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The external morphology and phylogeny of the bees
... More recent studies have dealt with the internal anatomy indicating the relevance of the internal skeleton as a data source for phylogenetic reconstructions at the family and genus levels (Porto et al., 2016(Porto et al., , 2017Meira and Gonçalves, 2018;Porto and Almeida, 2019). Michener (1944) pioneered exploration of the internal skeleton for bee systematics, and the seminal phylogenetic studies of Roig-Alsina and Michener (1993) with LT bees and Alexander and Michener (1995) with ST bees included some internal morphology characters. Non-skeletal tissues have been somewhat explored in bee systematics. ...
... Furthermore, works with more specific muscle groups for Apidae were consulted (Wolff, 1875;Morison, 1927;Urban, 1963;Graf, 1965Graf, , 1972. For the topological characterization of muscle attachments, we examined the studies of Michener (1944Michener ( , 2007, Roig-Alsina and Michener (1993), Alexander and Michener (1995), Porto et al. (2016), and Porto and Almeida (2019). Additionally, the pertinent literature available related to the head's extrinsic musculature in several Hymenoptera groups (Duncan, 1939;Alam, 1951;Matsuda, 1957Matsuda, , 1965Vilhelmsen et al., 1996;Beutel and Vilhelmsen, 2007) and the HAO (Yoder et al., 2010) were also consulted. ...
... Fig. 6). The variation of the maxillary process of hypostoma was already noted by Michener (1944), Graf (1972), Eickwort (1969), Alexander and Michener (1995;character 22, state 2), Michener (2007), and by Porto et al. (2016;character 22, state 1). According to the literature (Snodgrass, 1942;Youssef, 1971;Mik o et al., 2007), the fixed point of attachment of the gen-car is located in the occipital region of the head capsule but all examined Apoidea have the origin in the genal region. ...
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.
... The classification of Colletinae, one of the eight subfamilies currently recognized in the bee family Colletidae (see: Almeida et al., 2012Almeida et al., , 2019, has been the subject of a long-standing controversy amongst bee systematists. In an early paper on the phylogeny and classification of bees (Michener, 1944), Colletinae was considered in its broadest sense to include Paracolletini s.l. [= Paracolletes Smith, 1853 plus Callomelittinae, Neopasiphaeinae and Scrapterinae of Almeida et al. (2012Almeida et al. ( , 2019] and Caupolicanini (now classified among the Diphaglossinae; see : Michener, 2007). ...
... This view follows the results of recently published molecular phylogenies of colletid bees (Almeida & Danforth, 2009;Almeida et al., 2012Almeida et al., , 2019. As treated herein, Colletinae corresponds to the tribe Colletini of various authors, such as Michener (1944Michener ( , 1989Michener ( , 2007, Moure & Urban (2002), Melo & Gonçalves (2005) and Moure et al. (2007Moure et al. ( , 2012. ...
... Currently, Colletinae contains 542 valid species, the generic placement of which has also been the subject of debate. Michener (1944) considered all species in Colletes without recognizing any of the subgenera that had previously been erected within the genus. Later, the same author also adopted Mourecotelles Toro & Cabezas, 1977, which was subdivided by him into subgenera to accommodate Hemicotelles Toro & Cabezas, 1977and Xanthocotelles Toro & Cabezas, 1978(Michener, 1989; see also : Ascher & Pickering, 2020). ...
... Stingless bees are a eusocial insect that lives together in a colony and has many important life roles. These bees are classified in Apidae, subfamily Apinae, and tribe Meliponini (Borror, D. Triplehorn, C., & Johnson 1992;Michener 1944;Trianto, M., Kaini, Saliyem, Warsih, E. 2020). Stingless bees include bees with a large number of genera, which is about 55 genera divided into 61 sub-genera with different morphological, morphometric, and nest structure characters for each species (Rasmussen 2008;Trianto, M., Kaini, Saliyem, Warsih, E. 2020). ...
... mm) (Efin et al. 2019). Meanwhile, when compared with the specimen by Michener, (1944) and Smith, (2012), who used specimen from Singapore (3.50 mm) and who used specimen from Sulawesi (3.40-3.43 mm) (Suriawanto et al. 2017), who used samples from Special Region of Yogyakarta (3.64-3.68 ...
... The nest structure described in this study, similar to the description of Karimah, (2017). Batumen is made from a mixture of wax and resin called cerumen (Michener, 1944). Batumen is a layer of cerumen that covers the entire surface of the nest. ...
Stingless bees is one group of eusocial insects living together in the hive. There are around 46 species of stingless bee in Indonesia with different morphological, morphometric and hive structure characteristics. This study aimed to describe the morphological, morphometric and beehives structure of Tetragonula laeviceps from Yogyakarta. Methods: Survey method is used by taking three sampling points of bamboo, house building and livestock crates in Bantul Regency. Sampling points determined by Purposive sampling method. An XSZ-107 BN binocular microscope analyzed samples of T. laeviceps with Optilab viewer and Image Raster software. Results: The results showed morphological characters of T. laeviceps are dominated shiny-black body, brownish-yellow antennas, klipeus on a head covered by fine silver hair, brownish-yellow mandible with two teeth, mesonotum in thorax covered by brownish to black hair, scutellum extended to propodeum, the ribbon of hair on the dorsal thorax is not very clear, and the hind tibia is rather hairy. Morphometric of T. laeviceps included body length between 3.44-3.76 mm, head width 1.55-1.70 mm, front wing length with tegula 3.76-4.37 mm, length of rear limbs tibia 1.37-1.57 mm, and the number of hamuli as many as 5. The beehive structure consisted of oval-shaped entrance formed funnel and varying internal hive in terms of the number of saplings, pollen cells, and honey cells. Conclusions: T. laeviceps have morphological, morphometric and hive structure characteristics that are different from other species and variated compared to similar species from other regions.
... The classification of Colletinae, one of the eight subfamilies currently recognized in the bee family Colletidae (see: Almeida et al., 2012Almeida et al., , 2019, has been the subject of a long-standing controversy amongst bee systematists. In an early paper on the phylogeny and classification of bees (Michener, 1944), Colletinae was considered in its broadest sense to include Paracolletini s.l. [= Paracolletes Smith, 1853 plus Callomelittinae, Neopasiphaeinae and Scrapterinae of Almeida et al. (2012Almeida et al. ( , 2019] and Caupolicanini (now classified among the Diphaglossinae; see : Michener, 2007). ...
... This view follows the results of recently published molecular phylogenies of colletid bees (Almeida & Danforth, 2009;Almeida et al., 2012Almeida et al., , 2019. As treated herein, Colletinae corresponds to the tribe Colletini of various authors, such as Michener (1944Michener ( , 1989Michener ( , 2007, Moure & Urban (2002), Melo & Gonçalves (2005) and Moure et al. (2007Moure et al. ( , 2012. ...
... Currently, Colletinae contains 542 valid species, the generic placement of which has also been the subject of debate. Michener (1944) considered all species in Colletes without recognizing any of the subgenera that had previously been erected within the genus. Later, the same author also adopted Mourecotelles Toro & Cabezas, 1977, which was subdivided by him into subgenera to accommodate Hemicotelles Toro & Cabezas, 1977and Xanthocotelles Toro & Cabezas, 1978(Michener, 1989; see also : Ascher & Pickering, 2020). ...
The bee subfamily Colletinae includes 542 species, the vast majority of which (518 spp.) belong to Colletes. The generic placement of the remaining 24 species has been controversial, resulting in several classifications being proposed. Despite several recently published molecular phylogenies of Colletinae, it remains unknown (1) what morphological synapomorphies support the recognized genera, (2) in which direction some relevant functional traits (e.g. basitibial plate) have evolved and (3) whether morphology supports the available molecular data. Herein, we provide a morphological phylogeny of Colletinae, which was constructed through parsimony analyses of 186 characters. In total, 50 ingroup species were included representing all major lineages of Colletes (29 spp.), plus all but three of the non-Colletes species of Colletinae (21 spp.). Trees were estimated through equal weights and extended implied weighting. Both provide strong support for the monophyly of Colletinae and indicate that the subfamily is defined by four unique synapomorphies. Our results also confirm recent phylogenetic hypotheses showing that Colletinae can be subdivided into two major clades: one comprising the reciprocally monophyletic Mourecotelles and Xanthocotelles; the other includes Colletes plus Hemicotelles, which are also reciprocally monophyletic. We also provide a fully illustrated key to facilitate generic identification of the Colletinae.
... It was Popov (1935) who first drew clearer attention to the affinities between Tarsalia and Ancyla, and documented the peculiar asymmetry in the male seventh metasomal sternum of the former. In his world clas¬ sification of bees, Michener (1944) brought Tarsalia and Ancyla together formally into a tribe Ancylaini4 (Michener, 1944), although the characters purportedly shared between these genera were problematic and applied only to one or the other of the two (Baker, 1998). Michener (1944) did not have material from which to base his character assessments, relying on generally inadequate published descriptions, and even excluded the tribe from his final phylogenetic scheme due to the dearth of substantive information. ...
... It was Popov (1935) who first drew clearer attention to the affinities between Tarsalia and Ancyla, and documented the peculiar asymmetry in the male seventh metasomal sternum of the former. In his world clas¬ sification of bees, Michener (1944) brought Tarsalia and Ancyla together formally into a tribe Ancylaini4 (Michener, 1944), although the characters purportedly shared between these genera were problematic and applied only to one or the other of the two (Baker, 1998). Michener (1944) did not have material from which to base his character assessments, relying on generally inadequate published descriptions, and even excluded the tribe from his final phylogenetic scheme due to the dearth of substantive information. ...
... Michener (1944) did not have material from which to base his character assessments, relying on generally inadequate published descriptions, and even excluded the tribe from his final phylogenetic scheme due to the dearth of substantive information. Popov (1949) subsequently suggested a relationship between Ancylaini and Exomalopsini (echoing the notion of Cockerell, 1933), as 4 The tribe was originally established as Ancylini (Michener, 1944), but was emended to Ancylaini to remove homonymy with a family group among the Gastropoda (Engel et al., 2008;ICZN, 2010). Baker, 1998). ...
... Among the historical achievements that epitomize the advancements in the field of bee comparative morphology, two crucial events in the last century are worth remarking. The first was the publication of a monograph entitled "Comparative External Morphology, Phylogeny, and a Classification of the Bees" (Michener, 1944), which laid the ground for a higher-level classification of bees. About five decades later, two sequential works proved crucial to systematize bee morphological knowledge in a cladistic perspective: Roig-Alsina and Michener (1993), and Alexander and Michener (1995). ...
... The pharyngeal plate of a bee was first described in detail in the works of Snodgrass (1910Snodgrass ( , 1942, for Apis mellifera Linnaeus, 1758 (Apidae: Apini). Other works with detailed morphological treatments of single species of bees that also mention the pharyngeal plate include that of Michener (1944) for Anthophora edwardsii Cresson, 1878 (Apidae: Anthophorini), Camargo et al. (1967) for Melipona marginata Lepeletier, 1836 (Apidae: Meliponini) and Eickwort (1969) for Pseudaugochlora graminea (Fabricius, 1804) (Halictidae: Augochlorini). Characters incorporating morphological variation of the pharyngeal plate in comparative contexts are presented in the datasets of the works of Prentice (1991), Packer (2008), Praz and Packer (2014), Porto et al. (2016b), and Packer et al. (2017) dealing with particular taxa of bees. ...
The pharyngeal plate is a morphological complex with extensive anatomical variation among bees and, therefore, potential as a source of phylogenetic information. The pharyngeal plate of bees is divided into four morphologically distinct regions: sitophore, hypopharyngeal lobe, pharyngeal rods, and median oral plate. In this work we illustrate and document in detail for the first time the pharyngeal plate of 43 bee species, providing descriptions of the morphological variation and contrasting these findings with representatives of apoid wasps (Crabronidae and Sphecidae). We evaluate and discuss the potential of this structure as a rich source of morphological information in the context of bee phylogeny and any research potentially impacted by comparative morphological data. The shape of the hypopharyngeal lobe is highly variable among suprageneric taxa of bees and can be readily employed to characterise taxa at various levels. We argue that the global patterns in the variation of the pharyngeal plate can provide information for phylogenetic inference within bees and constructed and coded 10 characters that encompass the most noticeable morphological differences discussed herein.
... Tagma terminology follows Michener (1944): the mesosoma and metasoma refer to the apparent thorax and abdomen; the mesosoma includes the thorax plus the first true abdominal segment, referred to as the propodeum. The mesosternal area (= 'mesosternum' of Mutillidae researchers) refers to the ventral surfaces of the male mesopleura that are sometimes bilaterally armed with a tubercle, denticle, spine, carina(e), and/or process. ...
The exclusively New World velvet ant genus Timulla Ashmead, 1899 (Hymenoptera: Mutillidae: Mutillinae: Trogaspidiini) contains 180 species and ten subspecies. Most of these species are known from a single sex, and the validity of the subspecies has not been evaluated since their original description in 1937 and 1938. The Timulla fauna of the United States of America includes thirty species and eight subspecies, and the fauna of Canada includes three species. The faunas of these two countries were critically studied with the following results. Out of the eight total subspecies, seven were found to be structurally identical to
and sympatric with the nominate subspecies. The subspecific differences were limited to cuticle and/or setal coloration in males and intergrades between them were found in several cases. With the senior synonym listed first, seven subspecies-level synonymies are proposed, which include: Timulla barbigera (Bradley, 1916) = T. barbigera rohweri Mickel, 1937, new synonym; T. dubitata (Smith, 1855) = T. dubitata fugitiva Mickel, 1937, new synonym; T. hollensis (Melander, 1903) = T. hollensis melanderi Mickel, 1937, new synonym; T. ocellaria Mickel, 1937 = T. ocellaria rufidorsa Mickel, 1937, new synonym; T. suspensa (Gerstaecker, 1874) = T. suspensa jonesi Mickel, 1937, new synonym, = T. suspensa sonora Mickel, 1937, new synonym; T. vagans (Fabricius, 1798) = T. vagans rufinota Mickel, 1937, new synonym. The final remaining subspecies, Timulla navasota coahuila Krombein, 1951, is raised to a full species, Timulla coahuila Krombein, 1951, new status, based on its unique female morphology. Also, four new sex associations are proposed, which include: Timulla barbata (Fox, 1899) = T. wileyae Mickel, 1937, new synonym; T. euterpe (Blake, 1879) = T. compressicornis Mickel, 1937, new synonym; T. neobule Mickel, 1937 = T. nicholi Mickel, 1937, new synonym; T. subhyalina Mickel, 1937 = T. dubitatiformis Mickel, 1937, new synonym. The former species, T. dubitatiformis, which was previously known only from females, is here recognized as being a morphologically-conservative complex of species; its synonymy with T. subhyalina effectively associates the remaining male-based members of the Timulla ocellaria species-group with it as well, which includes Timulla hollensis (Melander, 1903), T. kansana Mickel, 1937, T. ocellaria Mickel, 1937, T. rufosignata (Bradley, 1916), T. sayi (Blake, 1871), T. subhyalina Mickel, 1937, and T. tolerata Mickel, 1937. Further, two species-level synonymies are proposed, which include: Timulla dubitata (Smith, 1855) = T. murcia Mickel, 1938, new synonym; T. vagans (Fabricius, 1798) = T. huntleyensis Mickel, 1937, new synonym. Finally, Timulla cyllene (Cameron, 1894) is newly recorded in the United States of America from the state of Arizona.
... Morphological terms and abbreviations used in this paper mainly follow Michener (1944Michener ( , 2007. AN -antenna, BL -body length, coll. ...
Andrena (Chlorandrena) harisi n. sp. from Libya is described and illustrated. Characters for separation of this new species from similar Andrena (Chlorandrena) pyrrhula Pérez, 1895 are provided.
... A defining characteristic of the genus is the weakened distal veins of the forewing, which is most evident in females (Danforth & Ji 2001;Michener 2007). The broad scope of Lasioglossum is due, in part, to the challenges in diagnosing taxa at lower levels (Ebmer 1969(Ebmer , 2002Gibbs et al. 2012b;Michener 1944Michener , 2007. The genus has among the highest diversification rates of all bees (Bossert et al. 2021;Cardinal 2018;Grab et al. 2019) and this has doubtlessly contributed to the well-documented taxonomic challenges of the group (Gibbs 2018b). ...
A problematic species complex within Lasioglossum subgenus Sphecodogastra with unusual metallic reflections on the mesosoma is described from North America. Three new species in this complex are described and illustrated: Lasioglossum (Sphecodogastra) iridescens sp. nov., Lasioglossum (Sphecodogastra) dilisena sp. nov., and Lasioglossum (Sphecodogastra) silveirai sp. nov. Our study addresses the challenge of diagnosing Lasioglossum subgenera within North America. We present an updated key for the North American Lasioglossum subgenera.
... Morphological terminology follows that of Michener (1944Michener ( , 2007 and Engel (2001). The density of integumental punctures is described using the following formula: puncture diameter (in μm) / ratio of distance between punctures to average puncture diameter, e.g. ...
Available information about bees of the genus Epeolus in Central Asia is summarized. Twenty species are currently known from this area. Two new species are described: E. albus Astafurova & Proshchalykin, sp. nov. and E. pesenkoi Astafurova, sp. nov. Two species are newly recorded from Central Asia: E. asiaticus Astafurova & Proshchalykin, 2022 and E. nudiventris Bischoff, 1930. The hitherto unknown male of E. mikhailovi Astafurova & Proshchalykin, 2021 is described, and lectotypes are designated for E. ruficornis Morawitz, 1875 and E. vinogradovi Popov, 1952.
... Greatly influenced by his parents, Michener admired the nature (Engel, 2016a). During his adolescence he started studying some of the insects he collected in California with his family, making draws and keys that would later appear in his dissertation and seminal monograph on the comparative morphology of bees (Michener, 1944). Michener admired one of the most proliferous naturalists of his time, Theodore Dru Alison Cockerell (1866Cockerell ( -1948 and in 1932, he wrote him asking for help with the identification of some bees (Engel, 2016a). ...
In this paper are presented notes on the primary types of some species of the oil-collecting bees of the genus Centris Fabricius, 1804 described by Alpheus Packard, Arturo Roig-Alsina, Charles Michener, Flamínio Ruiz, Haroldo Toro, James Crawford, Jesus Santiago Moure, Philip Timberlake, and Roy Snelling. Information on the type status, type locality and depository are provided.
... The taxonomy and synonymy of species generally follow those of Bogusch and Hadrava (2018), except we regard E. julliani and E. transitorius as separate species, following Le Divelec (2021). Morphological terminology follows that of Michener (1944Michener ( , 2007 and Engel (2001). The density of integumental punctures is described using the following formula: puncture diameter (in μm) / ratio of distance between punctures to average puncture diameter, e.g., 15-20 μm / 0.5-1.5. ...
The nine species of the Epeolus julliani species group from the Palaearctic region are reviewed. Two new species are described and illustrated: Epeolus rasmonti Astafurova & Proshchalykin, sp. nov. (Russia, Mongolia, China) and E. kyzylkumicus Astafurova, sp. nov. (Central Asia). Epeolus julliani Pérez, 1884 and E. laticauda Bischoff, 1930 are newly recorded from Kazakhstan and E. seraxensis Radoszkowski, 1893 is newly recorded from Kazakhstan and Tajikistan. An identification key for both sexes of all members of this species group is presented.
... Anthophorinae has a long and complicated taxonomic past, and the multitude of name changes and divisions of this group is covered in detail by Brooks (1988a). In one of the first tree-based, morphological studies on bees, Michener (1944) recovered Anthophorini sister to Hemisiini (Centridini) and with that clade closely related to Melectini, Emphorini, Eucerini, and Rhathymini + Ericrocini (Ericrocidini). Subsequently, Michener (1974) suggested Anthophorini was more proximal to Eucerini and Centridini (including some cleptoparasites), with Brooks (1988b) suggesting the latter without formal investigation. ...
Despite recent advances in phylogenomics, the early evolution of the largest bee family, Apidae, remains uncertain, hindering efforts to understand the history of Apidae and establish a robust comparative framework. Confirming the position of Anthophorinae—a diverse, globally distributed lineage of apid bees—has been particularly problematic, with the subfamily recovered in various conflicting positions, including as sister to all other Apidae or to the cleptoparasitic Nomadinae. We aimed to resolve relationships in Apidae and Anthophorinae by combining dense taxon sampling, with rigorous phylogenomic analysis of a dataset consisting of ultraconserved elements (UCEs) acquired from multiple sources, including low-coverage genomes. Across a diverse set of analyses, including both concatenation and species tree approaches, and numerous permutations designed to account for systematic biases, Anthophorinae was consistently recovered as the sister group to all remaining Apidae, with Nomadinae sister to (Apinae, [Xylocopinae, Eucerinae]). However, several alternative support metrics (concordance factors, quartet sampling, and gene genealogy interrogation) indicate that this result should be treated with caution. Within Anthophorinae, all genera were recovered as monophyletic, following synonymization of Varthemapistra with Habrophorula. Our results demonstrate the value of dense taxon sampling in bee phylogenomics research and how implementing diverse analytical strategies is important for fully evaluating results at difficult nodes.
... Anthophorinae has a long and complicated taxonomic past, and the multitude of name changes and divisions of this group is covered in detail by Brooks (1988a). In one of the first tree-based, morphological studies on bees, Michener (1944) recovered Anthophorini sister to Hemisiini (Centridini) and with that clade closely related to Melectini, Emphorini, Eucerini, and Rhathymini + Ericrocini (Ericrocidini). Subsequently, Michener (1974) suggested Anthophorini was more proximal to Eucerini and Centridini (including some cleptoparasites), with Brooks (1988b) suggesting the latter without formal investigation. ...
Despite recent advances in phylogenomics, the early evolution of the largest bee family, Apidae, remains uncertain, hindering efforts to understand the history of Apidae and establish a robust comparative framework. Confirming the position of Anthophorinae—a diverse, globally distributed lineage of apid bees—has been particularly problematic, with the subfamily recovered in various conflicting positions, including as sister to all other Apidae or to the cleptoparasitic Nomadinae. We aimed to resolve relationships in Apidae and Anthophorinae by combining dense taxon sampling, with rigorous phylogenomic analysis of a dataset consisting of ultraconserved elements (UCEs) acquired from multiple sources, including low-coverage genomes. Across a diverse set of analyses, including both concatenation and species tree approaches, and numerous permutations designed to account for systematic biases, Anthophorinae was consistently recovered as the sister group to all remaining Apidae, with Nomadinae sister to (Apinae, [Xylocopinae, Eucerinae]). However, several alternative support metrics (concordance factors, quartet sampling, and gene genealogy interrogation) indicate that this result should be treated with caution. Within Anthophorinae, all genera were recovered as monophyletic, following synonymization of Varthemapistra with Habrophorula. Our results demonstrate the value of dense taxon sampling in bee phylogenomics research and how implementing diverse analytical strategies is important for fully evaluating results at difficult nodes.
... Recent phylogenetic results based on molecular data (Freitas et al., 2021;Praz & Packer, 2014) supported the placement of Ancylaini as the sister group of Eucerini, as previously proposed (Michener, 1944;Silveira, 1993). These results suggested that Eucerinae colonized the Old World (OW) through North Atlantic land bridges during the Eocene (Praz & Packer, 2014), explaining the distribution of Ancylaini as an endemic Palaearctic taxon. ...
Aim
An antitropical pattern is characterized by the occurrence of closely related taxa south and north of the tropics but absent or uncommonly represented closer to the equator, in contrast to most taxa, which tend to have their highest diversity in the tropical regions. We investigate the antitropical distribution of eucerine bees with the aim of contributing to the characterization and understanding of this pattern.
Location
All continents except Antarctica and Australia.
Taxon
Eucerine bees (Hymenoptera: Apidae: Eucerinae).
Methods
We carried out phylogenomic dating under two different clock models and used multiple strategies to vary matrix composition, evaluating the overlapping of divergence times estimated across models using Bhattacharyya coefficients. Lastly, we reconstructed the biogeographic history of eucerine bees using a Bayesian implementation of the DEC model.
Results
Eucerinae is estimated to have started diversifying during the Palaeocene, with all its tribes originating during the Palaeocene/Eocene transition in southern South America. At least two range expansions happened into North America before the full closure of the Isthmus of Panama. We show that divergence between closely related groups with disjunct distributions would have happened in periods when the climate favoured the expansion of open habitats and became isolated when the forests were re‐established.
Main conclusions
We describe the early diversification of eucerine bees, revealing an intimate association with southern South America. Events of range evolution of Eucerinae were likely affected by periods of global cooling and aridification, and palaeoclimatic and vegetational conditions probably have been more relevant to the formation of the antitropical distribution of Eucerinae than the consolidation of the Isthmus of Panama connecting the Americas. We also demonstrate that most uncertainty in divergence time estimation is not due to the amount of molecular data being used, but more likely other factors like fossil calibrations and violations of clock models.
... The internal characters have been used in bee systematics since Michener (1944) with subsequent efforts by many authors (e.g. Roig-Alsina & Michener, 1993;Alexander & Michener, 1995;Packer, 2003). ...
Augochlorini comprise 646 described bee species primarily distributed in the Neotropical region. According to molecular and morphological phylogenies, the tribe is monophyletic and subdivided into seven genus groups. Our main objective is to propose a revised phylogeny of Augochlorini based on a comprehensive data set including fossil species as terminals and new characters from the internal skeleton. We also aim to develop a total-evidence framework incorporating a morphological-partitioned homoplasy approach and molecular data and propose a detailed biogeographic and evolutionary scenario based on ancestor range estimation. Our results recovered Augochlorini and most genus groups as monophyletic, despite some uncertainties about monophyly of the Megalopta and Neocorynura groups. The position of the cleptoparasite Temonosoma is still uncertain. All analyses recovered Augochloropsis s.l. as related to the Megaloptidia group. Internal characters from the head, mesosoma and sting apparatus provided important synapomorphies for most internal nodes, genus groups and genera. Augochlorini diversification occurred in the uplands of the Neotropical region, especially the Brazilian Plateau. Multiple dispersals to Amazonia, Central America and North America with returns to the Atlantic endemism area were recovered in our analysis. Total evidence, including morphological partitioning, was shown to be a reliable approach for phylogenetic reconstruction.
... In the past, the phylogenetic relationships of Scrapter to other colletid lineages proved difficult to establish based on morphology alone (McGinley 1981;Alexander and Michener 1995;Plant and Paulus 2016). Previously considered to be part of Paracolletini (e.g., Michener 1944), albeit without strong morphological evidence (McGinley 1981), analyses of nucleotide sequence data strongly indicated a sister group relationship of Scraptrinae and Euryglossinae, an Australian-endemic lineage of Colletidae (Almeida and Danforth 2009;Almeida et al. 2012;Kayaalp et al. 2017;Cardinal et al. 2018). Interestingly, a close relationship to Euryglossinae has been discussed as early as 1933 (Cockerell and Ireland 1933; specifically discussing Euryglossidia Cockerell) and a sister-group relationship is supported by certain morphological characters of the mature larvae (McGinley 1981). ...
Scrapter is a genus of colletid bees with a primary distribution centered in Southern Africa. The genus currently comprises 68 recognized species, which are divided into nine species groups, ranging from one to 29 included species. The Scrapter heterodoxus group is presently considered to be the only monotypic group, because of synonymization of Scrapter heterodoxus with Scrapter peringueyi in a previous revision of the genus. A comparative examination of these two species using both morphological assessment and molecular sequence data from the COI barcode region supported the recognition of S. peringueyi as a valid species, which we accordingly resurrect as the second species of the Scrapter heterodoxus species group. We provide high resolution images of the type specimens for both species and updated diagnoses to enable their separation from all other species of Scrapter .
... The galeal lamina with the galeal comb is also found in other groups of Vespidae (Duncan, 1939) and in other Hymenoptera (e.g. Ulrich, 1924;Michener, 1944;Prentice, 1998;Hymenoptera Anatomy Consortium, 2020: HAO:0002136), which is indicating that it is a plesiomorphic structure in Masarinae. In adult masarine wasps, pollen grains are commonly found on the galeal comb (Krenn et al., 2002), indicating that it has an essential function in the collection of pollen. ...
While many pollen wasps nest in hard clayey soil or in rigid sand or use these kinds of substrates to build aerial earthen cells, all representatives of the genus Quartinia, in which nesting behavior has been studied so far, construct their nests in habitats with loose sand. The walls of the burrow are stabilized by a silky excretion that is applied on their inner surfaces during nest construction. The lining-behavior of Quartinia females is described. Representatives of the genus Quartinia and other pollen wasp taxa have been studied comparatively using histology, microcomputed tomography and scanning electron microscopy for identifying the structures correlated with lining-behavior. Thus, we can document a change in the morphology of the maxillary gland for females of the genus Quartinia, as well as the existence of a process of the galea which likely serves the production of the silky threads. The fact that these modifications are missing in males is corroborating their function in brood care. Two possible ways of silk thread production are discussed. The newly discovered structures are key adaptations of the genus Quartinia that enabled the inhabitation of new habitats and thus probably facilitated the species diversification of this genus.
... Emphorini can be distinguished from other groups by the convex vertex, with ocelli below the margin of the head and short antennae in males (Michener 2007). in eastern north america, Ptilothrix can be identified using the keys to family and genus in Mitchell (1960Mitchell ( , 1962. Specific characters include lack of arolia between tarsal claws; elongate first flagellomere, which is two or more times as long as its apical width; finely plumose and elongate hind tibial scopal hairs; both sexes with black clypeus; marginal cell acute apically and bent away from costal margin slightly; and three submarginal cells, with the second shorter than the first or third (Michener 1944;Mitchell 1962). Ptilothrix bombiformis are moderately large bees (body lengths: ♀ 13-17.5 mm, ♂ 12.5-17.5 ...
... The term antennal rim is used for the convex section of cuticle that surrounds the antennal foramen. Tagma terminology follows Michener (1944): the mesosoma and metasoma refer to the apparent thorax and abdomen; the mesosoma includes the thorax plus the first true abdominal segment, referred to as the propodeum. The scutellar area as defined by Bartholomay et al. (2018) is the region apparently composed of both mesoscutellar and metascutellar tissue that is found between the propodeal spiracles on the mesosomal dorsum in females; the scutellar scale is a transverse carina or lamella found anteromedially in this scutellar area in females. ...
The velvet ant genus Invreiella Suárez, 1966 is redescribed and revised. Females are grouped into five species-groups, with eleven new species described based on females: I. acuminata Waldren, sp. nov., I. australis Waldren, sp. nov., I. bimaculata Waldren, sp. nov., I. breviclypeata Waldren, sp. nov., I. chihuahuensis Waldren, sp. nov., I. cuernavaca Waldren, sp. nov., I. erythrocephala Waldren, sp. nov., I. manleyi Waldren, sp. nov., I. mesomexicana Waldren, sp. nov., I. suarezi Waldren, sp. nov., and I. tequila Waldren, sp. nov. One new combination is included, I. cephalargia (Mickel, 1924), comb. nov., formerly placed in Pseudomethoca Ashmead, 1896. Additionally, I. curoei Quintero & Cambra, 2011 is synonymized with I. cardinalis (Gerstaecker, 1874), syn. nov., and I. megacantha (Cockerell & Casad, 1894), stat. resurr., is treated as a nomen dubium. The lectotype of Mutilla satrapa Gerstaecker, 1874 originally designated by C. Mickel is here validated. An illustrated key to species is included.
... The tropical New World bee genus Rhathymus Lepeletier & Serville, part of the tribe Rhathymini, consists of cleptoparasitic, medium to large sized bees, most species with yellow or black integument, and wasp-like habitus. Within the Apidae, morphology based phylogenetic analyses have placed the Rhathymini in the subfamily Apinae (Michener 1944;Roig-Alsina & Michener 1993), while a relatively recent molecular based analysis places them within a larger cleptoparistic clade along with the Nomadinae (Cardinal et al. 2010). The genera of Rhathymini have been fully characterized by Engel et al. (2004aEngel et al. ( , 2004b, including a key to the two included genera, Rhathymus Engel et al. 2004a andNanorhathymus Engel et al. 2004b, classification scheme adopted by Michener (2007) and followed here. ...
A new species of the bee genus Rhathymus Lepeletier & Serville, 1828 is described and figured as Rhathymus atitlanicus Ayala, Hinojosa-Díaz and Armas-Quiñonez. The new species is from the department of Sololá, Guatemala, and can be distinguished based on the black integument contrasting with the orange color of the wings.
... Particularly if bees have behavioral or morphological adaptations to certain floral or pollen morphologies, this could help explain broad-scale evolutionary patterns of floral host use in bees. This pattern has been repeatedly noted in bee species groups adapted to large or coarse pollen such as in the genera Diadasia, Macrotera, and Eucera (Michener 1944;Linsley and MacSwain 1958;Danforth 1996;Dorchin et al. 2018). ...
Pollen is the primary protein and nutrient source for bees and they employ many different behaviors to gather it. Numerous terms have been coined to describe pollen gathering behaviors, creating confusion as many are not clearly-defined or overlap with existing terms. There is a need for a clear yet flexible classification that enables accurate, succinct descriptions of pollen gathering behaviors to enable meaningful discussion and comparison. Here, we classify the different pollen gathering behaviors into two main classes: active and incidental pollen collection. Active pollen collection is subdivided into six behaviors: scraping with the extremities, buzzing, rubbing with the body and/or scopae, rubbing with the face, tapping, and rasping. In addition to the active and incidental pollen gathering behaviors, many bees have an intermediate step in which they temporarily accumulate pollen on a discrete patch of specialized hairs. Each behavior is described and illustrated with video examples. Many of these behaviors can be further broken down based on the variations found in different bee species. Different species or individual bees mix and match these pollen collecting behaviors depending on their behavioral plasticity and host plant morphology. Taken together, the different behaviors are combined to create complex behavioral repertoires built on a foundation of simple and basic steps. This classification sets the groundwork for further research on various topics, including behavioral plasticity in different species, comparisons between generalists and specialists, and the relative effectiveness of different pollen gathering behaviors.
... Explanation of homology analysis. Because much of the confusion over endopterygote genital homologies stems from apparent discord between developmental theory (Snodgrass, 1935a(Snodgrass, , 1941(Snodgrass, , 1957 and skeletal observations of the Hymenoptera (Crampton, 1920(Crampton, , 1938Michener, 1944a), it is necessary to provide a detailed explanation of the hypothesis developed and proposed here. The first step in this dialectic is to recognize the historical terminology for hymenopteran genitalia. ...
No consensus exists for the homology and terminology of the male genitalia of the Hexapoda despite widespread acknowledgment of systematic value and well over a century of debate. Based on dissections and the literature, I compared genital skeletomusculature across the Hexapoda in contrast with the Remipedia, the closest pancrustacean outgroup. I found the pattern of origin and insertion for extrinsic and intrinsic genitalic and appendicular musculature to be consistent among the Ectognatha, Protura, and the Remipedia; from these consistencies, I have inferred a groundplan for the Hexapoda, and have provided an extended and explicitly diagrammed theory of genitalic homologies and evolution for all insect orders (Boudinot 2018). In the present talk, I will explain the fundamental concepts derived from this study, and I will illustrate via fine-grained transition series the evolutionary patterns leading to the Endopterygota and clades therein.
... Callomelitta is the only genus included in this subfamily; eleven species are recognized. Historically, it has been placed within the tribe Paracolletini (sensu Michener 1944), in the subfamily Colletinae (e.g., Michener 2000), or in Paracolletinae (e.g., Engel 2005). The distinctness of Callomelitta as compared to other members of Colletidae has been discussed by several authors (e.g., McGinley 1981;Michener 2000Michener , 2007Engel 2005), but a revised placement of the genus awaited the phylogenetic analysis of Colletidae. ...
Colletid bees are widely distributed, but most of their diversity is found in southern continents, particularly Australia and South America. A major obstacle for the classification of Colletidae has been the lack of a well-resolved phylogenetic hypothesis encompassing representatives from all the main lineages of this family. A molecular phylogenetic study recently completed shed light on the subfamilial delimitation and relationships within Colletidae, and included a thorough enough sampling of Paracolletinae that genus-level classification could be evaluated. In this article a revised genus-level classification is presented for the Australian Paracolletinae, an attempt to make it more congruent with the phylogenetic relationships, as well as consistent with the classification of the South American paracolletine generic classification. According to the classification proposal presented here, Paracolletinae comprise 425 species, 298 of them endemic to the Australian Region. All subgenera of Leioproctus and of Paracolletes are elevated to the level of genus, thus raising the number genera occurring in the Australian Region to 25. According to a recent phylogenetic analysis of Colletidae, the Australian genus Callomelitta does not form a monophyletic group with the remaining genera of Paracolletinae. Callomelitta is hence removed from Paracolletinae and placed on a subfamily of its own, Callomelittinae, subfamily nov. Additionally, a new species of the Australian Paracolletinae genus Andrenopsis Cockerell is described as Andrenopsis michenerianus, sp.n..
... Morphological terms are from Brothers (2018) and the Hymenoptera Anatomy Consortium (2018); with details on sculpture from Harris (1979) and prosternum from Michener (1944). The abbreviations F, S and T refer to numbered flagellomeres, metasomal sterna and metasomal terga, respectively; IOD = the smallest distance between a lateral ocellus and the median ocellus, OOD = the smallest distance between the eye margin and a lateral ocellus and POD = the smallest distance between lateral ocelli. ...
A new species of Limaytilla from Argentina is described and illustrated based on males, L. diaguita Torréns & Fidalgo sp. nov.. Diagnoses for both sexes of the genus and an illustrated key for the Argentinian species are provided as well as new record data for L. pampa Casal and L. pehuenche Casal. Habitus photographs of all Argentinian species and L. orlandoi Cambra, Quintero & Pagliano from Chile, are also provided.
... Terminology for structures follows Michener (1944Michener ( , 2000, except that metapostnotum is used instead of propodeal triangle (Brothers, 1976). The maximum diameter of the median ocellus (MOD) is used as a reference to express the length of the pubescence and other structures. ...
A new species, Hexantheda entrerriana, from the province of Entre Ríos, Argentina, is described. These bees have an unusual labial palp with eight segments. Features distinguishing the new species from the other two species of the genus are presented.
... Third, many bee taxa now widely accepted as genera today, were treated in the past as subgenera of much larger genera, just like in the case of Megachile s.l. For example, Michener (1944) and Schwarz (1948) treated the more than 20 genera of Meliponini recognized today as subgenera of Trigona. For these reasons, we follow the third proposal given its practical adavantages, its hierarchichal arragement, and congruence with modern generic concepts of bees. ...
A unique feature among bees is the ability of some species of Megachile s.l. to cut and process fresh leaves for nest construction. The presence of razors between the female mandibular teeth (interdental laminae) to facilitate leaf-cutting (LC) is a morphological novelty that might have triggered a subsequent diversification in this group. However, we have a limited understanding of the evolutionary origins of this behavior and associated structures. Herein, we use total-evidence tip-dating analyses to infer the origin of LC bees and patterns of variation of interdental laminae. Our datasets included five nuclear genes, representatives of all fossil taxa, 80% of the extant generic-level diversity of Megachilidae, and the full range of generic and subgeneric diversity of Megachilini. Our analyses support the notion of a recent origin of LC bees (15-25 Ma), casting doubts on Eocene trace fossils attributed to these bees. We demonstrate that interdental laminae developed asynchronicaly from two different structures in the mandible (teeth or fimbrial ridge), and differ in their phenotypic plasticity. Based on the phylogenetic results, we propose robust classificatory solutions to long-standing challenges in the systematics of Megachilidae. We discuss the implications of our findings as a foundational framework to develop novel evolutionary, ecological, and functional hypotheses on this behavior.
... No forrageio por pólen para aprovisionamento do ninho (Roubik 1992), as abelhas compactam os grãos em uma massa e a movem para estruturas especializadas localizadas nas pernas traseiras (escopa/corbícula) ou no metassoma (escopa ventral em alguns grupos) (Michener 1944, Buchmann 1983, Muth et al. 2017. O pólen compactado torna-se indisponível para polinização (Roubik 1992, Michener 2007, porém é possível que pólen residual fique aderido ao restante do corpo. ...
O estabelecimento das interações planta-polinizador é de grande importância para o funcionamento dos ecossistemas. Estas interações podem ser registradas diretamente, por observação focal, ou indiretamente, pela carga polínica presente no corpo do polinizador. Assim, a avaliação da especialização e da dependência de parceiros mutualísticos, propriedades que podem ser estudadas por meio de redes complexas, depende do método empregado no registro das interações. O principal objetivo deste estudo foi comparar uma rede de visitação floral e uma rede de transporte de pólen construídas a partir de interações entre plantas e abelhas amostradas em 16 parcelas em estágios iniciais de sucessão ecológica na Floresta Atlântica, em Antonina, Paraná, sul do Brasil. O registro das interações foi realizado inicialmente por observação focal, e posteriormente foram amostrados os grãos de pólen aderidos ao corpo de seis espécies de abelha, ficando as duas redes restritas às interações realizadas por estas seis espécies. A rede de transporte de pólen foi subdividida em uma rede de alimentação, utilizando apenas o pólen aderido às corbículas ou escopas das abelhas, e uma rede de polinização, com o pólen presente no corpo, exceto pernas, das abelhas. Estas redes foram comparadas em relação ao grau ( k ), ao índice de especialização ( d’ ) e à força das espécies ( F ), por meio de testes t pareados. A rede de transporte de pólen apresentou maior número de interações, maior k , menor d’ e maior número de espécies de planta que a rede de visitação floral. Não houve variação da F. As redes de alimentação e de polinização apresentaram propriedades semelhantes. A análise polínica revelou que as abelhas tendem a realizar turnos de forrageio exclusivos em flores de pólen. As interações adicionadas a partir da análise polínica resultaram em diferenças nas métricas avaliadas e complementaram o registro das interações observadas na comunidade. Portanto, a informação associada de ambos os tipos de redes pode ampliar a compreensão das interações, aproximando-se, dessa forma, à descrição da rede real. FLORAL VISITATION NETWORK VERSUS POLLEN-TRANSPORT NETWORK BETWEEN BEES AND FLOWERS IN THE ATLANTIC FOREST IN SOUTH BRAZIL . The establishment of plant-pollinator interactions is of great importance for the functioning of ecosystems. These interactions can be recorded directly, by focal observation, or indirectly, by the pollen load on the pollinator body. Thus, the evaluation of the specialization and dependence of mutualistic partners, properties that can be assessed through complex networks, depends on the method used for interaction record. The main objective of this study was to compare a floral-visitation network and a pollen-transport network constructed from interactions between plants and bees sampled in 16 plots in early successional areas in the Atlantic Forest, in Antonina, Paraná, southern Brazil. We initially recorded these interactions by focal observation and we sampled the pollen grains on the bees’ body of six bees, and the two networks were restricted to the interactions of these six species. We subdivided the pollen-transport network into a feeding-pollen network, using only the pollen adhered to the bees' corbicules or scopes, and a pollination network, with the pollen present on the bees’ body, except the legs. These networks were compared in relation to the degree ( k ), the specialization index ( d ') and species strength ( F ), through paired t -tests. The pollen-transport network showed a higher number of interactions, higher k , lower d ' and a greater number of plant species than the floral visitation network. There was no variation in F . The feeding- and pollination networks presented similar properties. Pollen analysis revealed that bees tend to perform exclusive foraging flights on pollen flowers. The interactions added from the pollen analysis resulted in differences in the metrics evaluated and complemented the record of the interactions observed in the community. Therefore, the associated information of both types of networks can broaden the understanding of the interactions, thus approaching the actual network description.
... Explanation of homology analysis. Because much of the confusion over endopterygote genital homologies stems from apparent discord between developmental theory (Snodgrass, 1935a(Snodgrass, , 1941(Snodgrass, , 1957 and skeletal observations of the Hymenoptera (Crampton, 1920(Crampton, , 1938Michener, 1944a), it is necessary to provide a detailed explanation of the hypothesis developed and proposed here. The first step in this dialectic is to recognize the historical terminology for hymenopteran genitalia. ...
No consensus exists for the homology and terminology of the male genitalia of the Hexapoda despite over a century of debate. Based on dissections and the literature, genital skeletomusculature was compared across the Hexapoda and contrasted with the Remipedia, the closest pancrustacean outgroup. The pattern of origin and insertion for extrinsic and intrinsic genitalic musculature was found to be consistent among the Ectognatha, Protura, and the Remipedia, allowing for the inference of homologies given recent phylogenomic studies. The penis of the Hexapoda is inferred to be derived from medially-fused primary gonopods (gonopore-bearing limbs), while the genitalia of the Ectognatha are inferred to include both the tenth-segmental penis and the ninth-segmental secondary gonopods, similar to the genitalia of female insects which comprise gonopods of the eighth and ninth segments. A new nomenclatural system for hexapodan genitalic musculature is presented and applied, and a general list of anatomical concepts is provided. Novel and refined homologies are proposed for all hexapodan orders, and a series of groundplans are postulated. Emphasis is placed on the Endopterygota, for which fine-grained transition series are hypothesized given observed skeletomuscular correspondences.
... Treatments of the male genitalia of Hymenoptera have employed an exceedingly large and confusing array of terms, in part because of differing theories of male genitalic origin in insects (Michener 1944(Michener , 1956Snodgrass 1941Snodgrass , 1957, but also because of varying patterns of usage (Yoder et al. 2010). In this case, we choose Snodgrass' terms because they have been used often for Chalcidoidea (Bouček 1952;Dominichini 1953;Gibson 1997;Pinto 1992). ...
Eucharitidae (Hymenoptera) are specialized ant (Formicidae) parasitoids. As we begin to develop a better understanding of their phylogenetic relationships, it is critical to establish baselines for morphological and biological data. A morphological review and the first report of life history data for Psilocharis afra Heraty is provided based on new material from the Mpumalanga Province of South Africa. Psilocharis Heraty is included in Eucharitinae, but it is unclear whether it is the sister group of all other members of the subfamily, or sister group to Neolosbanus Girault in a monophyletic Psilochari-tini, which would in turn be sister group to Eucharitini. The oviposition habits of P. afra differ from those of other Eucha-ritidae in that eggs are placed among trichomes under bracts at flower bases, instead of either being inserted into cavities formed in plant tissue by an enlarged ovipositor (as in Oraseminae and some Neolosbanus) or inserted into cavities in plant tissue, as in most Eucharitini. The egg and first-instar planidia larva are described, and adult morphology is discussed with reference to Eucharitidae and other parasitoid Hymenoptera.
... Various classifications and many taxonomic studies and notes were made by Griswold (1994), Griswold andMichener (1988 and1997), Michener (1944Michener ( , 1965Michener ( , 1883Michener ( , 1996Michener ( , 1997Michener ( and 2000 and Gupta (1993). Complete phylogentic analysis for the long tongued bees was published by Roig-Alsina and Michener (1993). ...
... This explains, at least in part, the specificity of bees for a specific plant. Anchusa azurea attracted high numbers of long-tongued bees in the sub-family Anthophorinae (4 species) [43,44], a group characterised by very rapid foraging [45].This choice may be related to the composition of its nectar, rich in sucrose and amino acids, found in a tube formed by fused petals [46,47]. More Anthophorinae species are very active in the spring on this same plant species [48]. ...
This study considered the plant floral resources visited by native wild and honey bees in Algiers (northern Algeria). Three botanical families accounted for almost 2/3 of all visits: Asteraceae (44.1%), Boraginaceae (15.3%) and Brassicaceae (13%). Plants in other families were visitedless frequently (e.g.Ranunculaceae (0.1%)). At the species level, the most frequently visited plant was Anchusa azurea (Boraginaceae), reaching 15.2% of the total bee visitation. Our work showed that Apis mellifera Linnaeus, 1758 is, typically, a polylectic species and that the majority of solitary bees exhibited the more specialised trait of oligolectics. The narrowest trophic niche varied between 0.01 and 0.06 bits for Halictus rufipes (Fabricius, 1793), Halictus scabiosae (Rossi, 1790) (Halictidae), Osmia pinguis Pérez, 1895 and Osmia tricornis Linnaeus, 1811 (Megachilidae).
... Body length was measured in lateral view as the sum of distances from the antennal sockets to the posterior end of the propodeum and from the latter to the tip of the metasoma. The vein separating the first and second cubital cells of the forewing (cu-v in Michener 1944) is referred to here as the nervulus. The nervulus is described as antefurcal ( Fig. 32), interstitial, or postfurcal ( Fig. 31), if its anterior end is situated proximal to, in alignment with, or distal to the posterior end of the basal vein, respectively. ...
Andrena (Poecilandrena) Hedicke is a subgenus of small solitary bees, with the greatest diversity in the Eastern Mediterranean Region—an important but understudied biodiversity hotspot for bees. We studied Andrena (Poecilandrena) collected mostly in Israel and the West Bank, and make several additions to the regional fauna. We provide the first comprehensive review of Andrena (Poecilandrena) species currently known from Israel, the West Bank, Jordan, Lebanon and Syria, including diagnostic keys to females and males, descriptions of new species and unknown sexes, and detailed information for each taxon regarding distribution, phenology and flower visitation. Our review includes fourteen species of Andrena (Poecilandrena) from the Levant, including five species new to science: A. freidbergi Pisanty & Scheuchl n. sp., A. galilaea Pisanty & Scheuchl n. sp., A. hierosolymitana Pisanty & Scheuchl n. sp., A. sedumella Scheuchl & Pisanty n. sp., and A. stenofovea Scheuchl & Pisanty n. sp. We also report four species as new to the region, and provide the first description of the male of A. rusticola Warncke. We exclude A. arabica Scheuchl & Gusenleitner from the subgenus, and synonymize it with A. helouanensis Friese. We anticipate that more collecting work in this region will yield additional undescribed taxa as well as new records, especially of taxa already known from Turkey. In the appendix to this work we provide information on 35 new records of other Andrena subgenera from Israel and the West Bank that have accumulated in recent years.
The Stelidium group is readily distinguished from all other members of the subgenus Stelis Panzer, 1806 by the combination of small body size (≤ 6 mm), pale maculations on the head adjacent to the inner margins of the compound eyes and laterally on the vertex in both sexes, and females with sternum 6 extended beyond tergum 6, the former with the dorsal lip trowel-shaped with the apex broadly rounded or subtruncate to more narrowly pointed. This monophyletic clade, which is endemic to North America, currently consists of members previously placed into two species groups: the permaculata group containing S. anasazi Parker & Griswold, 2013, S. ashmeadiellae Timberlake, 1941, S. permaculata Cockerell, 1898, and S. robertsoni Timberlake, 1941, and the palmarum group containing S. broemelingi Parker & Griswold, 2013, S. elongativentris Parker, 1987, and S. palmarum Timberlake, 1941; two additional species, S. herberti (Cockerell, 1916) from Mexico, and S. nyssonoides (Brues, 1903) from Texas, United States, have not been definitively placed in either species group. Two new species are herein described, one from southcentral British Columbia, Canada, the other from New Mexico, United States. A preliminary molecular phylogeny places both new species in the permaculata species group. In addition, S. herberti is also placed within the permaculata species group based on morphological similarity, sharing the multi-spotted maculation pattern on the terga. Based on molecular affinity, S. broemelingi also belongs to the permaculata species group. Because no type specimen for S. nyssonoides is seemingly available for examination, it is hereby considered nomen dubium until the specimen is found and its taxonomic status clarified in relation to the more recently described species in the permaculata species group. A key to females and diagnoses are provided for all known taxa.
Fernando A. Silveira had the unique combination of being a sagacious scientist and a remarkable human being. Throughout his career, he made significant contributions to understanding bee diversity and keenly spread this scientific information to the academic community at large and beyond the university walls. His rich character, warm heart, strong voice and laughter are missed by those privileged to be Fernando’s students, friends, mentors, and family. In this volume, we honor Fernando A. Silveira, who prematurely passed away at the age of 62, leaving three sons, his wife, and numerous friends.
The genus Ancyloscelis Latreille, 1829 (Hymenoptera: Apidae), a taxon restricted to the Neotropics and southern Nearctic (Michener 1942, 2000, 2007; Schaller and Roig-Alsina 2021; Melo 2022), has been difficult to place precisely within the higher classification of bees (Roig-Alsina and Michener 1993; Aguiar et al. 2019; Freitas et al. 2020), and even the genus name has a confusing history (see Michener 1942). Michener (1944) placed it together with Exomalopsis Spinola, 1853 within the Exomalopsini Vachal, 1909, and Michener and Moure (1957) later expanded this tribe to include ten additional genera, with Ancyloscelis the sole member of one of the five distinct sections they recognized (reviewed by Silveira 1993). Later, Jesus S. Moure (cited in Roig-Alsina and Michener 1993) suggested that the placement of Ancyloscelis should be within Emphorini Robertson, 1904, a position supported in that work. However, Roig-Alsina and Michener (1993) concluded that it differed enough from other members to recognize two subtribes, proposing Ancyloscelina Roig-Alsina and Michener, 1993 containing only the type genus, with the remaining Emphorini recognized at that time (i.e., Diadasia Patton, 1879, Diadasina Moure, 1950, Melitoma Lepeletier and Serville, 1828, and Ptilothrix Smith, 1853) placed in subtribe Emphorina Robertson, 1904. Michener (2000, 2007) and others (Silveira et al. 2002, Rodríguez and Roig-Alsina 2004) continued to recognize Ancyloscelis within Emphorini, but subtribal classifications were not used in those works.
First record of a gynandromorph of Osmia submicans MORAWITZ, 1870 (Hymenoptera, Megachilidae)-characterisation by morphological and morphometric parameters and critical note on gynander classification Anselm KRATOCHWIL
A b s t r a c t : The first record of a gynandromorph of Osmia submicans MORAWITZ, 1870 (Hymenoptera, Megachilidae) with a nearly complete bilateral asymmetry is presented. However, several alternating asymmetries of female and male traits occur additionally on the tagma level. The article presents an analysis of 25 morphological (non-meristic) characteristics (integument colour, pubescence, structural features) and 34 morphometric parameters of females and males. The morphometric differences were tested statistically. The left and right gynander sides were compared with those of females and males (using multivariate analysis for parameter measurements). The gynanders within the genus Osmia described so far in the literature were typified again and compared with the gynander presented here. The conventional type classification of gynanders is critically discussed. Criteria that may be relevant for an analysis of gynanders are presented.
K e y w o r d s : Anthophila, gynander, gynandromorphism, morphometric differentiation, principal component analysis.
Trifolium alexandrinum L. اﻟﻧ وﺗﺣدﯾد اﻟﯾوﻣﻲ ﺷﺎط ﻟﻬﺎ ﻟﻠﻌﺎم ان ﯾر وﺣز وﻣﺎﯾس ﻧﯾﺳﺎن ﺷﻬور وﻟﺛﻼث ٢٠١٠ اﻟﻘﺎدﺳﯾﺔ ﻣﺣﺎﻓظﺔ ﻓﻲ. ات وﺣﺷر اﻟﺑري اﻟﻧﺣل ﻣن ﻧوﻋﺎ ﻋﺷر اﺣد ﺗﺳﺟﯾل ﺗم ﺣﯾث وﻫﻲ اﻟﻧﺣل ﻟﻌﺎﺋﻠﺔ ﺗﻌود اع اﻧو ﺑﻊ ار وﻫﻲ اﻟﺑرﺳﯾم ﻣﺣﺻول ازﻫﺎر ﻣن اﺧرى melifera Apis و Megachil sp و Halictus sp. ﻟﻠﻧوع ﺑﺎﻻﺿﺎﻓﺔ Anderena sp وﻫﻲ اﻟذﺑﺎب ﺗﺑﺔ ر اﻟﻰ ﺗﻌود اع أﻧو وﺛﻼث Metasyrphus taeniops و Eristalis aeneus scopoit و Episyrphus balteatus وﻫﻲ اﻻﺟﻧﺣﺔ ﺣرﺷﻔﯾﺔ ﺗﺑﺔ ر اﻟﻰ ﺗﻌود اع اﻧو ﺑﻊ ار و Lycaeides Melissa(alpime) و Autograph gamma و Colias croceus(L.) اﻟﻧوع ا اﺧﯾر و Artogeia rapa. اﻟﻌددﯾﺔ اﻟﻛﺛﺎﻓﺎت اﺧﺗﻼف اﻟﻰ اﻟﻧﺗﺎﺋﺞ اﺷﺎرت اﻟﺣﺷري اﻟﻧوع اﺧﺗﻼف وﻛذﻟك اﻟﯾوم ﻣن اﻟوﻗت ﺑﺎﺧﺗﻼف ات ﻟﻠﺣﺷر ﻣﺎ اﻋﻠﻰ ﯾﺔ اﻟﺣﺷر اع اﻻﻧو ﻟﻛﺛﺎﻓﺔ اﻟﻛﻠﻲ اﻟﻣﻌدل ﺑﻠﻎ ﺣﯾث ﻓﻲ ﯾﻣﻛن اﻟﺳﺎﻋﺔ ﻋﻧد ﻣﺎﯾس ﺷﻬر pm ١٢ وﺑﻠﻎ ٧٦٨ ﻓرد / م ٢ ﻟﻠﻛﺛﺎﻓﺔ ﻛﻠﻲ ﻣﻌدل اﻗل اﻣﺎ ﻛﺎن اﻟﺳﺎﻋﺔ ﻋﻧد pm ٧ ﻟﺷﻬر وﺑﻠﻎ ﻧﯾﺳﺎن ٨١ ﻓرد / م ٢ ان اﻟﻰ اﻟﻧﺗﺎﺋﺞ اوﺿﺣت اﻟﻧوع Megachil sp ﻛﺎن اﻛﺛر ﻣﻌدل ﻛﺛﺎﻓﺔ اﺳﺔ اﻟدر ﺧﻼل وﺑﻠﻐت ٨٩٢ ﻓرد / م ٢ اﻣﺎ اع اﻧو ﺗﺑﺔ ر ﻓﻘد اﻟذﺑﺎب اﻟﻧوع ﺷﻛل Episyrphus balteatus ﻛﺛﺎﻓﺔ ﻣﻌدل وﺑﻠﻐت ٤٣٥ ﻓرد / م ٢ ﺑﯾﻧﻣﺎ اع اﻧو اﻟﻧوع ﺷﻛل ﻓﻘد اﻻﺟﻧﺣﺔ ﺣرﺷﻔﯾﺔ ﺗﺑﺔ ر Colias croceus(L.) ﻛﺛﺎﻓﺔ ﻣﻌدل و ﺑﻠﻐ ت ٧٣٠ ﻓرد / م ٢ ﻛﻣﺎ اﻟﻧﺗﺎﺋﺞ اوﺿﺣت اﻟﻣﻠﻘﺣﺔ اﻟﻧﺣل اع أﻧو ان اﻟﻰ وﻫﻲ melifera Apis ؛ Megachile Sp. ؛ Anderena Sp و Halictus Sp. اﻛﺑر ﺷﻛﻠت ات ﯾﺎر اﻟز ﻋدد ﻣﻌدل ﻣن ﻧﺳﺑﺔ اﻟﺳﺎﻋﺔ ﻋﻧد pm ١٢ وﺑﻣﻌدل ١٣.٧٦ ؛ ٨.٢٣ ؛ ٦.٣٤ ، ٣.٧٣ ة زﻫر / دﻗﯾﻘﺔ اﻟ ﺑﻊ اﻟﻣر ﻟﻠﻣﺗر اﺣد و اﻟﺗ ﻋﻠﻰ ﺗﯾب ر. اﻟﻧوﻋﺎن وﺧﺻوﺻﺎ اﻟذﺑﺎب ﺗﺑﺔ ﻟر ات ﯾﺎر اﻟز ﻋدد ﻣﻌدل اﻣﺎ Episyrphus balteatus و Metasyrphus taeniops ﻛﺎن ﻓﻘد ات ﯾﺎر اﻟز ﻋدد ﻣﻌدل ، اﻟﺛﻼﺛﺔ اﻻوﻗﺎت ﺧﻼل ﺑﺎ ﻣﺗﻘﺎر ﺑﯾﻧﻣﺎ ا زداد ﻻ ات ﯾﺎر اﻟز ﻋدد ﻣﻌدل اﻟﻧوع وﻣﻧﻬﺎ اﻻﺟﻧﺣﺔ ﺣرﺷﻔﯾﺔ ﺗﺑﺔ ر اع ﻧو Colias croceus(L.) ا ﻋﺻر اﻟﺧﺎﻣﺳﺔ اﻟﺳﺎﻋﺔ ﻋﻧد و ﺑﻣﻌدل ﺑﻠﻎ ١١.٩٥) ة زﻫر / دﻗﯾﻘﺔ (ﺑﻊ اﻟﻣر ﻟﻠﻣﺗر اﺣد اﻟو. ﻧﺗﺎﺋﺞ ﺧﻼل وﻣن اﺳﺔ اﻟدر ﺑﺎن ﯾﺗﺿﺢ اﻟذﺑﺎب اع اﻧو وﺑﻌض اﻟﻧﺣل اع اﺟدﻻﻧو اﻟﺗو و اﻟﻧﺷﺎط ﯾﻣﻛن ﻣﺎ اﻋﻠﻰ ﻛﺎن ﺑﯾن ﻣﺎ اﻟﺳﺎﻋﺔ pm ١٢-pm ٧ ﻣﺎ اﻋﻠﻰ ﻛﺎن اﻻﺟﻧﺣﺔ ﺣرﺷﻔﯾﺔ ﺗﺑﺔ ر اع اﻧو اﺟد وﺗو ﻧﺷﺎط ﺑﯾﻧﻣﺎ اﻟﺳﺎﻋﺔ ﻋﻧد ﯾﻣﻛن pm ٥. Abstract The aims of the present study are to identified the native pollinators of Trifolium alexandrinum L. Clover field and to determine their daily activities for three months April, May and June/2010 in Al-qadisiya governorate.Elven species of wild bee and other insect were recorded on the flower of the crop which are four species from hymenoptera ,Apis melifera ; Megachil sp.; Halictus sp.; Anderena sp; and three species of diptera its Metasyrphus taeniops; Eristalis anaeus; Episyphus balteatus ; and four species from Lepidoptera it were Lycaeides Melissa(alpime); Autograph gamma L; Colias croceus (L.); and finally Artogeia rapa. The density of insect species were different according to the different time and insect species, Results showed that total mean of insect density were higher levels in May month at 12pm as it reached 768 idividual/m2,but lower level of the total mean density in April at 7pm as it reached 81 idividual/m2.Megachil sp was larger density mean reached 892 idividual/m2. but Episyrphus balteatus of diptera species was formed 435 idividual/m2.but types of Lepidoptera species such as Colias croceus(L.) was formed 730 idividual/m2.resuts appear that bee pollinators species Apis melifera , Megachil sp.; Halictus sp. and; Anderena sp, These four species contained ١٣.٧٦ , ٨.٢٣ , ٦.٣٤ , ٣.٧٣ flower/min of the total ratio of visiting means respectively. In diptera species especially Metasyrphus taeniops, Episyrphus balteatus were formed similar levels of visiting means for three time.but hn Lepidoptera species such as Colias croceus(L.) was increasing of visiting means at 5pm to reach 11.95 flower/min of one meter squares. This result showed to different appears of species due to different time of same day.the optimal activity of bee and some diptera species were in 7am-12pm, but the optimal activity of Lepidoptera species was in 5 pm. .
Within the amazon region 130 species of stingless bees can be found, grouped into 26 genera, where Melipona and Trigona are the most represented. In Acre there is official record of 20 genera and 64 species of stingless bees, among those M. eburnea is one of the members, being very common in the Acre forests and rationally created by melipolinculturists, in the municipality of Rio Branco. The objectives that integrated this job were: (i) to identify botanical species used by M. eburnea to collect pollen and nectar; (ii) to determine the frequency of pollen types found in honey itself as well as in the pollen charges of the bees´ corbicles of M. eburnea; (iii) to point out the flowering seasonality of the plants used by M. eburnea to collect pollen and nectar; (iv) to identify the growth habits of M. eburnea´ s pollen and nectar suppliers; (v) to indicate the contribution of native, cultivated and ruderal species in the supply of pollen and nectar to M. eburnea; (vi) to verify the influence of rainfall and light intensity on the supply of food resources for M. eburnea, throughout the year; (vii) to assess the protein content and the specific concentration of amino acids content in the pollen collected by M. eburnea, throughout the year and (viii) to check if there is a link between the different levels of protein and amino acids with the diversity of pollen types collected by M. eburnea. Once a week, from April 2017 to March 2019, from 5 am to 9 am. worker bees from three colonies of M. eburnea have been sampled. The weekly samples have been gathered, thus creating a single monthly sample. The pollen sources used by M. eburnea have consequently been determined by sampling the pollen loads from the worker bees, when returning to the colony with pollen in their corbicles. The sources of nectar have been confirmed by checking on honey samples taken from the pots, still open. In order to quantify the protein content and the amino acid profile, the samples have been collected together with those of pollen and honey, in still open pollen pots and from the same selected colonies. After that, pollen has been collected and sampled from plants present in the study area, as well as from the dried ones, kept in the UFAC – UFACPZ herbarium, to preparing a reference “pollen library”. Microscopy slides have been set to perform qualitative and quantitative analysis of the pollen types contained in the samples of the bees´ charges, honey and pollen pots. Palynological analysis have shown that M. eburnea bees have harvested floral resources from 115 botanical species, belonging to 47 families and 96 genera: 61 species have been used both to collect pollen and nectar, 19 only for pollen collection and 35 just for nectar extraction. Fabaceae (Mimosoideae) and Myrtaceae
have been the main sources of pollen and nectar to M. eburnea, in all the seasons of the bee calendar (rainy, rain-dry transition, drought and dry-rain transition). Among the pollen types of the corbicle loads´ samples, Mimosa pudica represented the highest concentration 6.79 (log 10). When considering the volume of the pollen grain, Hybiscus was the most important. Among those classified in the honey samples, Crotalaria retusa obtained the highest concentration of 6.56 (log 10). In the main group of plants visited by M. eburnea, 50.61% were native; 36.59% cultivated and 12.81% ruderal. Referring to the habit, 40.25% were trees; 34.59% bushes; 13.21% herbs; 8.18% vines; 3.14% sub-shrubs and 0.63% epiphytes. The seasonality of the botanical species utilized by M. eburnea to collect floral resources has demonstrated wide variation throughout the study period: the pollen types Solanum and M. pudica have turned out being the least seasonal, in the corbicula loads´ samples. Among honey ones, Combretum. These pollen types have been present during the all assessed months. 13 amino acids have been found in the pollen stored by M. eburnea, nine of which are essential ones. In this group, Arginine was preponderant. Among the non-essentials, Proline has shown the highest concentration, being predominant in the set of identified amino acids as well, representing 68.5%. The levels of raw protein vary from 16.57 to 24.39%, within an average of 20.75%. The highest concentration of flowering has occurred in the dry season, characterized by a decrease in the volume of rainfalls and an increase in the amount of hours of sunshine. As for the totality of plants visited by M. eburnea, 38.39% offered pollen and/or nectar, during the dry season. It is concluded that (i) M. eburnea presents a generalist behavior in the use of pollen sources; (ii) the main sources of pollen and nectar for M. eburnea have been Fabaceae and Myrtaceae; (iii) native vegetation, mainly trees and bushes, have been the primary source of pollen and nectar for M. eburnea; (iv) an association among native, cultivated and ruderal plants provides appropriate amounts of proteins and amino acids for M. eburnea.
Keywords: Stingless bee, Rational breeding, Meliponiculture, Floral resources, Agroforestry system – AFS.
Corbiculate bees comprise a distinctive radiation of animals including many familiar species, such as honey bees and bumble bees. The group exhibits a broad variety of morphologies and behaviors, including solitary, social, and cleptoparasitic lifestyles. Since corbiculate bees play a critical role for the interpretation of eusocial behaviors, understanding their phylogeny is crucial to explain patterns and mechanisms of social evolution. Despite advances to unveil corbiculate relationships employing genomic data, the drivers of conflict between molecular and morphological hypotheses are still not fully understood. Morphological datasets favor a single origin for highly eusocial behaviors (i.e., Apini + Meliponini) whereas molecular datasets favor other scenarios (e.g., Bombini + Meliponini). Explanations for this incongruence have been suggested, including quality, quantity, and source of data or methodological issues. In this work we tackled this problem by generating the most extensive morphological dataset for the corbiculate bee species by exploring characters from all body regions, including external and internal adult skeletal anatomy. We produced a matrix with 289 characters for 53 taxa of Apidae, including 24 corbiculate bees. We explored different analyses and optimality criteria including extended implied weights parsimony and two partitioning schemes for Bayesian inferences. We contrasted hypotheses with Bayesian topological tests and conducted analyses to investigate if characters were prone to concerted convergence. Our results are congruent with the conclusions of previous studies based on morphology, recovering Apini sister to Meliponini and both of them together sister to Bombini. Finally, we provide our interpretations on the corbiculate controversy and provide a conciliatory scenario about this issue.
Stingless bees (Meliponini) belong to the corbiculate bees, a monophyletic group of bees characterised, among other things, by their particular pollen-carrying structures (“pollen baskets” or corbiculae) on the hind legs (Chap. 1; Fig. 1.1) (Michener 2007). The evolution of the concave pollen baskets allowed corbiculate bees to transport large amounts of pollen and especially resin in an efficient way (Martins et al. 2014). Three other tribes belong to the corbiculate bees, the highly eusocial honey bees (Apini), the primitively eusocial bumble bees (Bombini), and the mostly solitary orchid bees (Euglossini) (Grimaldi and Engel 2005; Michener 2007). The term higher eusocial (sometimes also advanced eusocial or superorganismal) is often used to refer to species with morphologically distinct queen and worker castes (Michener 1974, 2007; Danforth et al. 2013; Boomsma and Gawne 2018). Occasionally, a perennial colony lifestyle (Michener 2007) or extensive food exchange among the adults (Michener 1974) is mentioned as defining features of highly eusocial colonies. In primitively eusocial colonies, queens and workers appear morphologically similar and can be distinguished mainly by their size (Michener 1974, 2007; Danforth et al. 2013). Stingless bees can be distinguished from the other corbiculate bees by a number of morphological traits, including a reduced forewing venation, the penicillum, the absence of an auricle (pollen press), the presence of a jugal lobe in the hind wing, and the reduced sting (Figs. 1.1 and 1.3) (see Michener 2007 for details).
Sweat bees in the subgenus Lasioglossum (Dialictus) are one of the most diverse and abundant bee taxa, and a critically important component of bee biodiversity. Yet, the most basic taxonomic knowledge of these bees is lacking in many regions. As a step towards a better understanding of the L. (Dialictus) of the western Nearctic region, a revision of the ‘red-tailed’ L. (Dialictus) species was completed. Thirty-six species were revised, 20 of which are described as new, and two names are treated as junior subjective synonyms. Descriptions, figures, distribution maps, floral hosts, and keys to species for females and males are provided. The following 20 species are described as new: Lasioglossum (Dialictus) arenisaltans sp. nov., L. (D.) argammon sp. nov., L. (D.) austerum sp. nov., L. (D.) cactorum sp. nov., L. (D.) cembrilacus sp. nov., L. (D.) clastipedion sp. nov., L. (D.) clavicorne sp. nov., L. (D.) decorum sp. nov., L. (D.) festinum sp. nov., L. (D.) imbriumbrae sp. nov., L. (D.) julipile sp. nov., L. (D.) lilianae sp. nov., L. (D.) meteorum sp. nov., L. (D.) miltolepoides sp. nov., L. (D.) minckleyi sp. nov., L. (D.) perditum sp. nov., L. (D.) rufornatum sp. nov., L. (D. ) spivakae sp. nov., L. (D.) tessellatosum sp. nov., and L. (D.) torrens sp. nov. Previously unknown males of L. (D.) clematisellum (Cockerell, 1904), L. (D.) droegei Gibbs, 2009, L. (D.) kunzei (Cockerell, 1898), and L. (D.) pallidellum (Ellis, 1914) are described and figured for the first time. Lasioglossum (Dialictus) clarissimum (Ellis, 1914) (= Halictus clarissimus Ellis, 1914) and L. (D.) perexiguum (Sandhouse, 1924) (= Halictus (Chloralictus) perexiguus Sandhouse, 1924) are new subjective junior synonyms of L. mesillense (Cockerell, 1898) (= Halictus nymphalis var. mesillensis Cockerell, 1898). A lectotype specimen is newly designated for L. mesillense, for which the location of the type material has not previously been known. The following five new records for Mexico are reported: L. clematisellum, L. droegei, L. eophilus (Ellis, 1914), L. kunzei, and L. pallidellum.
Three species of Ammobatoides are reviewed. One new species, Ammobatoides radoszkowskii sp. nov. (Russia and North China) is described and illustrated. A male lectotype is designated for Paidia melectoides Radoszkowski, 1872. One new subjective synonym is established: Ammobatoides scriptus (Gerstaecker, 1869) =Paidia melectoides Radoszkowski, 1872, syn. nov. A key to species for both sexes is given.
Five Chinese species of Nomia (Gnathonomia) Pauly, 2005 are treated in this paper: Nomia fusciventris Zhang & Niu, sp. nov. from Fujian Province is described as a new species; N. aurata Bingham, 1897 and N. wahisi Pauly, 2009 are recorded from China for the first time, and the male of N. pieli Cockerell, 1931 is newly reported. An updated diagnosis is provided for the subgenus Nomia (Gnathonomia) to accommodate Chinese endemic species which lack the white integumental bands found in most previously described forms. A key to the known Chinese species is provided.
Dar, S.A., Dukku, U.H., Ilyasov, R.A., Kandemir, I., Lee, M.L., Özkan Koca, A. 2019. Chapter 4. The classic taxonomy of Asian and European honey bees. 107-137 pp. In: R.A. Ilyasov and H.W. Kwon. [eds.]. Phylogenetics of Bees. CRC Press, Taylor and Francis Group, Boca Raton, London, New-York, USA. 290 pp. ISBN 9781138504233.
Abstract. Honey bees of the genus Apis, belonging to the family Apidae and the superfamily Apoidea in the order of insects Hymenoptera. The number of Apis species and their identification methods are discussed. According to different authors, the number of species of the genus varied from 6 to 24. While Apis mellifera inhabits West Asia, Africa and Europe, the ranges of all other species, including Apis cerana, are limited to Asia. A. mellifera and A. cerana are two species widely used in agriculture for the pollination, the production of honey and other products. They have adapted to wide climatic conditions. Intraspecific taxonomy for both species is incomplete and contradictory. In this review, all available studies of A. mellifera and A. cerana are analyzed to ordering the modern taxonomy of honey bees. We found that there are 27 subspecies for A. mellifera and 7 subspecies for A. cerana. However, these data are not ultimate, since some subspecies of A. mellifera and A. cerana remain unexplored.
La morfología es la base fundamental sobre la cual se desarrolla la taxonomía. El objetivo de este capítulo es ofrecer un glosario en español y un léxico español-inglés de más de 400 términos utilizados en los trabajos de taxonomía de hormigas (Hymenoptera: Formicidae). Dibujos e imágenes en microscopía electrónica de barrido y en microscopía óptica acompañan las definiciones para facilitar la comprensión de los términos. Definimos las principales estructuras externas de la casta obrera, las castas reproductoras (venación, genitales y mesosoma, de machos y ginas), y los términos relativos a la posición y a la orientación de las estructuras en el cuerpo con respecto al mesión. Uno de los principales criterios que hemos usado para denominar algunas estructuras es la homología existente entre las partes de las hormigas y las de otros himenópteros. En consecuencia, recomendamos el uso de unos términos sobre otros.
A female of the mining bee Andrena anatolica Alfk. has been collected from Mãcin National Park, North of Dobrogea. This species has not been recorded previously in Romania; its geographical distribution and a short morphological description is presented. Résumé. Une femelle de l'abeille mineure Andrena anatolica Alfk., a été capturée dans le Parc National Mãcin, nord du Dobrogea. Cette espèce n'a pas été signalée jusqu'à présent en Roumanie. De même, on présente sa répartition géographique et une description morphologique courte.
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