Suction feeding in orchid bees (Apidae: Euglossini)

Energy flux during nectar feeding is maximized at an intermediate sugar concentration, the value of which depends on the morphology of the feeding apparatus and the modality of fluid feeding. Biomechanical models predict that a shift from capillary-based lapping to suction feeding will lead to a decrease in this optimal sugar concentration. Here, I demonstrate that the four major genera of orchid bees (Apidae: Euglossini) are suction feeders and provide experimental evidence that the feeding optimum for one species, Euglossa imperialis, falls below the optimum for bee taxa that lap.     

Generic relationships within the Euglossini (Hymenoptera: Apidae)

ABSTRACT. .The relationships among the euglossine genera are examined. These genera form two natural groups, Exaerete+Euglossa , and Eufriesea+(Eulaema+Aglae). Each genus is discussed separately giving generic synonymy, biological notes and a complete species list. In addition, a key to the genera is provided.     

Phylogeny of the orchid bees (Hymenoptera: Apinae: Euglossini): DNA and morphology yield equivalent patterns

Orchid bees (Euglossini) are spectacular long-tongued Neotropical bees important in the pollination of Neotropical long-corolla flowers, particularly some orchids. Besides remarkably long tongues, males in particular exhibit other flower-related adaptations, including setal brushes on the foretarsi used for rasping the petals of orchids while collecting aromatic compounds. These compounds are stored in large swollen tibiae and are thought to play an important role in courtship behavior. Euglossini are also unusual in lacking sociality; they are the only tribe among the corbiculate bees that are not eusocial, and two of the genera are cleptoparasitic. Each genus exhibits distinct behavioral traits including nest architecture and host-parasite interactions, yet their evolution is unknown. Despite previous phylogenetic studies of on morphological characters, the relationships among the five euglossine genera remain under debate. We investigate euglossine generic relationships using DNA sequence data from four genes and new morphological characters. The morphological and molecular data yield congruent evolutionary patterns, and combining the data gives a fully resolved and well supported phylogeny of Euglossini.     

Effectiveness of two sampling protocols to survey orchid bees (Hymenoptera: Apidae) in the Neotropics

We compared the performances of two different methodologies for collecting orchid bees (Hymenoptera: Apidae): active sampling (at chemical baits with hand nets) versus passive sampling (with baited traps). Data were retrieved from previous field studies in which both methodologies were employed simultaneously. We evaluated the effectiveness of each method in terms of the number of species collected and the composition of the bee species through paired t test and Permutational Multivariate Analysis of Variance, respectively. Active sampling collected significantly more bee species than passive sampling in all but one study. The composition of bee species collected by each of the two methods was also significantly different. Larger bees belonging to Eulaema and Eufriesea species tended to be more frequent in traps. In contrast, all Euglossa species presented comparatively higher relative frequencies with active sampling. We challenge previous assertions that both methodologies are equally effective and argue that insisting on the sole use of baited traps in orchid-bee inventories may result in retrieving strongly biased raw data which will eventually lead to incorrect conclusions about the actual composition of orchid-bee assemblages. Since orchid bees have been more and more used as key organisms in studies on environmental impact in the Neotropics, particularly in Brazil, and the passive sampling methodology has been adopted as the “official” one by the Brazilian environmental agency (IBAMA), we suggest a re-evaluation of this practice and the adoption of the more expensive, but more reliable, methodology of using well-trained field researchers to actively collect data on these important Neotropical pollinators.     

Effects of habitat type change on taxonomic and functional composition of orchid bees (Apidae: Euglossini) in the Brazilian Amazon

Land use change impact species richness and functional diversity (FD). In the Brazilian Amazon, we examined the impacts of oil palm plantations on orchid bee (Apidae: Euglossini) species using abundance and FD. We collected male orchid bees in oil palm plantation (PALM), legal reserves (LR), and riparian corridors (APP), and then we used morphological and life-history traits to characterize each species. We evaluated differences in bee body size by comparing intertegular span values. We tested the influence of habitat on taxonomic and functional parameters of orchid bees by applying a partial redundancy analysis (pRDA). We contrasted FD by calculating species richness, functional richness, and functional dispersion. We sampled 1176 bees from 30 species in 18 sampling days across 2015 and 2016. Males from PALM were 13.6% bigger than those in LR areas, and bees from APP showed a similar pattern compared to LR and PALM. Less than 15% of the variation in species composition was related to the distance among sampling sites, and 8% was due to habitat structure. In our pRDA, the spatial difference explained 6% of the variation in orchid bee traits, but there were no effects of habitat parameters upon FD. FD was reduced with land use change caused by oil palm plantations. Our findings support the belief that many bees are impacted by cultivated lands. Nevertheless, the functional similarity between LRs and APPs reflects common structural elements between them, although we did not find significant relationship between functional composition and habitat structure that we evaluated.     

Microsatellite loci for Euglossa annectans (Hymenoptera: Apidae) and their variability in other orchid bees

Orchid or euglossine bees are conspicuous Hymenoptera of the Neotropics, where they pollinate numerous plants, including orchids. Allozyme-based analyses have suggested that their populations suffer from inbreeding, as evidenced by so-called diploid male production. We have developed nine polymorphic microsatellite loci for the widespread Euglossa annectans, with observed heterozygosities ranging from 0.143 to 0.952 and between 2 and 9 alleles per species. These loci will be useful for analysis of relatedness, population genetic structure and diploid male production in this and related species.     

Diversity and distribution of orchid bees (Hymenoptera: Apidae) with a revised checklist of species

The aim of this study was to investigate the diversity and distribution patterns of orchid bees (Euglossina). Cluster and correlation analyses were applied to data extracted from 28 orchid-bee surveys throughout the Neotropical Region. The 28 sampling sites were grouped in three main biogeographic areas that roughly correspond to the Amazonian Basin, the Atlantic Forest and Central America. These three regions, as well as subregions within each of them, correspond approximately to biogeographic components identified through phylogeny-based analyses for other bees and organisms. The Amazonian Forest as a whole has the richest fauna and the highest levels of endemism. The Atlantic Forest, on the other hand, showed the poorest fauna and the lowest levels of endemism. However, a major neotropical biome, in which orchid bees are known to occur, has not been sampled yet, the savanna-like cerrado. At least 30% of the species are endemic to each biome. An updated checklist of the species of Euglossina is provided.     

Diversity and community composition of euglossine bee assemblages (Hymenoptera: Apidae) in western Amazonia

Tropical forests are known for their diverse insect fauna. We aimed to determine the effect and relative importance of latitude, elevation and climatic factors affecting species richness and turnover in euglossine bee assemblages along a gradient of 18° latitude from tropical rainforests to subtropical, deciduous dry forests in Peru and Bolivia. Sixteen forest sites were sampled during the dry season. Variance partitioning techniques were applied to assess the relative effects of the spatial and environmental variables on species richness and composition. Furthermore, we conducted a Species Indicator Analysis to find characteristic species for the biogeographic zones. There was a significant decrease in species richness towards the subtropical area. The best predictors of species richness were precipitation and its consequences on soil properties as well as temperature seasonality. The abundance of euglossines was most closely related to precipitation and soil-pH, but the causal links of abundance to these factors is unclear since soil-pH itself is correlated to a drastic turnover of vegetation structure. Based on the analysis of assemblage composition we propose three different assemblages with a transitional zone at the southern tropical area. The biogeographical distribution of euglossine bees along our study transect appears to be primarily related to climatic conditions and does not reflect the common subdividion of Amazonia into drainage systems.     

Effects of fragments and landscape characteristics on the orchid bee richness (Apidae: Euglossini) in an urban matrix,southwestern Amazonia

Orchid bees are important pollinators in tropical forests. Although studies have already detected effects of habitat loss and forest fragmentation on bee assemblages, little is known about orchid bees in urban forest fragments. The aim of this study was to analyse the influence of forest fragments (size and edge index) and landscape features (forest cover area and built-up area around the forest fragments, connectivity and spatial distance from the urban center) on the abundance, richness and composition of orchid bees. Male bees were attracted by odoriferous baits and collected in ten forest fragments of different sizes. In total, we collected 3166 male bees belonging to 4 genera and 38 species. The increase of the built-up area and the reduction of the forest cover area around the forest fragments decreased the abundance and richness of bees. We recorded a smaller number of bees in areas closer to the urban center. We failed to find a significant relationship between abundance/richness of bees and forest fragment area, edge index, and connectivity. We observed that the faunistic dissimilarity was positively correlated with the geographic distance between forest fragments. The forest fragments that presented greater landscape dissimilarity also presented greater faunistic dissimilarity. Our results suggest that the matrix structure around the forest fragments is an important factor that influences the Euglossini bee assemblages inside these forest fragments. Based on our results, we believed that the conservation of fragments with a larger forest cover area and smaller built-up areas around them contribute to the maintenance of the diversity of orchid bees and their pollination services.     

Drinking with a very long proboscis: Functional morphology of orchid bee mouthparts (Euglossini,Apidae, Hymenoptera)

Neotropical orchid bees (Euglossini) possess the longest proboscides among bees. In this study, we compared the feeding behavior and functional morphology of mouthparts in two similarly large-sized species of Euglossa that differ greatly in proboscis length. Feeding observations and experiments conducted under semi-natural conditions were combined with micro-morphological examination using LM, SEM and micro CT techniques. The morphometric comparison showed that only the components of the mouthparts that form the food tube differ in length, while the proximal components, which are responsible for proboscis movements, are similar in size. This study represents the first documentation of lapping behaviour in Euglossini. We demonstrate that Euglossa bees use a lapping-sucking mode of feeding to take up small amounts of fluid, and a purely suctorial technique for larger fluid quantities. The mouthpart movements are largely similar to that in other long-tongued bees, except that the postmentum in Euglossa can be extended, greatly enhancing the protraction of the glossa. This results in a maximal functional length that is about 50% longer than the length of the food canal composing parts of the proboscis. The nectar uptake and the sensory equipment of the proboscis are discussed in context to flower probing.     

The behaviour of male orchid bees (Apidae,Hymenoptera, Insecta) and the question of leks

The male territories of two species of orchid bees, Eulaema meriana and Euglossa imperialis, are described. These territories consist of a perch, where the males display on the trunk of a tree, and a route flown from and back to the perch. Territories are located in treefalls or other large light gaps in tropical forest. The territories of these two species differ in the height of the perch and amount of light reaching it, size of the perch tree, and period of activity. A definition of lek is given. These male euglossines form facultative leks in large treefalls. The relationships between male territories, female foraging ranges, and collection of aromatic material by males are discussed.     

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.     

Population genetic structure of orchid bees (Euglossini) in anthropogenically altered landscapes

Habitat degradation and fragmentation are widespread phenomena in tropical regions. Negative effects on the biota are numerous, ranging from interruption of gene flow among populations, to the loss of genetic diversity within populations, to a decline in species richness over time. Orchid bees (Hymenoptera: Apidae: Euglossini) are of major conservation interest due to their function as pollinators of numerous orchid species and other tropical plants. Here, we used microsatellite markers to investigate the effects of geographic distance and habitat fragmentation on gene flow among populations. Populations of Euglossa dilemma in three geographic regions??the Yucat??n peninsula (Mexico), Veracruz (Mexico), and Florida (USA)??were genetically structured predominantly across the regions, with the strength of differentiation among populations being positively correlated with geographic distance. Within geographic regions only little substructure was found, suggesting that dispersal is substantial in the absence of geographic or ecological barriers. In a second study, patterns of genetic differentiation among eight species of Euglossa were not related to habitat fragmentation following deforestation in southern Mexico (Veracruz). Specifically, most bee populations in the 9,800?ha forest remnant of Los Tuxtlas (Volcano San Martin) were neither differentiated from, nor had less genetic diversity than, populations in near-continuous forest separated from Los Tuxtlas by 130?km of agricultural land. Either occasional long distance dispersal across open areas has buffered the expected genetic effects of fragmentation, or the history of fragmentation in southern Mexico is too recent to have caused measurable shifts in allelic composition.     

A re-evaluation of the phylogenetic relationships in the Apidae (Hymenoptera)

ABSTRACT .
The relationships between the four tribes in the bee family Apidae are re-examined. Characteristics of the postgena, presternum, antenna cleaner, arolium, female hind tibia and male genitalia, among others, support the placement of these tribes in three subfamilies: Meliponinae (Meliponini), Apinae (Apini) and Bombinae (Euglossini + Bombini). The Apinae is the sister group of the Bombinae. This tribal arrangement was originally proposed, using different characteristics, by Winston & Michener (1977).     

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.     

Nesting biology and nest re-use ofEulaema nigrita (Hymenoptera: Apidae,Euglossini)

Summary The nesting behavior of the euglossine beeEulaema nigrita was observed in the laboratory after being transferred from three nests to observation boxes. Nests were re-used by successive generations with more than one female working in each re-use process. Associated females were always of the same generation. All females that shared a nest foraged for construction materials (mud, excrement and resin) and each constructed, provisioned and oviposited in her own cells. The number of cells constructed by each female ranged from one to 23. Females stayed in the nests for periods ranging from 15 to 59 days during the hot and wet season and from five to 78 days during the cool and dry season. The egg-to-adult period was related to climatic conditions, and in all re-use processes it was longer than the time of residence of the females in the nests. The meloid beetleMeloetyphlus attacephalus was the only nest parasite. Due to the attack method employed by this parasite, the presence of more than one female in the nest did not result in improved nest defense.     

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.     

Molecular identification of species of the genus Euglossa Latreille (Hymenoptera: Apidae, Euglossini)

Euglossine bees interact with more than 60 plant families of the Neotropical region. The richness and abundance of these bees have been intensively studied in different ecosystems using the methodology of capturing males with chemical baits. Females are poorly known for most of the species and morphological characters for their taxonomic classification have not yet been described. The purpose of this study was to use allozymes and restriction patterns of the mitochondrial regions 16S and Cyt b to identify species of Euglossa Latreille. Bees were collected while visiting Thevetia peruviana (Apocynaceae) flowers in five cities of the state of S?o Paulo, Brazil. Three Euglossa species were identified among the 305 individuals collected. Euglossa cordata (L.) was the only species found in all cities, while E. securigera Dressler and E. townsendi Cockerell were restricted to two and one cities respectively. EST-3 was a diagnostic marker, whereas ICD, MDH, ME and PGM were informative for species identification when used in combination. Restriction by VspI of the amplified 16S fragment differentiated the three species and showed intraspecific polymorphism for E. cordata and E. securigera. The Cyt b region showed distinctive patterns for E. townsendi but it was not possible to differentiate the other two species. Our results describe potentially useful genetic markers for the identification of Euglossa spp. at the species and group level.     

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 .