Phylogeny of bumble bees in the New World subgenus Fervidobombus (Hymenoptera: Apidae): congruence of molecular and morphological data

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 (Bombus transversalis and Bombus pullatus) comprise a monophyletic clade, along with a third species, Bombus atratus, which is widely distributed in South America, including lowland subtropical habitats.     

Allozyme variation in bumble bees (Hymenoptera: Apidae)

Allozyme variation at an average of 37.3 loci was assessed in queens of 16 Bombus and 2 Psithyrus bumble bee species from North America. The mean expected heterozygosity (H) for the Bombus species was 0.008 +/- 0.006 (95% confidence limits) and that for the Psithyrus was 0.007 +/- 0.007. These levels are significantly lower than found in other Hymenoptera but are comparable to those found in previous studies of bumble bees based on far fewer loci. Neutral mutation and random genetic drift can account for the observed variation, but this implies a very small effective population size for species of bumble bees.     

Queen-worker conflict and social parasitism in bumble bees (Hymenoptera: Apidae)

Psithyrus ashtoni (Hymenoptera: Apidae) is an obligate, workerless bumble, bee social parasite which invades nests of Bombus affinis. Although parasites are limited by host worker defence to invading very small colonies, there is considerable flexibility in the way parasites control host brood bionomics once they are accepted inside the nest. A study of 46 parasitized and 22 non-parasitized laboratory colonies of B. affinis showed that P. ashtoni females cohabited with host queens and workers while the worker force increased, but not to the maximum normally achieved in non-parasitized nests. While colony reproductive success was correlated with the number of workers reared, parasites risked being killed or ejected from the comb by workers, after the queen had lost dominance. Host bees usually succeeded in rearing offspring, and Psithyrus reproductive success was related to the ability of parasites to control proportional investment in the two species. In addition to displacing P. ashtoni females, host bees ate the eggs of parasites and ejected their larvae. These behaviours were also exhibited by workers in the later stages of development of non-parasitized colonies. These results indicate that social parasites are at least partially subject to the conflict of genetic self-interest between the queen and her workers which is believed to influence the control of reproductive investment in haplodiploid Bombus societies.     

Bombus folsomi and the origin of Philippine bumble bees (Hymenoptera: Apidae)

Abstract The type female of Bombus folsomi (Frison) is redescribed and synomymized with B.ephippiatus Say. It is concluded that the putative type locality of B.folsomi , Mt Kinabalu in Borneo, is in error and that the genus appears to be absent from Borneo. This lends support to the hypothesis that bumble bees immigrated to the Philippines by an unusual northern route through Taiwan. More rigorous testing of this idea will require further cladistic analysis of the genus Bombus .     

Temporal dynamics of facultative social parasitism in bumble bees (Hymenoptera: Apidae)

Parasitical behaviour is thought to play an important, yet so far largely unknown role in the evolution and maintenance of insect sociality. In this study, the influence of interspecific, facultative social parasitism upon bumble bee, Bombus, nesting biology was explored. Queens of B. affinis were introduced into 38 various-sized laboratory colonies of B. terricola. Foreign queens were very successful at killing the resident queen and usurping pre-worker nests (77% wins), although usurpation success decreased to 30% in colonies that contained workers. In addition, B. affinis queens were unable to suppress ovarian development in B. terricola workers that emerged prior to nest usurpation, resulting in the eventual death or expulsion of the foreign queen. In contrast, ovarian development was supporessed in workers that eclosed in the presence of a foreign queen, and in workers that were less than 5 h old when confined in small experimental boxes with a B. affinis queen. Foreign queens that usurped a nest prior to worker emergence were assisted by B. terricola workers, and achieved similar reproductive success when compared to a cohort of 22 B. affinis queens that initiated their own nests in the laboratory. These results suggest that there are periods in the ontogeny of Bombus nest defence, recognition and dominance during which bees are sensitive to the effects of nest parasitism. In bumble bees, and in other social insects, individuals of some species may exploit these weaknesses for their own reproductive benefit.     

Phylogeny of the World Anthophorini (Hymenoptera: Apoidea: Apidae)

Abstract.  The phylogenetic relationships of members of the long-tongued bee tribe Anthophorini were investigated. Twenty-six ingroup taxa, representing all known genera and the most common subgenera of the Anthophorini, were included in a cladistic analysis of the tribe based on 51 morphological characters. The analysis yielded the generic relationships: (( Habrophorula , Elaphropoda ) ( Habropoda ( Deltoptila ( Pachymelus ( Amegilla , Anthophora ))))). The monophyly of the Anthophorini and each of its genera was confirmed. A second tribe, Habropodini, as postulated by some authors, is paraphyletic. On the basis of the results of the cladistic analysis and the biogeographical data of Anthophorini, an evolutionary scenario is postulated in which all genera of Anthophorini probably evolved in the Late Cretaceous, except Amegilla and Anthophora , which originated in the Oligocene. The northern part of south-east Asia (India to south-east China) is regarded as the most probable place of origin and radiation of the Anthophorini. The New World was probably colonized three times independently by Habropoda (upper Cretaceous to Tertiary), by an ancestral lineage of Deltoptila (upper Cretaceous to Tertiary) and by Anthophora (Tertiary to Quaternary).     

A preliminary cladistic investigation of relationships among the bumble bees (Hymenoptera, Apidae)

ABSTRACT. 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.     

The tracheal mite Locustacarus buchneri in South American native bumble bees (Hymenoptera: Apidae)

As in other regions of the world, bumble bees (Bombus spp.) are important pollinators in the neotropics. Despite its relevance, knowledge on their health is still limited in the region. While external acari are known to occur in these insects, presence of the internal, tracheal mite Locustacarus buchneri is here reported for first time. After the examination of 2,508 individuals of eight Bombus species from Argentina, two workers of Bombus bellicosus and one of Bombus atratus were found parasitized by L. buchneri in localities within San Luis and Buenos Aires provinces, respectively. The rare occurrence recorded agrees with findings from elsewhere in the world.     

Phylogeny and biogeography of the oil-collecting bee subgenus Centris (Wagenknechtia) (Hymenoptera,Apidae)

Centris (Wagenknechtia) is the only subgenus of centridine bees that occurs almost exclusively in the Andean Region. This study investigates the monophyly of C. (Wagenknechtia) proposing a hypothesis for the relationships among its species using 42 morphological characters of adults. The analysis resulted in one most parsimonious tree corroborating the monophyly of this group of oil-collecting bees, with the relationships among its species as follows: ((C. escomeli + C. moldenkei) (C. muralis (C. rhodophthalma + C. vardyorum) (C. cineraria + C. orellanai))). Based on these results plus available distributional records, a cladistic biogeographic analysis was performed. The resulting hypothesis of the relationships between the endemism areas in the Andean Region is as follows: (((Atacama–Puna) (Desert–Atacama–Puna)) ((Pampean–Chaco–Puna) ((Monte–Coquimban) (Coquimban (Patagonian–Valdivian Forest–Coquimban–Maule–Monte–Santiago–Magellanic Forest))))).     

Phylogeny,diversification patterns and historical biogeography of euglossine orchid bees (Hymenoptera: Apidae)

The orchid bees constitute a clade of prominent insect pollinators distributed throughout the Neotropical region. Males of all species collect fragrances from natural sources, including flowers, decaying vegetation and fungi, and store them in specialized leg pockets to later expose during courtship display. In addition, orchid bees provide pollination services to a diverse array of Neotropical angiosperms when foraging for food and nesting materials. However, despite their ecological importance, little is known about the evolutionary history of orchid bees. Here, we present a comprehensive molecular phylogenetic analysis based on ～4.0 kb of DNA from four loci [cytochrome oxidase (CO1), elongation factor 1‐α (EF1‐α), arginine kinase (ArgK) and RNA polymerase II (Pol‐II)] across the entire tribe Euglossini, including all five genera, eight subgenera and 126 of the approximately 200 known species. We investigated lineage diversification using fossil‐calibrated molecular clocks and the evolution of morphological traits using disparity‐through‐time plots. In addition, we inferred past biogeographical events by implementing model‐based likelihood methods. Our dataset supports a new view on generic relationships and indicates that the cleptoparasitic genus Exaerete is sister to the remaining orchid bee genera. Our divergence time estimates indicate that extant orchid bee lineages shared a most recent common ancestor at 27–42 Mya. In addition, our analysis of morphology shows that tongue length and body size experienced rapid disparity bursts that coincide with the origin of diverse genera (Euglossa and Eufriesea). Finally, our analysis of historical biogeography indicates that early diversification episodes shared a history on both sides of Mesoamerica, where orchid bees dispersed across the Caribbean, and through a Panamanian connection, thus reinforcing the hypothesis that recent geological events (e.g. the formation of the isthmus of Panama) contributed to the diversification of the rich Neotropical biota. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 552–572.     

Historical biogeography, divergence times, and diversification patterns of bumble bees (Hymenoptera: Apidae: Bombus)

Bumble bees (Bombus) are a cold-adapted, largely alpine group that can elucidate patterns of Holarctic historical biogeography, particularly in comparison to the alpine plants with which they likely coevolved. A recently published molecular phylogeny of bumble bees provides uniquely comprehensive species sampling for exploring historical patterns of distribution and diversification. Using this phylogeny and detailed data on extant distributions, I reconstruct the historical distribution of bumble bees in a temporal framework, estimating divergence times using fossil data and molecular rates derived from the literature. The nearly comprehensive phylogeny allows assessment of the tempo of diversification within the bumble bees using lineage-through-time plots and diversification statistics, which have been performed with special consideration to confidence intervals. These analyses reveal movements of Bombus concordant with geographic and climatic events of the late Cenozoic. The initial diversification of extant bumble bee lineages was estimated at around 25 to 40 Ma, near the Eocene-Oligocene boundary 34 Ma, a period of dramatic global cooling. Dispersal-vicariance analysis (DIVA) predicted an Old World Bombus ancestor, with early diversification events largely restricted to the eastern Old World. The numerous intercontinental dispersal events occurred mostly in the direction of Old World to New World and North America to South America. Early movements from the Palearctic into the Nearctic most likely took place after 20 Ma and may have coincided with a period of Miocene cooling that gave rise to taiga habitat across Beringia. Subsequent dispersal between these regions is estimated to have occurred among boreal and tundra-adapted species mostly in the last 5 million years. Radiations are estimated in both Nearctic and Neotropical regions at approximately 6 to 8 Ma and after 3.5 Ma, concordant with the opening of land corridors between the continents.     

The communities of bumble bees (Hymenoptera: Apidae, Bombini) in the city of Novosibirsk and its environs

The diversity and abundance of bumble bees in the city of Novosibirsk and its environs have been investigated. A list of registered species is given with an estimation of their relative abundance. For the first time Bombus pseudobaicalensis Vogt, 1911, B. saltuarius (Skorikov, 1931), Psithyrus barbutellus (Kirby, 1802), Ps. norvegicus (Sparre-Schneider, 1918), and Ps. sylvestris (Lepeletier, 1832) are recorded in Novosibirsk oblast. The seasonal dynamics of the number and diversity of these insects has been studied.     

The New World oil‐collecting bees Centris and Epicharis (Hymenoptera,Apidae): molecular phylogeny and biogeographic history

We present a first comprehensive time‐calibrated phylogeny for two Neotropical genera of bees, Centris and Epicharis, whose females collect floral oil together with pollen for larval provisioning, and that traditionally have been grouped in the tribe Centridini. Our analyses rely on a matrix of 167 taxa and 4228 aligned nucleotides for the subfamily Apinae, with denser sample of Centris and Epicharis. Centris and Epicharis are strongly supported monophyletic groups, but Centridini is paraphyletic in relation to the corbiculate bees. The inner phylogenetic relationships of Epicharis agree with the current taxonomic classification. In Centris, three main clades were recovered, namely the Centris, Trachina and Melacentris groups. Inner relationships in Centris suggest the reinstatement of some subgenera and proposition of new ones. Early diversification of Centris and Epicharis took place at the tropical regions of South America. Epicharis and the Melacentris group in Centris mostly diversified in that region, expanding over Central America and tropical North America only recently in the last 3 My. The groups Trachina and Centris present a complex biogeographic history, with expansions to the Nearctic region, the Antilles, and temperate regions of South America in the late Oligocene and Miocene.     

Hazards of insecticides to the bumble bees Bombus impatiens (Hymenoptera: Apidae) foraging on flowering white clover in turf

Insecticides used on turf are sometimes applied to areas with flowering weeds that attract honey bees and native pollinators. We tested residual effects of such treatments on colony vitality and behavior of the bumble bees Bombus impatiens Cresson foraging on turf containingwhite clover, Trifolium repens L. Imidacloprid, a syst emic chloronicotinyl used for preventive control of root-feeding grubs, was applied as granules, followed by irrigation, or sprayed as a wettable powder, with or without irrigation. Hives were confined on the plots in large field cages after residues had dried and colony vitality (i.e., numbers of brood, workers, and honey pots, and weights of queens, workers, and whole colonies with hives) was evaluated after 28-30 d. Workers' foraging activity and defensive response to an aggressive stimulus also were evaluated. In another test, weedy turf was sprayed with chlorpyrifos, carbaryl, or cyfluthrin at labeled rates for surface-feeding pests. Bee colonies were confined on the plots after residues had dried, with effects on colony vitality evaluated after 14 d. Finally, foraging activity of wild bumble bees was monitored on open plots to determine if insecticide-treated areas were avoided. Imidacloprid granules, and imidacloprid sprays applied with posttreatment irrigation, had no effect on colony vitality or workers' behavior, suggesting that such treatments pose little systemic or residual hazard to bumble bees. In contrast, exposure to dry nonirrigated residues of all of the aforementioned insecticides had severe impact on colony vitality. Foraging workers did not avoid insecticide-treated areas. Means by which turf managers can reduce hazards of insecticide applications to pollinators are discussed.     

Microsatellite loci for euglossine bees (Hymenoptera: Apidae)

The Neotropical Euglossini (Hymenoptera: Apidae) are important pollinators of many flowering plants, particularly orchids. Lack of highly polymorphic genetic markers for euglossine species has limited the study of their social organization and inbreeding. We therefore developed microsatellite markers for two species, Eulaema nigrita (11 loci) and Euglossa cordata (nine loci), most of which were highly polymorphic in the source species and in a range of related euglossine bees.     

American foulbrood and African honey bees (Hymenoptera: Apidae)

We have taken samples of honey from individual beekeepers (N = 64), and of domestic (N = 35) and imported honey (N = 15) retailed in supermarkets in several sub-Saharan countries and cultivated these samples for Paenibacillus larvae subsp. larvae Heyndrickx et al. causing American foulbrood in honey bee colonies. The results are compared with samples of similar backgrounds and treated the same way but collected in Sweden (N = 35). No P. larvae subsp. larvae spores were found in any honey produced in Africa south of the Sahara although honey imported into this region frequently contains the pathogen. Swedish honey frequently contains P. larvae subsp. larvae spores although the general level of visibly infected bee colonies is low (roughly 0.5%). The results suggest that large parts of Africa may be free from American foulbrood. Behavioral studies (hygienic behavior) on Apis mellifera subsp. scutellata Lepeletier in Zimbabwe suggest that hygienic behavior of African bees could influence the apparent low level, or even absence of American foulbrood in large parts of Africa.     

Phylogeny of the bee family Melittidae (Hymenoptera: Anthophila) based on combined molecular and morphological data

The bee family Melittidae comprises a small, but biologically fascinating, group of mostly oligolectic bees, some of which are oil collecting. Phylogenetic relationships within this family are poorly understood and some genera cannot be placed with confidence at the subfamily level. We analysed melittid phylogeny using a combined dataset of five nuclear genes [28S, elongation factor‐1α (EF‐1α, F2 copy), long‐wavelength rhodopsin, Na‐K ATPase and RNA polymerase II] spanning 4842 bp plus 68 adult morphological characters. Our study included 25% of the species‐level diversity and 81% of the generic‐level diversity and included all previously recognized tribes and subfamilies. We analysed the dataset using parsimony, maximum likelihood and Bayesian methods. All methods yielded congruent results. All topologies recovered the three previously recognized subfamilies (Dasypodainae, Melittinae, Meganomiinae), but two genera (Afrodasypoda and Promelitta) are transferred from Dasypodainae to Melittinae. On the basis of our tree topologies we identify four tribes (Dasypodaini comb.n. , Hesperapini stat.n. , Macropidini comb.n. and Melittini), only one of which (Melittini) matches a widely used classification. Lastly, we discuss the evolution of host‐plant association in the light of our new phylogenetic hypothesis. Our results strongly support multiple independent origins of oil‐collecting behaviour in the Melittinae.     

A molecular phylogeny of the Old World stingless bees (Hymenoptera: Apidae: Meliponini) and the non-monophyly of the large genus Trigona

Abstract.  We examined the inter- and infrageneric relationships of Old World Meliponini with a near-complete sampling of supra-specific taxa. DNA sequences for the taxa were collected from four genes (mitochondrial 16S rRNA, nuclear long-wavelength rhodopsin copy 1 (opsin), elongation factor-1α copy F2 and arginine kinase). Additional sampling of New World taxa indicated that Trigona sensu lato is not monophyletic: Trigona from the Indo-Malayan/Australasian Regions forms a large clade distantly related to the Neotropical Trigona . A separate clade comprises the Afrotropical meliponines, and includes the 'minute' species found in the Afrotropical, Indo-Malayan and Australasian Regions. The Neotropical genus Melipona , by contrast with previous investigations, is not the sister lineage to the remaining stingless bees, but falls within the strongly supported Neotropical clade. These results constitute the framework for a revised classification and ongoing biological investigations of Meliponini. A single taxonomic change, Heterotrigona bakeri stat.n. , is proposed on the basis of sequence divergence.     

Phylogenetic divergence of the Australian allodapine bees (Hymenoptera: Apidae)

Allodapine bees are most diverse in Africa but are distributed throughout the Old World tropical and Austral regions. They are considered useful for studies into the evolution of social behaviour since they exhibit the full range of social organisation from solitary to highly eusocial (sensu; ). Five genera are found in Australia, namely Braunsapis, Exoneurella, Exoneura, Brevineura, and Inquilina. Sociality and life histories are well documented for the exoneurine genera (review in ) and Inquilina is an obligate social parasite of species of Exoneura (). In this paper, maximum parsimony and maximum likelihood methods using molecular sequence data from two mitochondrial gene regions (cyt b and COI) and a single nuclear gene region (EF-1alpha) are used to reconstruct phylogenetic relationships of the Australian allodapine genera. Results suggest that the exoneurine group (Brevineura, Exoneurella, and Exoneura+Inquilina) diverged very rapidly and are monophyletic to the exclusion of other (primarily African) allodapine genera. A clade containing Australian species of Braunsapis is also monophyletic to the exclusion of African congeners. Braunsapis is not phylogenetically close to, and is a more derived group than the exoneurine group and probably came to occupy the Australian plate via a later dispersal through the southern Asian region. It is unclear at this point how the exoneurine group came to occupy the Australian plate and possible scenarios are discussed.