Living in the sunlight: micro-environments with higher exposure of sunlight have more abundance and diversity of Hymenoptera in a Brazilian Atlantic Forest fragment

ABSTRACT Tropical forests are three-dimensional with the presence of numerous micro-environments formed by horizontal and vertical gradients. Such micro-environments can affect the nesting preference of organisms, including the trap-nesting Hymenoptera. Bees and wasps are key elements in ecosystems and are considered as sensitive to environmental changes, and trap-nests sampling methodology is widely used in their ecological and conservation studies. However, many uncertainties remain about nesting site preferences. From this perspective, our aim is to assess the diversity descriptors of trap-nesting Hymenoptera in different micro-environments. The sampling was conducted on three micro-environments (canopy, understory and treefall gaps) replicated on ten permanent plots within a Brazilian Atlantic forest hotspot. In each micro-environment, we installed trap-nest stations made by a set of twenty bamboo artificial nests. We found 762 brood cells from ten wasp and five bee species. The rarefaction curves indicate the lower species richness in understory stations, while treefall gaps and canopy stations were not significantly different. We analyzed abundance, mortality and parasitism rates using generalized linear models, but only abundance varies significantly among micro-environments. Our data indicates that trap-nesting Hymenoptera prefer to nest in micro-environments with higher exposure of sunlight. Canopy and treefall gap assemblages are consistently more abundant and diverse than understory probably due the higher temperature and lower humidity. On the other hand, mortality, parasitism rates, and the species composition were similar among environments. Our hypothesis is that the species composition was not affected as these species have a foraging range that encompasses nearby micro-environments.

Comparative morphology and evolution of the cranial musculature in bees (Hymenoptera: Apoidea).

Comparative morphological studies in bees are mostly restricted to the skeleton, and the musculature of bees has not been explored much from this perspective. Here we investigate the head extrinsic musculature under an evolutionary perspective. The musculature of 34 bee species belonging to six major lineages and 26 tribes plus two apoid wasps is described, illustrated, and compared. A standardized terminology for the extrinsic musculature is proposed and aligned with the Hymenoptera Anatomy Ontology (HAO). A total of 12 characters derived from the analysis were optimized onto a summary phylogenetic tree. The musculature was found to be conserved among the main bee lineages, and most variation was found in the proboscis; no modification was found in antennal muscles. Four characters are interpreted as synapomorphies for bees, one for the long-tongued bees, three for Halictinae, and one for Megachilinae. We also found that the Apinae clade of cleptoparasites is supported by some character state transformations and that stingless bees have a unique posterior fronto-pharyngeal muscle. Both sexes display similar morphology, except for having two fixed attachment points for the anterior cranio-mandibular muscle in the males of Andreninae. This study provides a foundation for future investigations on bee head musculature on various taxonomic levels.

A revised genus-level classification for the Neotropical groups of the cleptoparasitic bee tribe Sphecodini Schenck (Hymenoptera, Apidae, Halictinae)

Abstract Sphecodini are a clade of obligatory parasitic Halictinae, comprising 332 species and five genera worldwide. Recently, Sphecodes Latreille was considered polyphyletic according to the results of a molecular phylogeny. Morphological analysis of Neotropical groups revealed groups of species in Sphecodes s.l. that can be recognized at genus level to improve the classification of the tribe. The main objective of this study is to review the generic classification of Sphecodini, with emphasis on Neotropical groups. I propose Austrosphecodes Michener as a separate genus from Sphecodes s.s. and describe a new genus, Melissocleptis gen. nov. for eight known species. Both genera comprise most of the Neotropical species. Nesosphecodes depressus sp. nov. is described for Brazil and a revised diagnosis for Nesosphecodes Engel is given to accommodate this new species and to include data on male terminalia. A revised key for the genera from the Western Hemisphere is provided.

New species of Augochlora (Oxystoglossella) Eickwort (Hymenoptera; Apoidea) from Northeastern Brazil with an identification key for the region.

Within Augochlorini, Augochlora Smith is one of the most speciose genera, with 118 species, and widely distributed groups, occurring from Argentina to Canada. The genus has a remarkable range of social behaviors and nesting substrate preferences. There are two recognized extant subgenera: Augochlora s.s., whose species nest in decaying wood cavities and Oxystoglossella, whose species nest in soil. To date, a single species revision is available for Central Argentina and Uruguay. In the present study we provide a revision of Oxystoglossella species occurring in Northeastern Brazil including the description of three new species: Augochlora bipunctata sp. nov., Augochlora modica sp. nov. and Augochlora tenax sp. nov. Herein Halictus eucnemis Vachal is resurrected from synonymy with Augochlora thalia Smith, and the lectotype of Halictus hemitomes Vachal is presently designated. Augochlora aurinasis is cited for Brazil for the first time. A key to the six species of Oxystoglossella occurring at Northeastern region in Brazil is provided. The three new species are apparently restricted to this region, being one of them probably linked to the Caatinga biome.

A revision of the bee genus Augochlora Smith (Hymenoptera; Apoidea) in Southern South America.

Even after more than 250 years of taxonomic research on bees, there are still many gaps in the knowledge about their identity, classification and distribution patterns. Regarding the New World tribe Augochlorini, many efforts have been made in the last years to describe and organize its diversity. Within the tribe, Augochlora Smith has the widest distribution range, as these bees occur from Argentina to Southern Canada, including Caribbean islands. The genus comprises 124 described species in two extant subgenera, and, to date, two partial revisions are available, accounting for central Argentina and Uruguay and for northeastern Brazil. In the present study we review the Augochlora species occurring in Argentina, Paraguay, Uruguay and the southern Brazilian states of Paraná, Rio Grande do Sul, and Santa Catarina. Specimens from other Brazilian states and South American countries were also examined to help with species circumscription. We found 27 species in the studied area, including nine new species (Augochlora atlantica sp. nov., A. australis sp. nov., A. genalis sp. nov., A. helena sp. nov., A. hestia sp. nov., A. hirsuta sp. nov., A. laevicarinata sp. nov., A. mendax sp. nov., and A. scabrata sp. nov.) and the remaining redescribed when necessary. A key for the species occurring in the studied area is provided. Five new synonymies are proposed and the lectotypes of Augochlora francisca and Halictus esox are presently designated. The geographic distribution of most studied species is associated with the Atlantic biome in many ways.

Ecological impact and population status of non-native bees in a Brazilian urban environment

Abstract The introduction of species is an important global threat to native ecosystems and yet little is known about invasion risks of non-native bees. Moreover, urbanization is increasing rapidly around the world with impact on the bee fauna. The city of Curitiba (Brazil) could be considered a model system for studies under this scope, with historical samplings and monitoring programs since the 1940s. This paper aims to quantify the current ecological impact of the non-native bee species present in Curitiba by using an integrative measure of impact and to define their current population status by classifying them into a unified framework for biological invasions. Ten sites were actively sampled in the metropolitan area of Curitiba during ten months. In addition to Apis mellifera Linnaeus, 1758, we found two others non-native species, Anthidium manicatum (Linnaeus, 1758) and Melipona scutellaris Latreille, 1811. None of them changed the native bee community structure, but they got different values of impact due to their ranges and abundances. While the honeybee is a known invasive species, A. manicatum was also considered invasive and has persisted in the city since the 1940s, interacting with the non-native plant Leonurus japonicus Houtt. M. scutellaris, a cultivated stingless bee species, had its first record for the city and fits the casual non-native category. We suggest monitoring the non-native bee species and actions centered on beekeepers to avoid future illegal introductions. The honey production or hobby interest on stingless bees can offer additional obstacles for species conservation when decoupled of scientific knowledge.

Phylogeny of the Augochlora clade with the description of four new species (Hymenoptera, Apoidea)

ABSTRACT The Augochlora clade includes four genera: Augochlora Smith, Augochlorella Sandhouse, Ceratalictus Moure, and Pereirapis Moure. This is one of the richest and most widespread groups of Augochlorini bees. There are about 150 species, which occur from Argentina to Canada. The species of Augochlora clade are considered solitary to facultatively social, except Ceratalictus for which nothing is known. Wood nesting behavior arose once in the clade, in Augochlora sensu strictu. The objective of this study is to describe four new species and to present a revised phylogenetic analysis of the Augochlora clade for the placement of these species. The morphological matrix comprised 77 characters and 42 terminals, and resulted in two most parsimonious trees. The monophyly of the Augochlora clade is corroborated. Ceratalictus and Pereirapis are considered as sister groups and Ceratalictus inflexus sp. nov. came as sister to other species of Ceratalictus. Augochlora and Augochlorella are monophyletic and sister groups. Both extant subgenera of Augochlora were corroborated as monophyletic. Augochlorella comis is considered as sister group to the rest of Augochlorella species. All Augochlorella new species described belong to the Augochlorella ephyra group. Augochlorella kelliae sp. nov. is phylogenetically related to Augochlorella una. Augochlorella procliva sp. nov. and Augochlorella mavricera sp. nov. constitute a clade with Augochlorella acarinata. Including the new species, Augochlorella has 19 species and Ceratalictus 11 species. A revised key for species of Augochlorella and Ceratalictus is also presented in the Supplementary Information.

New species of Neocorynura Schrottky from Brazil (Hymenoptera: Apoidea).

Neocorynura Schrottky is one of the most speciose genera in the Augochlorini with about 80 species. Neocorynura is primarily distributed in the Neotropical region, from Argentina to Mexico. New species have been described recently, but the Brazilian fauna has not been studied at the same rate. Thus, the objective of this study is to provide a revision of Neocorynura species from Brazil. More than 2,500 specimens were studied along with the examination of primary type specimens and/or high resolution photographs of type material. A total of 44 species were recognized, 24 new species are added to the fauna and figured as N. acuta sp.nov., N. aethra sp.nov., N. amabilis sp.nov., N. arethusa sp.nov., N. aurantia sp.nov., N. carmenta sp.nov., N. dictyata sp.nov., N. eliasi sp.nov., N. hebe sp.nov., N. hyalina sp.nov., N. insolita sp.nov., N. laevistriata sp.nov., N. lamellata sp.nov., N. meloi sp.nov., N. nambikwara sp.nov., N nicolle sp.nov., N. olivacea sp.nov., N perfida sp.nov., N. pilosifacies sp.nov., N. proserpina sp.nov., N. rubicunda sp.nov., N. surrufa sp.nov., N. truncata sp.nov., and N. veneta sp.nov. I designate the lectotype for Corynura oiospermi Schrottky and consider this name as a junior synonym of Halictus codion Vachal. Augochloropsis celaeno Schrottky is considered a junior synonym of Corynura pseudobaccha Cockerell. Casosoma semimarginata Cockerell is resurrected from synonymy with Cacosoma aenigma Gribodo. Neocorynura aenigma (Gribodo) and Neocorynura caligans (Vachal) are removed from the faunal list for Brazil. Half of species recognized here are known only for the female. The males of N. atromarginata (Cockerell), N. cuprifrons (Smith), N. dilutipes (Vachal), N. erinnys (Schrottky), N. melamptera (Moure), N. roxane (Schrottky) and N. semimarginata (Cockerell) and females of N. iopodion (Vachal) and N. jucunda (Smith) are described for the first time. Identification keys to females and males are presented.

Phylogeny and new species of the Neotropical bee genus Paroxystoglossa Moure (Hymenoptera, Apoidea)

ABSTRACT Paroxystoglossa is a solitary, ground-nesting bee genus. It was revised in 1960 and currently includes nine species from Argentina, Brazil and Paraguay. The objectives of this contribution are to provide a morphological phylogeny for the group and to describe two new species: P. levigata n.sp. and P. mourella n.sp. Paroxystoglossa is monophyletic and three species groups are recognized, jocasta species group: (P. mourella n.sp., (P. brachycera, (P. jocasta, P. barbata))), transversa species group: (P. transversa, P. levigata n.sp.), and crossotos species group: (P. mimetica, (P. crossotos, P. seabrai, (P. andromache, P. spiloptera))). The crossotos and transversa species groups were considered as sister groups. Interestingly Paroxystoglossa species have very similar male genital capsules an uncommon pattern among Augochlorini genera. The species groups have a widely redundant distribution indicating replication events in southeastern South America. An updated, illustrated key for species identification is also presented.

Is the capture success of orchid bees (Hymenoptera, Apoidea) influenced by different baited trap designs? A case study from southern Brazil

Orchid bees are increasingly applied on Neotropical biomonitoring and bioindication studies due to the relative easiness of sampling and identification when compared to other bee groups. A considerable number of orchid bee community studies have been adopting baited traps as a sampling method, especially for replication purposes. However, the trap attributes are variable, and hitherto no evaluation of different designs was carried out. Here, five attributes of baited traps were tested: trap volume, number of entrance holes, presence of landing platform, kind of landing platform, and fixation content. We use Mann-Whitney tests to access differences in richness and abundance capture rates for each trap design. We found that volume, number of entrance holes, and fixation content do not influence orchid bees capture. However, the design without landing platforms had a significantly higher capture rate for richness when compared with sanded landing platforms. On the other hand, analyzing the kind of landing platform, we detected a significantly higher richness and abundance for the trap with landing platforms glued with sand. Despite the fact that the effects of different designs tested here were very punctual, we consider that results from samples taken with different baited trap designs are comparable. Some adjustments on trap design can be done according to the particularities of future studies.

Diversidade de abelhas (Hymenoptera, Apidae) ao longo de um gradiente latitudinal na Mata Atlântica / Diversity of bees (Hymenoptera, Apidae) along a latitudinal gradient in the Atlantic Forest

A Mata Atlântica é um dos ambientes mais ricos e ameaçados do mundo, o que deveria ter estimulado em muito o estudo e a conservação do Bioma, mas a fauna de Hymenoptera permanece ainda relativamente pouco conhecida. Em especial, a fauna de abelhas da floresta ombrófila densa é pouco estudada em comparação à fauna das áreas abertas brasileiras. O projeto temático "Biodiversidade de Hymenoptera e Isoptera: riqueza e diversidade ao longo de um gradiente latitudinal na Mata Atlântica - a floresta úmida do leste do Brasil", integrante do Programa Biota-Fapesp, foi idealizado com o objetivo de catalogar térmitas, formigas e famílias selecionadas de vespas ao longo da Mata Atlântica, disponibilizando dados que permitam melhor embasar a conservação deste bioma. O protocolo de amostragem aplicada para a coleta de himenópteros (excluindo as formigas) empregou armadilhas Malaise, pratos-armadilha de cor amarela e esforço similar na varredura de vegetação ao longo de 17 localidades selecionadas, representando um gradiente de quase 20º de latitude na Mata Atlântica, dos Estados da Paraíba até Santa Catarina. Este protocolo foi definido para otimizar a coleta de vespas, sendo as abelhas um produto secundário da amostragem (levantamentos de abelhas em geral utilizam captura em flores ao longo das estações do ano). No entanto, devido à escala regional do projeto e ao grande esforço de amostragem, uma expressiva quantidade de abelhas foi coletada durante o projeto, incluindo novos registros de abelhas para a Mata Atlântica. Foi amostrado um total de 797 espécimes distribuídos em 105 espécies de abelhas; o grupo de abelhas mais rico e abundante foi Meliponina. Uma análise de correspondência "destendenciada" ('detrended correspondence analysis') aplicada à uma matriz de presença ou ausência de Meliponina revela a relativamente fraca influência do gradiente latitudinal na composição das assembléias de abelhas de Mata Atlântica. Uma listagem das espécies...

The Atlantic Forest is one of the most rich and endangered natural environments in the world. It is expected then that this situation would have encouraged the study and conservation of this biome; the Atlantic forest Hymenoptera fauna, however, remains virtually unknown. The bee fauna, for instance, is less studied than that of the Brazilian open areas. The overall ignorance on most of the Atlantic Forest insect faunas based the elaboration of the project "Biodiversity of Hymenoptera and Isoptera: richness and diversity along a latitudinal gradient in the Atlantic Forest - the humid eastern forest of Brazil", within the Biota-Fapesp Program, that aimed to catalog the termites, ants and several wasp families along the Atlantic forest and to publish data that could contribute to the biome conservation. Similar sampling protocols were applied for wasp surveys: the same number of Malaise and pan traps and similar effort on vegetation swapping, along 17 selected localities, representing a gradient of almost 20º of latitude in evergreen pristine Atlantic Forest, from Paraíba to Santa Catarina Brazilian states. These protocols were defined to optimize the collecting of wasps; bees represented a 'secondary product' of the sampling protocol. A sum of 797 specimens representing 105 bee species was sampled; the richest and more abundant bee taxon was the Meliponina. Unfortunately the applied sampling method does not afford comparisons among Atlantic Forest bee assemblages with those of other biomes. The detrended correspondence analysis using only the relative frequencies of Meliponina species revealed a weak correspondence of the stingless bee assemblages' composition with the latitudinal gradient. Despite that, the sampled taxa are possibly restricted to the Atlantic forest and additional data on their distribution deserves to be published. A complete checklist of the sampled bees, their relative frequencies, and the localities where they were...