Microbial Communities of Three Sympatric Australian Stingless Bee Species

Bacterial symbionts of insects have received increasing attention due to their prominent role in nutrient acquisition and defense. In social bees, symbiotic bacteria can maintain colony homeostasis and fitness, and the loss or alteration of the bacterial community may be associated with the ongoing bee decline observed worldwide. However, analyses of microbiota associated with bees have been largely confined to the social honeybees (Apis mellifera) and bumblebees (Bombus spec.), revealing – among other taxa – host-specific lactic acid bacteria (LAB, genus Lactobacillus) that are not found in solitary bees. Here, we characterized the microbiota of three Australian stingless bee species (Apidae: Meliponini) of two phylogenetically distant genera (Tetragonula and Austroplebeia). Besides common plant bacteria, we find LAB in all three species, showing that LAB are shared by honeybees, bumblebees and stingless bees across geographical regions. However, while LAB of the honeybee-associated Firm4–5 clusters were present in Tetragonula, they were lacking in Austroplebeia. Instead, we found a novel clade of likely host-specific LAB in all three Australian stingless bee species which forms a sister clade to a large cluster of Halictidae-associated lactobacilli. Our findings indicate both a phylogenetic and geographical signal of host-specific LAB in stingless bees and highlight stingless bees as an interesting group to investigate the evolutionary history of the bee-LAB association.     

Phylogeny of eusocial Lasioglossum reveals multiple losses of eusociality within a primitively eusocial clade of bees (Hymenoptera: Halictidae)

We performed a phylogenetic analysis of the species, species groups, and subgenera within the predominantly eusocial lineage of Lasioglossum (the Hemihalictus series) based on three protein coding genes: mitochondrial cytochrome oxidase I, nuclear elongation factor 1alpha and long-wavelength rhodopsin. The entire data set consisted of 3421 aligned nucleotide sites, 854 of which were parsimony informative. Analyses by equal weights parsimony, maximum likelihood, and Bayesian methods yielded good resolution among the 53 taxa/populations, with strong bootstrap support and high posterior probabilities for most nodes. There was no significant incongruence among genes, and parsimony, maximum likelihood, and Bayesian methods yielded congruent results. We mapped social behavior onto the resulting tree for 42 of the taxa/populations to infer the likely history of social evolution within Lasioglossum. Our results indicate that eusociality had a single origin within Lasioglossum. Within the predominantly eusocial clade, however, there have been multiple (six) reversals from eusociality to solitary nesting, social polymorphism, or social parasitism, suggesting that these reversals may be more common in primitively eusocial Hymenoptera than previously anticipated. Our results support the view that eusociality is hard to evolve but easily lost. This conclusion is potentially important for understanding the early evolution of the advanced eusocial insects, such as ants, termites, and corbiculate bees.     

Molecular phylogeny and historical biogeography of the large carpenter bees, genus Xylocopa (Hymenoptera: Apidae)

The biogeographical history of major groups of bees with worldwide distributions have often been explained through hypotheses based on Gondwanan vicariance or long distance dispersal events, but until recently these hypotheses have been very difficult, if not impossible, to distinguish. New fossil data, comprehensive information on Mesozoic and Cenozoic coastline positions and the availability of phylogenetically informative DNA markers now makes it feasible to test these hypotheses for some groups of bees. This paper presents historical biogeographical analyses of the genus Xylocopa Latreille, based on phylogenetic analyses of species belonging to 22 subgenera using molecular data from two nuclear genes, elongation factor‐1α (EF‐1α) and phosphoenolpyruvate carboxykinase (PEPCK), combined with previously published morphological and mitochondrial data sets. Phylogenetic analyses based on parsimony and likelihood approaches resulted in several groups of subgenera supported by high bootstrap values (>85%): an American group with the Oriental/Palaearctic subgenera Nyctomelitta and Proxylocopa as sister taxa; a geographically diverse group (Xylocopa s.l); and a group consisting of African and Oriental subgenera. The relationships among these three clades and the subgenus Perixylocopa remained unresolved. The Oriental subgenus Biluna was found to be the sister group of all other carpenter bee subgenera included in this study. Using a relaxed molecular clock calibrated using fossil carpenter bees, we show that the major splits in the carpenter bee phylogeny occurred well after the final breakup of Gondwanaland (the separation of South America and Africa, 100 Mya), but before important Miocene fusion events. Ancestral area analysis showed that the genus Xylocopa most likely had an Oriental‐Palaearctic origin and that the present world distribution of Xylocopa subgenera resulted mainly from independent dispersal events. The influence of Pleistocene glaciations on carpenter bee distributions is also discussed. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society, 2002, 77 , 249–266.     

Co-dependency between a specialist <Emphasis Type="Italic">Andrena</Emphasis> bee and its death camas host, <Emphasis Type="Italic">Toxicoscordion paniculatum</Emphasis>

Among associations of plants and their pollinating bees, mutually specialized pairings are rare. Typically, either pollen specialist (oligolectic) bees are joined by polylectic bees in a flowering species’ pollinator guild, or specialized flowers are pollinated by one or more polylectic bees. The bee Andrena astragali is a narrow oligolege, collecting pollen solely from two nearly identical species of death camas (Toxicoscordion, formerly Zigadenus). Neurotoxic alkaloids of these plants are implicated in sheep and honey bee poisoning. In this study, T. paniculatum, T. venenosum and co-flowering forbs were sampled for bees at 15 sites along a 900-km-long east–west transect across the northern Great Basin plus an altitudinal gradient in northern Utah’s Bear River Range. Only A. astragali bees were regularly seen visiting flowering panicles of these Toxicoscordion. In turn, this bee was never among the 170 bee species caught at 17 species of other prevalent co-occurring wildflowers in the same five state region (38,000 plants surveyed). Our field pollination experiments show that T. paniculatum is primarily an outcrosser dependent on pollinator visitation for most capsule and seed set. Thus, both A. astragali and two sister species of Toxicoscordion are narrowly specialized and co-dependent on each other for reproduction, illustrating a rare case of obligate mutual specialization in bee–plant interactions.     

Ambush Predation of Stingless Bees (<Emphasis Type="Italic">Tetragonisca angustula</Emphasis>) by the Solitary-Foraging Ant <Emphasis Type="Italic">Ectatomma tuberculatum</Emphasis>

Social insect colonies are high-value foraging targets for insectivores, prompting the evolution of complex colony defensive adaptations as well as specialized foraging tactics in social insect predators. Predatory ants that forage on other social insects employ a diverse range of behaviors targeted at specific prey species. Here, we describe a solitary foraging strategy of the ant Ectatomma tuberculatum, on nest guards of the stingless bee Tetragonisca angustula. We observed multiple instances of E. tuberculatum ambushing and successfully capturing the hovering and standing guards of T. angustula near nest entrances. The unique hovering behavior of the guard caste of this bee species, an adaptation to frequent cleptoparasitism by other stingless bees, may make these guards particularly vulnerable to ground-based, ambush attacks by E. tuberculatum. Likewise, the behavior of the foraging ants appears to adaptively exploit the defensive formations and activity patterns of these bees. These observations suggest an adaptive and targeted predatory strategy aimed at gathering external guard bees as prey from these heavily fortified nests.     

Foraging ecology of a local wild bee community in an abandoned Satoyama system in Kanazawa, Central Japan

Satoyama is a traditional land management system in Japan that emphasizes the importance of a harmonious relationship between humans and their environment. It has been in a state of abandonment and destruction since the late 1960s. In 2000, the Japanese government initiated a new environment policy encouraging the use of Satoyama in biodiversity reservation areas. Before the initiation of active restoration works, an assessment of the potential of the Satoyama system is needed. To assess this potential, a study that focused on the foraging ecology of wild bees in a small abandoned Satoyama terraced paddy was carried out from June to October 2003. A total of 372 foraging bees belonging to 35 species in nine families was collected: two species of social bee ( Apis cerana and Bombus diversus ) and two of solitary bee ( Lasioglossum japonicum and Lasioglossum ohei ) dominated. The foraging activities of wild bees were highly influenced by the requirement of food for the colony and the active time of solitary bees. In the study, most wild bees showed a strong preference for abundant flowering plant species that produced a large number of flowers ( Achyranthes japonica and Polygonum thunbergii ). The results showed that an abandoned Satoyama system was an agricultural system that benefited pollinating bees. Careful and planned restoration practices to increase the diversity of foraging resources, microenvironment diversity and potential nesting areas for wild bees are needed to improve the ability of the Satoyama system to support more diverse and abundant wild bee populations.     

Nesting biology of tropical solitary and social sweat bees,Lasioglossum (Dialictus) figueresi Wcislo andL. (D.) aeneiventre (Friese) (Hymenoptera: Halictidae)

Summary The nesting biology of a mainly solitary bee,Lasioglossum (Dialictus) figueresi, is compared with that of a possible relative and mainly eusocial bee,L. (D.) aeneiventre. These bees nest in the ground in highly disturbed areas in the Meseta Central of Costa Rica. Information is provided on social organization, male production, diel and seasonal activity patterns, pollen utilization, natural enemies and nest architecture. L. (D.) figueresi nests within aggregations in vertical earthen banks, and 80–90 % of females are solitary during the nest-provisioning phase. Social nests contain two (or rarely three) females which may be either equal or unequal in reproductive status (i.e. mated with developed ovaries or not). Solitary nests and two-female nests do not experience different rates of parasitism. Mid-way through the dry season, females cease provisioning at a time when otherL. (Dialictus) remain active. Females typically remain within their nests, although they occasionally forage for nectar. This behavior is similar of that of spring gynes of temperate eusocial species. The egg-to-adult developmental rate ofL. (D.) figueresi is unusually slow for halictine bees, however, so that all the adult females die before their brood eclose in April and May, precluding overlap of generations. The eclosed offspring remain in open cells within their natal nests until mid-June, when both males and females emerge to mate. These newly mated females either establish new nests or re-use old ones. L. (D.) aeneiventre nests within aggregations in horizontal ground or in vertical banks. A foundress female digs a nest at the beginning of the dry season, although some re-activate old nests. Foundress nests develop into colonies with various kinds of social organization. In contrast toL. (D.) figueresi, L. (D.) aeneiventre is active nearly all year round, except during periods of heavy rain, and produces up to three broods per year.Sweat bees (Hymenoptera: Halictinae) are a socially heterogeneous group of mainly ground-nesting bees which are abundant world-wide. Intra-specific variation in social behavior is prevalent both within and among populations, presumably indicating social and environmental control of behavioral modifications (see e.g. Sakagami and Munakata, 1972; Eickwort, 1986; Packer, 1990; Yanega, 1988; reviewed in Michener, 1990). The initial stages of hymenopteran social evolution are represented by solitary individuals and those in undifferentiated societies, yet their biology is not well known, as is true for the numerous tropical halictine species or populations (see Michener, 1990).The subgenusDialictus ofLasioglossum is a primarily New World group of several hundred species (Moure and Kurd, 1987). These bees are monotonously similar in structure and appearance, yet diverse in social behavior. FemaleLasioglossum (Dialictus) figueresi are usually solitary, and structurally are very similar to their social relativeL. (D.) aeneiventre (Wcislo, 1990 a). The systematic placement of these species with respect to otherL. (Dialictus) is uncertain, but they have no obvious affinities to other recognized species groups (G. C. Eickwort, pers. comm.). Unusual morphological features, such as large size, yellowish wings and pubescence, and features of the genital organs, may indicate thatL. (D.) figueresi is the more derived of the pair, and may therefore be secondarily solitary, as is known for other sweat bees (Packer, 1991).     

Antennal Asymmetry in Social Behavior of the Australian Stingless Bee, <Emphasis Type="Italic">Tetragonula carbonaria</Emphasis>

The left and right antennae of stingless bees have different roles in learning and recall of olfactory memory. Antennal asymmetry in social behavior is reported here. Approaches and physical contacts were scored in dyads of stingless bees (Tetragonula carbonaria): dyads in which both bees had only their right antennae (left antennae removed) made significantly more physical contacts with each other than dyads in which both bees had only their left antennae. In dyads of one left and one right, it was found, unexpectedly, that the bee with a left antenna approached the bee with the right antenna more often that the other way around, and the bee with the left antenna often attacked (by biting) its hive mate. Hence, the low number of contacts in dyads of bees using their left antennae appears to be due to mutual avoidance. Whereas use of the right antenna stimulates positive contact, the left stimulates avoidance or attack. Via such left-right asymmetries, intact bees may compute behavior directed towards friend and foe. Such antennal asymmetry may have evolved concomitantly with eusocial behavior. We found no evidence that it was associated with significant differences in the number of olfactory or non-olfactory sensilla on the left versus right antenna.     

A Comparison of Social Bee–Plant Networks between Two Urban Areas

In the last decade, several studies demonstrated the effectiveness of ecological network analysis to a better understanding of the structure bee–plant interaction networks; however, such approaches involving urban areas are still scarce. Here, we analyzed two assemblages of corbiculate bees (Apoidea, Apidae) in two geographically distinct urban areas in Brazil. In both study areas, apid bees visiting flowers were captured with an insect net. Surveys were performed biweekly and alternately in each area, over a 1-year period. Both urban areas were very similar for most indices. The two social bee–plant networks were significantly nested, a pattern usually described for bee–plant networks and somehow expected in our study, considering the recognized behavior of social apid bees in exploring a wide range of plant species. The modularity measures were low and very similar for the networks of both urban areas, a finding that could be due at least in part to the low phylogenetic distance between corbiculate bees and the broad dietary habits of the social apid bees. Network-level indices showed that both bee assemblages had a relatively low niche overlap, indicating that the set of social apid species studied exploited differently the arrays of plants available. Species level index (resource range) showed that in both urban areas, Trigona spinipes (Fabr.) and Apis mellifera L. showed the higher number of interactions, a result that demonstrates the importance of these species in social bee–plant interaction networks in urban areas. Similarly to other ecosystems, these two apid species behaved as super-generalists in the two urban areas surveyed herein.     

Subsociality in halictine bees

To look for the occurrence and the significance of brood care in social evolution, I reared six eusocial halictine bee species in laboratory cages enabling the observation of intranest behaviour: Lasioglossum (Evylaeus) laticeps, L. (E.) pauxillum, L. (E.) nigripes, L. (E.) euboeensis, Halictus (Halictus) scabiosae and L. (E.) fulvicorne. All of them were subsocial, each mother caring for her brood. Brood cells were sealed after oviposition with earthen plugs; they were then reopened, visited and closed again. These observations plus the reports in the literature on eleven eusocial species indicate that seventeen species of eusocial halictine bees provide parental care, i.e. are subsocial. Brood care, subsociality, is strongly associated with eusociality. To study reversal from eusociality to subsociality, I have reared the non-eusocial form of two species within which there are or have been eusocial forms: Halictus (H.) rubicundus and Lasioglossum (E.) fratellum. They are secondarily solitary, having lost worker brood. However, both species still show brood care. This suggests that in transitions to eusociality, brood care antedated eusociality. To further examine this issue I reared two truly solitary species that are not derived from eusocial ancestors: Lasioglossum (E.) villosulum and L. (L.) quadrinotatum. Unlike secondarily solitary species, females of both these species close their brood cells after oviposition and ignore their progeny thereafter. This association strongly suggests that the subsocial route with maternal brood care is the route to eusociality in halictine bees.     

Population genetics of wild and managed pollinators: implications for crop pollination and the genetic integrity of wild bees

Loss of habitat and chemical use associated with agriculture can cause population declines of wild pollinators. Less is known about the evolutionary consequences of interactions between species used in commercial agriculture and wild pollinators. Given population declines of many wild bee species, it is crucial to understand if commercial queens become established in natural areas, if wild bees visit agricultural fields and have the potential to interact with commercial bees, and if gene flow occurs between commercial and wild bees. We drew on a long-term data set that documents commercial bumble bee (Bombus impatiens) use in New England, and we conducted genetic analyses of foraging B. impatiens from areas with varying intensities of commercial bee use. In agricultural areas with a history of commercial bee use we also sampled bees directly from commercial hives. We found significant genetic differences among foraging B. impatiens and B. impatiens sampled directly from hives (average pairwise F′_ST = 0.14), but not among samples of foraging bees from natural areas (average F′_ST among foraging bees?=?0.002). Furthermore, Bayesian analysis of population structure revealed that foraging bees caught in areas with a history of commercial bee use grouped with samples from natural areas. These results document an agricultural setting where there was no widespread introgression of alleles from commercial bumble bees to wild bumble bees, commercial bumble bees did not become established in natural areas, and wild bees were providing pollination services to crops.     

Socially peaceful: foragers of the eusocial bee Lasioglossum malachurum are not aggressive against non-nestmates in circle-tube arenas

Due to the universally found nestmate recognition in eusocial insects, it is predictable that non-nestmates interact aggressively. In sweat bees (Hymenoptera: Halictidae), this trend was largely shown for queen–queen interactions, but data on worker–worker interactions are still scarce and somehow controversial. We studied behavioural interactions between foragers of the eusocial and ground-nesting bee Lasioglossum malachurum within circle-tubes. Independently of colony membership, bees exhibited high frequencies of cooperative behaviours, together with lack of aggression and moderate avoidance of social interactions. The cooperative mutual passing was the most frequently recorded behaviour. Size difference between the opponents had no effects on cooperation or avoidance. In a heterospecific experiment, bee foragers were observed to react more aggressively and to pass very rarely towards cuckoo bees, suggesting that our results were not biased by the circle-tube methodology. Our results and comparisons with other bee species suggest that studying worker interactions may be not enough to predict the social organisation in bees. Whatever the evolutionary meaning of this generalised tolerance towards conspecifics, the present findings are somehow in agreement with recent studies showing that L. malachurum colonies may have imperfect nestmate recognition and often include a mixture of related and unrelated workers.     

The cuticular profiles of Australian stingless bees are shaped by resin of the eucalypt tree Corymbia torelliana

Bees are known to collect pollen and nectar to provide their larvae and themselves with food. That bees, especially the tropical stingless bees (Apidae: Meliponini), also collect plant resins has, however, been barely addressed in scientific studies on resource use in bees. Resins are used for nest construction, nest maintenance and nest defence. Furthermore, some South‐East Asian species transfer resin‐derived terpenes to their cuticular profiles. The resin requirement of bees is in turn used by certain plant species, which attract bees either for pollination by providing resin in their inflorescences, or for seed dispersal by providing resin in their seed capsules (mellitochory). Mellitochory is found in the eucalypt tree Corymbia torelliana, the resin of which is collected by Australian stingless bees. We investigated how the interaction between C. torelliana and resin‐collecting bees affects the chemical ecology of two Australian stingless bee genera by comparing the chemical profiles of eight bee species with resin from C. torelliana fruits. The two bee genera differed significantly in their chemical profiles. Similar to South‐East Asian stingless bees, 51% of all compounds on the body surfaces of the five Tetragonula species were most likely derived from plant resins. Up to 32 compounds were identical with compounds from C. torelliana resin, suggesting that Tetragonula species include C. torelliana compounds in their chemical profiles. By contrast, few or none resinous compounds were found on the body surfaces of the three Austroplebeia species sampled. However, one prominent but as yet unknown substance was found in both C. torelliana resin and the chemical profiles of all Tetragonula and four Austroplebeia colonies sampled, suggesting that most colonies (76%) gathered resin from C. torelliana. Hence, C. torelliana resin may be commonly collected by Australian stingless bees and, along with resins from other plant species, shape their chemical ecology.     

Hunting for healthy microbiomes: determining the core microbiomes of <Emphasis Type="Italic">Ceratina,Megalopta</Emphasis>, and <Emphasis Type="Italic">Apis</Emphasis> bees and how they associate with microbes in bee collected pollen

Social corbiculate bees such as honey bees and bumble bees maintain a specific beneficial core microbiome which is absent in wild bees. It has been suggested that maintaining this microbiome can prevent disease and keep bees healthy. The main aim of our study was to identify if there are any core bacterial groups in the non-corbiculate bees Ceratina and Megalopta that have been previously overlooked. We additionally test for associations between the core bee microbes and pollen provisions to look for potential transmission between the two. We identify three enterotypes in Ceratina samples, with thirteen core bacterial phylotypes in Ceratina females: Rosenbergiella, Pseudomonas, Gilliamella, Lactobacillus, Caulobacter, Snodgrassella, Acinetobacter, Corynebacterium, Sphingomonas, Commensalibacter, Methylobacterium, Massilia, and Stenotrophomonas, plus 19 in pollen (6 of which are shared by bees). Unlike Apis bees, whose gut microbial community differs compared to their pollen, Ceratina adults and pollen largely share a similar microbial composition and enterotype difference was largely explained by pollen age. Megalopta displays a highly diverse composition of microbes throughout all adults, yet Lactobacillus and Saccharibacter were prevalent in 90% of adults as core bacteria. Only Lactobacillus was both a core bee and pollen provision microbe in all three species. The consequences of such diversity in core microbiota between bee genera and their associations with pollen are discussed in relation to identifying potentially beneficial microbial taxa in wild bees to aid the conservation of wild, understudied, non-model bee species.     

Global stingless bee phylogeny supports ancient divergence,vicariance, and long distance dispersal

Stingless bees (Meliponini) are one of only two highly eusocial bees, the other being the well studied honey bee (Apini). Unlike Apini, with only 11 species in the single genus Apis, stingless bees are a large and diverse taxon comprising some 60 genera, many of which are poorly known. This is the first attempt to infer a phylogeny of the group that includes the world fauna and extensive molecular data. Understanding the evolutionary relationships of these bees would provide a basis for behavioural studies within an evolutionary framework, illuminating the origins of complex social behaviour, such as the employment of dance and sound to communicate the location of food or shelter. In addition to a global phylogeny, we also provide estimates of divergence times and ancestral biogeograhic distributions of the major groups. Bayesian and maximum likelihood analyses strongly support a principal division of Meliponini into Old and New World groups, with the Afrotropical+Indo‐Malay/Australian clades comprising the sister group to the large Neotropical clade. The meliponine crown clade is inferred to be of late Gondwanan origin (approximately 80 Mya), undergoing radiations in the Afrotropical and Indo‐Malayan/Australasian regions, approximately 50–60 Mya. In the New World, major diversifications occurred approximately 30–40 Mya. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 206–232.     

Review of bees as guests in termite nests,with a new record of the communal bee, <Emphasis Type="Italic">Gaesochira obscura</Emphasis> (Smith, 1879) (Hymenoptera,Apidae), in nests of <Emphasis Type="Italic">Anoplotermes banksi</Emphasis> Emerson, 1925 (Isoptera,Termitidae, Apicotermitinae)

Associations between bees and termites are documented infrequently, but records are available for bee species ranging in behavior from solitary to highly eusocial. The subtribe Meliponina (stingless bees) is the most common bee group reported in termite nests, and, for some species, the occupancy of termite nests may be obligatory. The records of solitary bees nesting within termite nests include species of the tribes Emphorini, Centridini, Megachilini, and Paracolletini. Most of these bees can probably nest in other substrates, and their relationships with termite nests are presumably opportunistic. This study provides a review of published records of bees as guests in termite nests, and also describes the aggregation of nests of Gaesochira obscura within one nest of Anoplotermes banksi in Brazilian Amazonia. One termite nest contained at least ten nest entrances of G. obscura, with burrows 4–6 mm in diameter and about 10 cm long. Each burrow ended in brood cells in different stages of food provisioning and larval development. As commonly reported for other associations of this nature, there was no connection between the tunnels of bees and those of termites. This record adds important data on the biology of A. banksi. Because this is a single record, it is impossible to classify G. obscura either as a termitophile or termitariophile; this species may be opportunistic in relation to nesting substrate.     

The effect of habitat fragmentation on the bee visitor assemblages of three Australian tropical rainforest tree species

Tropical forest loss and fragmentation can change bee community dynamics and potentially interrupt plant–pollinator relationships. While bee community responses to forest fragmentation have been investigated in a number of tropical regions, no studies have focused on this topic in Australia. In this study, we examine taxonomic and functional diversity of bees visiting flowers of three tree species across small and large rainforest fragments in Australian tropical landscapes. We found lower taxonomic diversity of bees visiting flowers of trees in small rainforest fragments compared with large forest fragments and show that bee species in small fragments were subsets of species in larger fragments. Bees visiting trees in small fragments also had higher mean body sizes than those in larger fragments, suggesting that small‐sized bees may be less likely to persist in small fragments. Lastly, we found reductions in the abundance of eusocial stingless bees visiting flowers in small fragments compared to large fragments. These results suggest that pollinator visits to native trees living in small tropical forest remnants may be reduced, which may in turn impact on a range of processes, potentially including forest regeneration and diversity maintenance in small forest remnants in Australian tropical countryside landscapes.     

Phylogeny of the bee genus Lasioglossum (Hymenoptera: Halictidae) based on mitochondrial COI sequence data

The bee genus Lasioglossum Curtis is a model taxon for studying the evolutionary origins of and reversals in eusociality. This paper presents a phylogenetic analysis of Lasioglossum species and subgenera based on a data set consisting of 1240 bp of the mitochondrial cytochrome oxidase I (COI) gene for seventy-seven taxa (sixty-six ingroup and eleven outgroup taxa). Maximum parsimony was used to analyse the data set (using paup *4.0) by a variety of weighting methods, including equal weights, a priori weighting and a posteriori weighting. All methods yielded roughly congruent results. Michener’s Hemihalictus series was found to be monophyletic in all analyses but one, while his Lasioglossum series formed a basal, paraphyletic assemblage in all analyses but one. Chilalictus was consistently found to be a basal taxon of Lasioglossum sensu lato and Lasioglossum sensu stricto was found to be monophyletic. Within the Hemihalictus series, major lineages included Dialictus + Paralictus, the acarinate Evylaeus + Hemihalictus + Sudila and the carinate Evylaeus + Sphecodogastra. Relationships within the Hemihalictus series were highly stable to altered weighting schemes, while relationships among the basal subgenera in the Lasioglossum series (Lasioglossum s.s., Chilalictus, Parasphecodes and Ctenonomia) were unclear. The social parasite of Dialictus, Paralictus, is consistently and unambiguously placed well within Dialictus, thus rendering Dialictus paraphyletic. The implications of this for understanding the origins of social parasitism are discussed.     

Phylogenetic relationships in Iranian <Emphasis Type="Italic">Scutellaria</Emphasis> (Lamiaceae) based on nuclear ribosomal ITS and chloroplast <Emphasis Type="Italic">trn</Emphasis>L-F DNA data

The genus Scutellaria (Lamiaceae, Scutellarioidae) includes taxa with a wide range of pharmacological traits, which make this genus economically important. Previous taxonomical studies on Scutellaria were mainly based on morphological and chromosomal characters. The infrageneric classification of the genus has been the subject of a long controversy among authors, and delimitation of taxa remains problematic. In this study, the phylogeny of the genus was evaluated using nrDNA ITS and trnL-F sequences, with a focus on the 42 Iranian taxa belonging to subgenera Apeltanthus and Scutellaria. In both ITS and trnL-F trees, there were two main clades within the genus corresponding to the two subgenera Scutellaria and Apeltanthus. In the subg. Scutellaria, the sect. Anaspis was monophyletic, but the monophyly of sect. Scutellaria was not supported. In this subgenus, S. galericulata was located away from other species in both trees, supported the isolation of this species from the sect. Scutellaria and inclusion in the sect. Galericularia as was previously considered by some authors. In subg. Apeltanthus, sect. Apeltanthus and sect. Lupulinaria were not monophyletic; furthermore, the monophyly of subsect. Lupulinaria was not supported. Based on the results of this study, it can be suggested that the sectional classification of both subgenera Apeltanthus and Scutellaria should be revised.     

Decreasing Abundance,Increasing Diversity and Changing Structure of the Wild Bee Community (Hymenoptera: Anthophila) along an Urbanization Gradient

Background

Wild bees are important pollinators that have declined in diversity and abundance during the last decades. Habitat destruction and fragmentation associated with urbanization are reported as part of the main causes of this decline. Urbanization involves dramatic changes of the landscape, increasing the proportion of impervious surface while decreasing that of green areas. Few studies have investigated the effects of urbanization on bee communities. We assessed changes in the abundance, species richness, and composition of wild bee community along an urbanization gradient.

Methodology/Principal Findings

Over two years and on a monthly basis, bees were sampled with colored pan traps and insect nets at 24 sites located along an urbanization gradient. Landscape structure within three different radii was measured at each study site. We captured 291 wild bee species. The abundance of wild bees was negatively correlated with the proportion of impervious surface, while species richness reached a maximum at an intermediate (50%) proportion of impervious surface. The structure of the community changed along the urbanization gradient with more parasitic species in sites with an intermediate proportion of impervious surface. There were also greater numbers of cavity-nesting species and long-tongued species in sites with intermediate or higher proportion of impervious surface. However, urbanization had no effect on the occurrence of species depending on their social behavior or body size.

Conclusions/Significance

We found nearly a third of the wild bee fauna known from France in our study sites. Indeed, urban areas supported a diverse bee community, but sites with an intermediate level of urbanization were the most speciose ones, including greater proportion of parasitic species. The presence of a diverse array of bee species even in the most urbanized area makes these pollinators worthy of being a flagship group to raise the awareness of urban citizens about biodiversity.