Proventricular structure in solitary bees (Hymenoptera: Apoidea)

Proventricular structure, analyzed by scanning electronic microscopy, is compared among 28 species of solitary bees representing four families. Observations on the shapes of proventricular folds and on hair-like cuticular projections are presented, discussed, and suggested as useful to future studies of bee systematics.     

A cladistic analysis of the nomadine bees (Hymenoptera: Apoidea)

Abstract. This study compares the results of Rozen's cladistic analysis of the larvae of fifteen genera of cleptoparasitic bees in the subfamily Nomadinae with an independent data set of adult characters for the same genera. Adult characters exhibited considerably higher levels of homoplasy and poorer resolution of cladistic relationships, with multiple equally parsimonious cladograms. However, comparison of a Nelson consensus tree based on adult characters with the cladogram based on larval characters reveals three components consistently supported in both analyses (the tribes Epeolini and Ammobatini, and Neopasites + Neolarra) , one component supported only by adult characters (Isepeolus + Protepeolus) , and one terminal component supported only by larval characters (Nomada + Ammobatini), as well as several more inclusive groupings based on larval characters that are difficult to compare with the adult consensus tree because it shows so much less resolution. When adult and larval characters are combined in a single data matrix, the resulting cladogram closely resembles the cladogram based on larval characters alone, although levels of homoplasy are considerably higher than in the larval analysis.
A preliminary analysis of adult characters for thirty-four genera in the Nomadinae also exhibited high levels of homoplasy and very large numbers of equally parsimonious cladograms. Nevertheless, certain consistent monophyletic groupings, most notably the Epeolini and Ammobatini, were also supported in this analysis. The one currently recognized tribe whose monophyly has received no support from any analysis is the Nomadini.
The relevance of these phylogenetic hypotheses to our understanding of host associations and variable features of egg morphology and oviposition behaviour in nomadine bees is briefly discussed.     

Guidelines on sampling intensity of bees (Hymenoptera: Apoidea: Apiformes)

Changes in bee fauna, such as the disappearance of certain species or increasing abundance of others, are very important. The common belief that pollinating insects are facing problems also prompts detailed studies of the bee fauna in order to track on-going changes. Assessment of the state of bee communities in particular ecosystems or ecosystem complexes within a landscape, tracing the course of trends in fauna and also predicting their future structures resulting from current changes, are only possible if sampling is carried out at an appropriate frequency, so that representative materials are obtained. The aim of the present study was to determine what sampling intensity during the growing season would enable the collection of representative materials to evaluate species diversity of bees in a study area. Repeated bee sampling at monthly intervals throughout the period of activity of bees resulted in the identification of 73 bee species, corresponding to 51.4 % of the estimated number of bee species in the study area. When samples were obtained twice a month, 93 bee species were captured, accounting for 65.5 % of the estimated number of bee species. When sampling took place nearly four times a month, 108 bee species were captured, making up 76 % of the estimated number of bee species; the materials obtained at this sampling rate may be regarded as representative. The importance of accounting for the phenology of a given animal group during a sampling effort is also emphasised.     

The major opsin in bees (Insecta: Hymenoptera): A promising nuclear gene for higher level phylogenetics.

We report the phylogenetic utility of the nuclear gene encoding the long-wavelength opsin (LW Rh) for tribes of bees. Aligned nucleotide sequences were examined in multiple taxa from the four tribes comprising the corbiculate bees within the subfamily Apinae. Phylogenetic analyses of sequence variation in a 502-bp fragment (approx 40% of the coding region) strongly supported the monophyly of each of the four tribes, which are well established from previous studies of morphology and DNA. Trees estimated from parsimony and maximum likelihood analyses of LW Rh sequences show a strongly supported relationship between the tribes Meliponini and Bombini, a relationship that has been found uniformly in studies of other genes (28S, 16S, and cytochrome b). All of the tribal clades as well as relationships among the tribes are supported by high bootstrap values, suggesting the utility of LW Rh in estimating tribal and subfamily rank for these bees. The sequences exhibit minimal base composition bias. Both 1st + 2nd and 3rd position sites provide information for estimating a reliable tree topology. These results suggest that LW Rh, which has not been reported previously in studies of organismal phylogenetics, could provide important new data from the nuclear genome for phylogeny reconstruction.     

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.     

The evolution of food-producing glands in eusocial bees (Apoidea,Hymenoptera)1

A food-producing role for cephalic exocrine glands has arisen independently in both taxa of highly eusocial bees, Apis and Meliponini. With several exceptions, there is little evidence that food is produced by glands of solitary bees or by most bees at lower levels of sociality. We suggest that this association with sociality is due to four adaptive features of these glands: (1) food from the glands allows feces from queens and larvae to have a small volume, (2) the queen's fecundity can be increased, (3) nutrient recovery via cannibalism can be facilitated, and (4) rearing of emergency replacement queens is accelerated. Acceleration of the rearing of other castes and of queens in the normal process of colony fission is not clearly an advantage ascribed to these glands. Trophic eggs produced by meliponine colony workers are analogous to the secretions from food-producing glands in Meliponini and Apis workers.     

Parasitism rates in relation to nest site in bees and wasps (Hymenoptera: Apoidea)

To account for differences in occurrence of social behavior in different lineages of bees, Michener (1985) hypothesized that ground nests are more easily located by parasites than are twig nests. In the former case parasites search in two dimensions, while in the latter they search in three-dimensional space. One prediction derived from this hypothesis is that ground nests will have higher rates of parasitism than twig nests. A survey of published reports on rates of cell parasitism for 92 species of nesting bees and wasps (Apoidea) shows no significant differences in mean parasitism rates between these two classes of nests. The analyses were repeated at the generic level (N=44), yielding the same pattern. These data may be biased due to phylogenetic effects. Paired comparisons (n=11 pairs) of related taxa that differ in nest site show that ground-nesting taxa more often have higher rates of parasitism than twig-nesters. The use of artificial trap-nests to study twig-nesters significantly enhances the success rate of parasites. This bias, as well as several other limitations, suggests that experimental studies of the host-searching capabilities of parasites and predators may be more efficacious than such comparative tests.     

Wild bees (Hymenoptera: Apoidea) in an urban botanical garden in Buenos Aires,Argentina

We assessed bee diversity and abundance in a botanical garden in the city of Buenos Aires during two consecutive spring–summer periods. Every 15 days, we collected by hand-netting bee specimens seen foraging on flowers during five-minute censuses at every entomophilous herbaceous plant. A total of 66 bee species (Hymenoptera: Apoidea) were recorded. Richness and abundance of native, generalist, and above-ground nesting (cavities) species were higher than that of exotic, specialist, and below-ground (soil) nesting ones, respectively. Social bees were more abundant, while the richness of solitary species was higher than that of social bees. Cleptoparasitic species were represented by a high number of species, even though only a few individuals were captured. Our results suggest that the studied area is an important bee reservoir within the city.     

Floral resource partitioning by social wasps (Hymenoptera: Vespidae) and bees (Hymenoptera: Apoidea) in an area of caatinga in Brazil

Wasps and bees are important components of flower visitor guilds in the Neotropical region. The abundance of social wasps and bees, collected during flower visits, was used to calculate niche breadth and niche overlap indices, in order to characterize the utilization of the floral resources by those insects. Samples were taken monthly, for 13 months (156h), by two collectors which captured the specimens on flowers using entomological nets, while walking along a way in an area of caatinga at Itatim, State of Bahia, Brazil. Wasps and bees heavily visited a few sources of floral resources, but the most visited plants by each group were different, and few plant species were important for both groups, resulting in low niche overlap between bees and wasps. The niche overlap among wasp species was generally higher than among bee species. The general overlap, which takes into consideration all species together was low (< 30%).     

Contribution of native bees and Africanized honey bees (Hymenoptera:Apoidea) to Solanaceae crop pollination in tropical México

In the tropics the contribution of bees as pollinators of important economic indigenous crops remains largely undocumented. We studied the diversity of bee species visiting indigenous tomato (Solanum lycopersicon) and habanero pepper (Capsicum chinense) in subtropical Yucatán, México. The contribution of two native bees, Exomalopsis (E) and Augochloropsis (AG) was compared with that of the introduced Africanized Apis mellífera (HB) for pollinating unvisited flowers in both crops. Apis mellífera and stingless bees were dominant in habanero pepper but solitary bee species were important visitors of tomato. In spite of both crops being autogamic, there was a significant contribution of native bees for pollination of both tomato and habanero pepper. The comparison of fruit weight, number of seeds, and a pollination index based on the latter showed that E and AG were more effective pollinators compared to HB in both crops (ca. Spear’s index of ca. 0.7 vs. 0.35 respectively). In tomato, a further evaluation of the contribution to pollination provided by the three bee species was made using the rate of visits to flowers. Although E and AG were the most efficient pollinators at single flower visits in tomato, none of the three species (including HB) were able to provide single visits to all flowers per unit time to the crop. Our results underline the importance of maintaining diverse assemblages and abundant populations of bee species that can synergically contribute to the productivity of tomato and hot pepper in the Neotropics.     

New species and unexpected diversity of socially parasitic bees in the genus Inquilina Michener (Hymenoptera: Apoidea: Apidae)

Abstract  Allodapine bees present particular problems for taxonomy due to a high level of morphological conservatism in adults, even between genera. However, this tribe of bees also presents a unique opportunity to explore the evolution of social parasitism because of the comparatively large number of origins of socially parasitic species. Morphological differences presented here, along with DNA sequence data and molecular phylogenetic analyses, indicate a much larger number of Australian social parasite species in the genus Inquilina than previously anticipated, and suggest that the final number of socially parasitic species may be considerable. We describe five new species and present sequence data that will help elucidate the delineation of further new species. Inquilina provides a unique opportunity to study the evolution of social parasitism in social insects, but further studies will need to encompass both population genetic and phylogenetic approaches.     

Records of bees (Hymenoptera: Apoidea) new to both Georgia and Transcaucasia revealed by DNA barcoding

Community composition of local pollinator assemblages, population dynamics and distribution patterns of many species are still poorly known, especially in little studied regions like Transcaucasia. We generated a total of 189 DNA barcodes from bees collected in 14 apple orchards in central and eastern Georgia to extend our knowledge of their distribution. Sequences (ca. 700 bp of the mitochondrial cytochrome oxidase I gene) were compared to the NCBI nucleotide database. Using a threshold of 98% similarity, analyses suggested that the sampled bees belong to 36 species of Apidae, Andrenidae, Halictidae, and Megachilidae. We report five bee species for the first time for Georgia: Andrena alfkenella Perkins, 1914, Halictus cochlearitarsis Dours, 1871, Nomada goodeniana (Kirby, 1802), Nomada ruficornis (Linnaeus, 1758) and Eucera nigrescens Pérez, 1879, four of which are also new for the Transcaucasia region. With these additions, the number of bee species known to occur in Georgia increases to 371.     

Forested landscapes promote richness and abundance of native bees (Hymenoptera: Apoidea: Anthophila) in Wisconsin apple orchards

Wild bees provide vital pollination services for many native and agricultural plant species, yet the landscape conditions needed to support wild bee populations are not well understood or appreciated. We assessed the influence of landscape composition on bee abundance and species richness in apple (Malus spp.) orchards of northeastern Wisconsin during the spring flowering period. A diverse community of bee species occurs in these apple orchards, dominated by wild bees in the families Andrenidae and Halictidae and the honey bee, Apis mellifera L. Proportion of forest area in the surrounding landscape was a significant positive predictor of wild bee abundance in orchards, with strongest effects at a GIS (Geographic Information Systems) buffer distance of 1,000 m or greater. Forest area also was positively associated with species richness, showing strongest effects at a buffer distance of 2,000 m. Nonagricultural developed land (homes, lawns, etcetera) was significantly negatively associated with species richness at buffer distances >750 m and wild bee abundance in bowl traps at all distances. Other landscape variables statistically associated with species richness or abundance of wild bees included proportion area of pasture (positive) and proportion area of roads (negative). Forest area was not associated with honey bee abundance at any buffer distance. These results provide clear evidence that the landscape surrounding apple orchards, especially the proportion of forest area, affects richness and abundance of wild bees during the spring flowering period and should be a part of sustainable land management strategies in agro-ecosystems of northeastern Wisconsin and other apple growing regions.     

Comparative morphology of the antenna cleaner in bees (Apoidea)

The morphology of the antenna cleaners of 175 species of bees representing more than 50 genera was investigated and described comparatively. An ancestral type of antenna cleaner was found in most short-tongued bees (except Oxaeidae and Ctenoplectridae) as well as in Nomadinae, Ceratini, and some Megachilidae. A further ancestral character (shallow notch), however, was only found in some of these bees (but throughout Colletidae and Halictidae). A derived type of antenna cleaner was found in Anthophora, Eucera and related genera, as well as in Melecta, Ancyloscelis, and Ctenoplectra. Several further (derived) types of antenna cleaners were found in Anthophoridae and Apidae. A large variety of antenna cleaners (ancestral as well as derived ones) were found in the Megachilidae. Possible phylogenetic consequences are suggested in Figure 37.     

The mycoflora of domesticated and wild bees (Apoidea)

Fungi including yeasts are common in the honey stomachs and provisions of diverse bees. They may be parasites, commensals or mutualistic. Yeasts, singly or in association with bacteria, are pioneer colonizers during a microbial succession in larval cells of many subterranean bees. They are followed by fungi such asAspergillus, Penicillium, Emericellopsis, Sartorya, Pseudoarachniotus, Gymnoascus, Carpenteles andFusarium. Aspergillus flavus andSaccharomyces spp. are pathogenic to many species of bees, and fungi are the main cause of declining alkali bee populations. There are 124 species of fungi, including 36 new records, associated with Apoidea; 49 species are associated with alkali bees.Cooperative investigations of the Plant Sciences Research (L. R. Batra) and Entomology (G. E. Bohart) Divisions, Agricultural Research Service, U.S. Department of Agriculture, and the Utah Agricultural Experiment Station, Logan, Utah.     

Quantitative Aspects of Antenna Grooming in Bees (Apoidea: Hymenoptera)

Antenna grooming in more than 100 species of bees (Apoidea), representing 34 genera of the 7 major bee families is recorded and quantitatively analysed. Most species of bees fall into one of two groups with respect to repetitive antenna cleaning: “Uniscrapers” predominantly clean their antennae with one stroke, “biscrapers” mostly with two subsequent strokes. Uniscrapers are more consequent in their behaviour than biscrapers. Most biscrapers occasionally clean their antenna with one or three strokes. Individual variation in the ratio of stroke repetition is considerably larger in bi- than in uniscrapers. In several species males and females differ with respect to their antenna cleaning behaviour, females tend to be more uniscraping. Most species of the families Colletidae, Halictidae, and Andrenidae, as well as the species of the genera Ceratina and Nomada (Anthophoridae) are biscrapers. Almost all species of Melittidae, Megachilidae, Apidae, and Anthophoridae (except Ceratina and Nomada) are uniscrapers. Bees with an antenna cleaner with ancestral (plesiomorphic) morphology are mostly, but not always, biscrapers, those with a derived antenna cleaner are always uniscrapers. Bees with a derived antenna cleaner perform on average less cleaning actions and strokes than those with an ancestral antenna cleaner. Uniscrapers with an ancestral strigilis do on average more cleaning actions per minute than biscrapers, thus they compensate partly for the fewer number of strokes. But nevertheless the uniscrapers do fewer strokes than the biscrapers (both with an ancestral strigilis). Females clean their antennae on average more often than males. It is interpreted that the behaviour of uniscraping and a derived morphology of the antenna cleaner result in greater efficiency than the status which is ancestral for Apoidea (biscraping and a plesiomorphic antenna cleaner).     

基于线粒体和核基因序列的蜜蜂属系统发育分析

文章测定了中国分布的蜜蜂属(Apis)5种蜜蜂22个样本的线粒体基因ND2、CO2、16S rRNA以及核基因ITPR的序列,对序列的碱基组成和蜜蜂种间的遗传距离进行了分析。结合下载的蜜蜂属其他4个种的相关序列,采用最大简约法、邻接法和最大似然法重建了蜜蜂属系统发育关系。系统发育分析结果支持蜜蜂属划分为3个类群,即小蜜蜂类群(包括小蜜蜂和黑小蜜蜂)、大蜜蜂类群(包括大蜜蜂和黑大蜜蜂)和穴居蜜蜂类群(西方蜜蜂、东方蜜蜂、沙巴蜂、苏拉威西蜂、绿努蜂),且小蜜蜂类群较早分化。结果还显示,我国海南岛的大蜜蜂和大陆的大蜜蜂之间可能存在较大的遗传分歧。