analysis along with a diversity of other bumble bee species representing most major extant lineages, and particularly the subgenus Bombus s.s. The morphometric analysis supports the placement of the Randeck Maar species within Bombus s.s., as a species distinct from all others in the subgenus. It shows that extant subgenera of bumblebees were already derived in the early/middle Miocene. The Randeck Maar fossil is formally described as Bombus (Bombus) randeckensis Wappler & Engel sp. n..

New fossil evidence of the early diversification of bees: Paleohabropoda oudardi from the French Paleocene (Hymenoptera, Apidae, Anthophorini)

Phylogenetic relationships among and within the major groups of bees (Apoidea Apiformes) were recently reconsidered using extensive molecular and morphological datasets. The next step in the study of bee evolution will consist in estimating the antiquity of the nodes within the inferred topologies. We describe here the third oldest bee fossil, Paleohabropoda oudardi gen. n. sp. N. (Apidae, Apinae, Anthophorini) from the Paleocene of Menat (France, Puy‐de‐Dôme; 60 Myr). Phylogenetic analysis of 17 morphological characters and morphometric analysis of the wing shape were used to recover, respectively, its taxonomic position and morphological affinities. Our results indicate that Paleohabropoda oudardi gen. n. sp. n. clearly belongs to the Anthophorini. Paleohabropoda is therefore the oldest fossil that can be confidently date and place to an extant tribe. Its wing shape is surprisingly close to the extant genus Habropoda. The discovery of Paleohabropoda oudardi gen. n., sp. n. brings further evidence for the Cretaceous diversification of major lineages of bees and for strong constancy of wing shape within the Anthophorini.

Methods for species delimitation in bumblebees (Hymenoptera, Apidae, Bombus): towards an integrative approach

Delimitation of closely related species is often hindered by the lack of discrete diagnostic morphological characters. This is exemplified in bumblebees (genus Bombus). There have been many attempts to clarify bumblebee taxonomy by using alternative features to discrete morphological characters such as wing shape, DNA, or eco‐chemical traits. Nevertheless each approach has its own limitations. Recent studies have used a multisource approach to gather different lines of speciation evidence in order to draw a strongly supported taxonomic hypothesis in bumblebees. Yet, the resulting taxonomic status is not independent of selected evidence and of consensus methodology (i.e. unanimous procedure, majority, different weighting of evidence). In this article, we compare taxonomic conclusions for a group of taxonomically doubtful species (the Bombus lapidarius‐group) obtained from the four commonly used lines of evidence for species delimitation in bumblebees (geometric morphometric of wing shape, genetic differentiation assessment, sequence‐based species delimitation methods and differentiation of cephalic labial gland secretions). We ultimately aim to assess the usefulness of these lines of evidence as components of an integrative decision framework to delimit bumblebee species. Our results show that analyses based on wing shape do not delineate any obvious cluster. In contrast, nuclear/mitochondrial, sequence‐based species delimitation methods, and analyses based on cephalic labial gland secretions are congruent with each other. This allows setting up an integrative decision framework to establish strongly supported species and subspecies status within bumblebees.

Taxonomic affinity of halictid bee fossils (Hymenoptera: Anthophila) based on geometric morphometrics analyses of wing shape

Taxonomic assignment of fossils can be difficult because descriptions and phylogenetic analyses are often based on a limited number of discrete visible morphological characters. Quantitative, morphometric analyses can help to assign fossil specimens to modern groups, especially when two-dimensional features, such as insect wings, provide easily identifiable landmarks that are not likely to be deformed in the fossil specimens. Here we study taxonomic attribution of bee fossils by analysing wing shape of fossil and contemporary taxa. Our study focuses on the bee family Halictidae (Hymenoptera: Halictidae: Halictinae). Halictinae is a well-known cosmopolitan group including more than 2500 species in five tribes described from robust molecular and morphological analyses. We analysed 202 specimens of 48 species of Halictinae and Nomiinae. To analyse wing shape, we performed a Geometric Morphometrics analysis by using 19 2D-landmarks. Ordination methods, such as canonical variates analysis (CVA) and linear discriminant analysis (LDA), were used to discriminate tribes based on wing shape. CVA and LDA were both powerful enough to recover tribes previously delimited by adult morphology and DNA sequences. We then assigned fossils in CVA and LDA groups based on Mahalanobis distances. The ‘morphometric affinity’ of the two fossil taxa did not confirm their previous taxonomic attribution. By re-examining the fossil description we propose a new taxonomic attribution for Electrolictus antiquus.

Wing Shape of Four New Bee Fossils (Hymenoptera: Anthophila) Provides Insights to Bee Evolution

Bees (Anthophila) are one of the major groups of angiosperm-pollinating insects and accordingly are widely studied in both basic and applied research, for which it is essential to have a clear understanding of their phylogeny, and evolutionary history. Direct evidence of bee evolutionary history has been hindered by a dearth of available fossils needed to determine the timing and tempo of their diversification, as well as episodes of extinction. Here we describe four new compression fossils of bees from three different deposits (Miocene of la Cerdanya, Spain; Oligocene of Céreste, France; and Eocene of the Green River Formation, U.S.A.). We assess the similarity of the forewing shape of the new fossils with extant and fossil taxa using geometric morphometrics analyses. Predictive discriminant analyses show that three fossils share similar forewing shapes with the Apidae [one of uncertain tribal placement and perhaps near Euglossini, one definitive bumble bee (Bombini), and one digger bee (Anthophorini)], while one fossil is more similar to the Andrenidae. The corbiculate fossils are described as Euglossopteryx biesmeijeri De Meulemeester, Michez, & Engel, gen. nov. sp. nov. (type species of Euglossopteryx Dehon & Engel, n. gen.) and Bombus cerdanyensis Dehon, De Meulemeester, & Engel, sp. nov. They provide new information on the distribution and timing of particular corbiculate groups, most notably the extension into North America of possible Eocene-Oligocene cooling-induced extinctions. Protohabropoda pauli De Meulemeester & Michez, gen. nov. sp. nov. (type species of Protohabropoda Dehon & Engel, n. gen.) reinforces previous hypotheses of anthophorine evolution in terms of ecological shifts by the Oligocene from tropical to mesic or xeric habitats. Lastly, a new fossil of the Andreninae, Andrena antoinei Michez & De Meulemeester, sp. nov., further documents the presence of the today widespread genus Andrena Fabricius in the Late Oligocene of France.

Discrimination of haploid and diploid males of Bombus terrestris (Hymenoptera; Apidae) based on wing shape

Presence of diploid males in wild bees reflects inbreeding and provides information about the health status of a colony or population. Detection of diploid males, and discrimination from haploid males and workers, has, however, been limited to molecular diagnostics. Here we present a novel method based on differences in wing shape, e.g., venation patterns in wings. The method is easy to apply and results, for Bombus terrestris, in very high discrimination success. Possible applications of the method are discussed.

A new interpretation of the bee fossil Melitta willardi Cockerell (Hymenoptera, Melittidae) based on geometric morphometrics of the wing

Abstract Although bees are one of the major lineages of pollinators and are today quite diverse, few well-preserved fossils are available from which to establish the tempo of their diversification/extinction since the Early Cretaceous. Here we present a reassessment of the taxonomic affinities of Melitta willardi Cockerell 1909, preserved as a compression fossil from the Florissant shales of Colorado, USA. Based on geometric morphometric wing shape analyses M. willardi cannot be confidently assigned to the genus Melitta Kirby (Anthophila, Melittidae). Instead, the species exhibits phenotypic affinity with the subfamily Andreninae (Anthophila, Andrenidae), but does not appear to belong to any of the known genera therein. Accordingly, we describe a new genus, Andrenopteryx gen. n., based on wing shape as well as additional morphological features and to accommodate M. willardi. The new combination Andrenopteryx willardi (Cockerell) is established.

Microsatellite Analysis of Museum Specimens Reveals Historical Differences in Genetic Diversity between Declining and More Stable Bombus Species

Worldwide most pollinators, e.g. bumblebees, are undergoing global declines. Loss of genetic diversity can play an essential role in these observed declines. In this paper, we investigated the level of genetic diversity of seven declining Bombus species and four more stable species with the use of microsatellite loci. Hereto we genotyped a unique collection of museum specimens. Specimens were collected between 1918 and 1926, in 6 provinces of the Netherlands which allowed us to make interspecific comparisons of genetic diversity. For the stable species B. pascuorum, we also selected populations from two additional time periods: 1949–1955 and 1975–1990. The genetic diversity and population structure in B. pascuorum remained constant over the three time periods. However, populations of declining bumblebee species showed a significantly lower genetic diversity than co-occurring stable species before their major declines. This historical difference indicates that the repeatedly observed reduced genetic diversity in recent populations of declining bumblebee species is not caused solely by the decline itself. The historically low genetic diversity in the declined species may be due to the fact that these species were already rare, making them more vulnerable to the major drivers of bumblebee decline.

Chemical reproductive traits of diploid Bombus terrestris males: Consequences on bumblebee conservation

The current bumblebee decline leads to inbreeding in populations that fosters a loss of allelic diversity and diploid male production. As diploid males are viable and their offspring are sterile, bumblebee populations can quickly fall in a vortex of extinction. In this article, we investigate for the first time a potential premating mechanism through a major chemical reproductive trait (male cephalic labial gland secretions) that could prevent monandrous virgin queens from mating with diploid males. We focus our study on the cephalic labial gland secretions of diploid and haploid males of Bombus terrestris (L.). Contrary to initial expectations, our results do not show any significant differentiation of cephalic labial gland secretions between diploid and haploid specimens. Queens seem therefore to be unable to avoid mating with diploid males based on their compositions of cephalic labial gland secretions. This suggests that the vortex of extinction of diploid males could not be stopped through premating avoidance based on the cephalic labial gland secretions but other mechanisms could avoid mating between diploid males and queens.