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Preferred total evidence dated phylogeny of Megachilidae. Majority-rule consensus tree from Bayesian analysis using fossils as terminals under the FBD tree prior. Blue bar at each node represents the 95% highest posterior density age range. Posterior probability below 100 indicated above each node.
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A unique feature among bees is the ability of some species of Megachile Latreille s.l. to cut and process fresh leaves for nest construction. The presence of a razor 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 grou...
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... phylogenies: In the preferred Bayesian analysis for the family Megachilidae ( Fig. 9), Pararhophitinae resulted as the sister group of Lithurginae, both taxa sister to Megachilinae. Within the latter subfamily, Dioxyini were the sister group of Glyptapini while Aspidosmiini rendered Ctenoplectrellini paraphyletic. Megachilini also rendered Osmiini paraphyletic, as they clustered with the osmiine genera Afroheriades ...
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... sister to Megachilinae. Within the latter subfamily, Dioxyini were the sister group of Glyptapini while Aspidosmiini rendered Ctenoplectrellini paraphyletic. Megachilini also rendered Osmiini paraphyletic, as they clustered with the osmiine genera Afroheriades Peters and Pseudoheriades Peters. The remaining osmiines are together in another clade (Fig. 9). The origin of crown Megachilidae was estimated at a median age of 111.3 Ma (95% highest posterior density 80.94-127.56 Ma) with crown Megachilini at 42.0 Ma (24.75-49.55 Ma). A similar topology resulted from the ML analysis of the combined dataset with extant taxa only. However, Pararhophitinae were the sister group of Lithurginae + ...
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... the fossil tribes Ctenoplectrellini and Glyptapini, whose phylogenetic positions were consistent among analyses (Fig. 9), that of the Dominican fossil M. glaesaria varied from being near Chelostomoides (Fig. 8) to be the sister group of all Megachilini (Figs. 10, 11). Interestingly, Engel (1999) discussed the possibility of both phylogenetic positions. Such instability might be the result of the low number of characters that we were able to score from ...
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... of Osmiini whereas the molecular analysis of Praz et al. (2008) placed them either as sister of either Anthidiini or Megachilini. Griswold (1985) also suggested a relationship with Megachilini based on the modified setae on the fifth sternum of the male. Our analyses consistently placed these two genera and Megachilini in a well-supported clade (Fig. 9). All other Osmiini clustered in another clade. Thus, in order to recognize a monophyletic Osmiini, we would need to either transfer them to Megachilini or distinguish them in their own tribe. Tranferring them to Megachilini weakens the recognition and diagnosis of this tribe because a morphological synapomorphy unambiguously present in ...
Citations
... Globally, seven subgenera of Stelis are recognized (i.e., Michener 2007), but only three of these occur in North America. The subgenus Dolichostelis Parker & Bohart, 1979 occurs from southern Ontario, Canada (i.e., S. louisae Cockerell, 1911-see Paiero & Buck 2004, extending south into northern South America (Gonzalez et al. 2012), and parasitizes nests of Megachile Latreille, 1802, subgenus Chelostomoides Robertson, 1901(Parker et al. 1987, a chalicodomiform or Group 2 (sensu Michener 2007) more recently considered a valid genus of Megachilini by Gonzalez et al. (2019); the six species of Dolichostelis occurring in North and Central American were revised by Parker & Bohart (1979), with an additional species described more recently (Gonzalez et al. 2012). Members of the Holarctic subgenus Heterostelis Timberlake, 1941 are nest parasites of Trachusa; the six North American and sharing several morphological characteristics, including the more narrowly pointed dorsal lip of the sternum 6; these latter two species also have more extensive pale maculations on the metasoma than most members of the permaculata species group (Timberlake 1941;Parker, in Rozen 1987). ...
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.
... This is consistent with the reconstructed phylogeny of the genus Hoplitis, which placed H. curvipes and H. mitis as members of the same clade amidst and not basal to species of Alcidamea, which use leaf pulp as nest building material (Sedivy et al. 2012). Within the family Megachilidae, the use of leaf fragments for brood cell construction is most typical for numerous species of the genus Megachile Latreille, 1802 (Megachilini), which, however, differ from H. curvipes and H. mitis by their habit to usually cut regularly circular to elliptical leaf pieces (Michener 2007;Ivanov and Zhidkov 2010;Gonzalez et al. 2019). Some Megachile species, however, masticate the margins of the cut leaf pieces in order to glue them together (Ivanov and Filatov 2008). ...
... Some Megachile species, however, masticate the margins of the cut leaf pieces in order to glue them together (Ivanov and Filatov 2008). As in the H. curvipes species group, the leaf-cutting behaviour of Megachile probably evolved from ancestors, which used leaf pulp as the main nest building material, but did not completely masticate the cut leaves (Michener 2007;Gonzalez et al. 2019). ...
Two nests of Hoplitis curvipes are described from Apulia (Italy) and Dagestan (Russia). Both nests consisted of two brood cells placed side by side under a stone. The cells were neither attached to each other nor to the substrate. They were constructed from leaf fragments, which were imbricately arranged, forming a cone-like structure; each leaf fragment consisted of a basal part that was masticated to leaf pulp and an api-cal part that protruded freely from the cell wall. The cell wall was formed by the fusion of the masticated basal parts of the leaf fragments and thus entirely consisted of leaf pulp. The cell was sealed with a closing plug made of pure leaf pulp; a few leaf fragments were glued to its outer surface. The cocoon consisted of two layers: the outer layer was restricted to the anterior portion of the cell and had several longitudinal air-exchange slits on its lateral surface, while the inner layer had an air-exchange orifice in its most anterior dome-shaped top. Results of measurements of brood cell dimensions and contents are provided. The nesting biology of species of the H. curvipes group is discussed.
... From its origin estimated to be in the latest Cretaceous-Paleocene (nearly 66 My ago) and diversification during late Eocene (deposits of Baltic amber), bees of the family Megachilidae have colonized all kinds of habitats around the world (Michener, 2007;Michez et al., 2012;Danforth et al., 2019). The huge genus Megachile originated presumably in the Early to Middle Eocene, is globally distributed and currently consists of approximately 1500 species (Raw, 2004(Raw, , 2007Engel and Perkovsky, 2006;Michener, 2007;Gonzalez et al., 2019). In the Neotropics, Megachile is very numerous with more than 400 species (Raw, 2004(Raw, , 2007Moure et al., 2007). ...
... Bees were identified by Arturo Roig Alsina and deposited in the Entomology National Collection (MACNEn). Most nests belonged to two identified Megachile species: Megachile infima Vachal (nests 10 and 11 and scopal pollen Sc2) (see Table I), belonging currently to the subgenus Chrysosarus (Gonzalez et al., 2019) although another study classified it in Dactylomegachile (Durante et al., 2020). This species is native to Argentina and is distributed in parts of South America from the Mendoza province in Argentina to the São Paulo state in Brazil (Silveira et al., 2002;Durante et al., 2020); and M. pusilla Pérez (nests 5-9 and 12-13 and scopal pollen Sc1) (see Table I), an exotic species belonging to the subgenus Eutricharaea whose area of origin is around the Mediterranean Sea (Soltani et al., 2017). ...
... In the urban microhabitats studied, the native M. infima nested in semi-exposed spaces as well as different human-made structures such as water taps and hosepipes and exclusively used petals to line brood cells. Among Megachile, the use of only petals, without leaf pieces or mud, has been reported a few times in the subgenera Megachiloides and Megachile (Bohart and Youssef 1972;Gonzalez et al., 2019) and appears to be related to the absence in the development of interdental laminae in M. infima (cited as M. parsonsiae Schrottky in Gonzalez et al., 2019 [Fig. 2c]). ...
... Megachile Latreille (Hymenoptera: Megachilidae) is a large (approximately 1505 spp.) genus of solitary bees distributed worldwide (Ascher and Pickering 2021). Megachile females nest in pre-existing natural or artificial cavities in the ground or above ground, lining their breeding cells using small pieces of elliptical and circular fresh leaves to protect them from desiccation and natural enemies (Marinho et al. 2018;Sabino and Antonini 2017;González et al. 2019). The nests consist of a linear sequence of brood cells within the tube, each with a supply of food (a mixture of pollen and nectar) where a single egg is deposited (Michener 2007). ...
Megachile amparo (González, Revista Colombiana De Entomología 32(1):93–96, 2006) is the only high Andean leaf-cutter bee reported in Colombia and is possibly endemic to the Colombian Andes. Although it is frequently observed, even in urban areas, its biology and ecology remain unknown. The present study aimed to describe detailed aspects of its bionomy. Trap-nests were installed on the Campus of the Nueva Granada University (Cajicá, Colombia) from June/2018 to March/2020. The trap-nests were wooden blocks (25 × 15 × 14 cm) with 30 cavities of Ø = 1 cm and different lengths (50 mm, 75 mm, and 100 mm) lined with waxed paper straws. During the observations, an increasing number of trap-nests were installed, increasing from 250 to 720 cavities. The trap-nests were monitored three times a week, recording both the date the start and end building by female. Most of the nest were maintained in the field to estimate the sex ratio, cell survival, and total development time under natural conditions. Thirty-two nests were removed at different times of the observation period to establish number of cells per nest, and cells built per female per day. We incubated 20 cells from different nests at 18 °C, 22 °C, 26 °C, and 32 °C to estimate the base temperature, thermal constant k (developmental time in degree days), and cell survival. Young cells of different positions were dissected and weighed to characterize food provision and brood cells. Computerized tomography-CT scans were performed in 30 brood cells to determine if diapause occurred during prepupal stage. Females nested 7- and 10-cm-long cavities and the number of cells per nest varied with cavity length. The brood cells had a length of 1.23 ± 0.12 cm and a diameter of 0.92 ± 0.05 cm. The female spends 1.17 ± 0.29 days to build a brood cell. Food provision varied according to the position of the brood cell in the nest. The adults of M. amparo present a marked seasonality being more active during dry months. Base temperature and thermal constant k were different for males and females. The sex ratio is female biased (1.9:1), and cell survival in the field was 89% with no cleptoparasites or predators recorded.
... These species had been previously assigned to either the subgenus Ptilosaroides Mitchell (Schlindwein, 1998), Ptilosarus Mitchell (Raw, 2002(Raw, , 2007Silveira et al. 2002), or Rhyssomegachile Mitchell (Moure et al., 2007;Moure & Melo, 2022). All these subgenera, together with Neochelynia Schrottky and Zonomegachile Mitchell, form a group of closely related Neotropical subgenera (Gonzalez et al., 2018;Gonzalez et al., 2019). The morphological distinctiveness of Chalepochile among all these subgenera is hampered by the lack of knowledge of the female sex, because females bear important features used for characterizing subgenera. ...
The subgenus Chalepochile Gonzalez & Engel of Megachile Latreille was known only from males of two species. In this contribution the female of M. ardua Mitchell, type species of Chalepochile, is described, as well as a new species based on both sexes. The new species M. (Chalepochile) bilineata occurs in the provinces of Córdoba, La Rioja and Mendoza in Argentina. The phylogenetic relationships of the subgenus Chalepochile are discussed in the light of characters of the females.
... Individual species in other orders of insects (and arthropods) have been the subject of micro-CT techniques, including certain flies (Wasserthal et al., 2018), locusts (Harrison et al., 2013;Greco et al., 2014), bees (Greco et al., 2008;Gonzalez et al., 2019;Herhold et al., 2019), and even centipedes (Hilken et al., 2021). Studies have also focused on tracheal compression using high-resolution synchrotron imaging (Westneat et al., 2003), including hypoxiainduced compression (Greenlee et al., 2013); 3D printing of tracheal structures (Greco et al., 2014); quantitative analysis of the distribution of air spaces (Shaha et al., 2013); and even capturing insects during a variety of behaviors (Alba-Tercedor et al., 2021). ...
A broad comparative study of insect respiratory morphology is presented. Tracheae, epidermal invaginations extending into the body in branching networks of tubes, supply tissues with direct access to air for gas exchange. While previous tracheal studies focused on a handful of taxa and lacked in consistency, here a unified system of tracheal nomenclature is established using visualiza- tions from micro-CT scanning of representatives from apterygotes, Paleoptera, and Polyneoptera, totaling 29 species, 29 genera, and 26 families in 13 insect orders. Three-dimensional visualizations of named tracheal branches establish robust assessments of homology and provide a framework for further studies across class Insecta. Patterns in respiratory architecture are presented along with a discussion of future investigations into phylogenetic and physiological questions.
... Leafcutter bees, for instance, need young leaves for constructing brood cells 17 (Fig. 1). Our previous studies [15][16][17][18] suggest that the leafcutter bees prefer plants of specific lineages to forage leaf fragments. Megachilidae is a cosmopolitan solitary bee family, having over 4100 species worldwide 19 ; the genus Megachile, which comprises leafcutters, has about half of these species and is globally distributed 19 . ...
Pollinator conservation is a global priority. Efforts are taken to restore pollinators by improving flower resources, a crucial driver of pollinator diversity and population growth. It helped gardening and landscaping supply chains, which introduced lists of bee-friendly plants and bee hotels, yet, desirable results seem distant. One shortcoming of the present schemes is that they lack a cohesive planning for nesting opportunities and nesting provisions for wild solitary bees, the crucial pollinators of crop and wild plants. We tested whether the world’s popular ornamental plant, rose (Rosa chinensis Jacq.)—a hitherto unlisted bee-friendly plant—can aid in conserving leafcutter bees, which require fresh leaves for constructing nest cells. We surveyed 2360 rose plants in 136 sites in rural and urban places and lowlands and highlands of south (8°N–12°N) and northeastern India (26°N–27°N) for the characteristic notches the bees leave on foraged leaves. We reared brood constructed with rose and non-rose leaves to examine the brood success rate. About a quarter of all the roses surveyed had the notches of leafcutter bees on the leaves. However, the proportion of cut roses varied considerably among sites. Bees used roses much higher in urban areas and lowlands than in rural areas and highlands. The selection of plants was negatively associated with pesticide application. The brood success rate was 100% for the brood that was constructed by the leaves of rose and non-rose plants. Rose flowers do not support bees, but rose leaves indeed do. We recommend rose plants in leafcutter bee conservation and restoration schemes, particularly in urban environment.
... In the extant ant species that we examined for this study, the shovel shape is very clearly developed, although with a comparatively long and narrow blade in Protanilla (see also . A very similar shape also occurs in a few other groups of Aculeata such as the megachilid bees (Gonzalez et al. 2019). However, given the distant relationship to these bees, we consider this to be the result of independent evolution. ...
The fossil record allows a unique glimpse into the evolutionary history of organisms living on Earth today.
We discovered a specimen of the stem group ant †Gerontoformica gracilis (Barden and Grimaldi, 2014) in
Kachin amber with near-complete preservation of internal head structures, which we document employing
μ-computed-tomography-based 3D reconstructions. We compare †Gerontoformica to four outgroup taxa and
four extant ant species, employing parsimony and Bayesian ancestral state reconstruction to identify morphological
differences and similarities between stem and crown ants and thus improve our understanding of
ant evolution through the lens of head anatomy. Of 149 morphological characters, 87 are new in this study,
and almost all applicable to the fossil. †Gerontoformica gracilis shares shortened dorsal tentorial arms,
basally angled pedicels, and the pharyngeal gland as apomorphies with other total clade Formicidae. Retained
plesiomorphies include mandible shape and features of the prepharynx. Implications of the reconstructed
transitions especially for the ant groundplan are critically discussed based on our restricted taxon sampling,
emphasizing the crucial information derived from internal anatomy which is applied to deep time for the first
time. Based on the falcate mandible in †Gerontoformica and other Aculeata, we present hypotheses for how
the shovel-shaped mandibles in crown Formicidae could have evolved. Our results support the notion of
†Gerontoformica as ‘generalized’ above-ground predator missing crucial novelties of crown ants which may
have helped the latter survive the end-Cretaceous extinction. Our study is an important step for anatomical
research on Cretaceous insects and a glimpse into the early evolution of ant heads.
... The megachiline bee genus Noteriades Cockerell is a seemingly relict genus, with comparatively few species occurring in both temperate and tropical regions of sub-Saharan Africa and southern Asia (Michener 2007). Traditionally, the genus was classified in the tribe Osmiini and among the Heriades-group of genera owing to its distinctly hoplitiform body habitus and size (Michener 2007), but as originally hypothesized by Griswold (1985) Noteriades has recently been recovered as the sister group to Megachilini (Praz et al. 2008;Gonzalez et al. 2012Gonzalez et al. , 2019. Today, the genus is classified as the extant sister group to all other Megachilini (Gonzalez et al. 2019), a noteworthy position as the tribe otherwise includes the famous leaf-cutter and resin bees of the genus Megachile Latreille and its relatives (Michener 2007;Gonzalez et al. 2019). ...
... Traditionally, the genus was classified in the tribe Osmiini and among the Heriades-group of genera owing to its distinctly hoplitiform body habitus and size (Michener 2007), but as originally hypothesized by Griswold (1985) Noteriades has recently been recovered as the sister group to Megachilini (Praz et al. 2008;Gonzalez et al. 2012Gonzalez et al. , 2019. Today, the genus is classified as the extant sister group to all other Megachilini (Gonzalez et al. 2019), a noteworthy position as the tribe otherwise includes the famous leaf-cutter and resin bees of the genus Megachile Latreille and its relatives (Michener 2007;Gonzalez et al. 2019). Unfortunately, nothing is known of the biology of any species of Noteriades, the discovery of which melittologists are encouraged to seek. ...
... Traditionally, the genus was classified in the tribe Osmiini and among the Heriades-group of genera owing to its distinctly hoplitiform body habitus and size (Michener 2007), but as originally hypothesized by Griswold (1985) Noteriades has recently been recovered as the sister group to Megachilini (Praz et al. 2008;Gonzalez et al. 2012Gonzalez et al. , 2019. Today, the genus is classified as the extant sister group to all other Megachilini (Gonzalez et al. 2019), a noteworthy position as the tribe otherwise includes the famous leaf-cutter and resin bees of the genus Megachile Latreille and its relatives (Michener 2007;Gonzalez et al. 2019). Unfortunately, nothing is known of the biology of any species of Noteriades, the discovery of which melittologists are encouraged to seek. ...
The little-known megachiline genus Noteriades Cockerell, 1931 is recorded from Vietnam for the first time. A new species, Noteriades hangkia Tran, Engel & Nguyen sp. nov. is described and figured based on a series of females collected from the provinces of the northern and central highlands of Vietnam. The genus is briefly discussed and a new subtribe is established, Noteriadina Engel, Tran & Nguyen subtrib. nov. of Megachilini. Lastly, an identification key and distribution map are provided for those species occurring in Southeast Asia.
... The suprageneric classification of Megachilinae, including the recognition and description of new taxa, has been reviewed in recent studies, supported by phylogenetic hypotheses (Praz et al., 2008;Litman et al., 2011;Gonzalez et al., 2012Gonzalez et al., , 2019. Some of these works investigated the relationships at tribal level and demonstrated the paraphyly of some lineages, such as the fideliine bees (Litman et al., 2011), the Osmiini (Praz et al., 2008) and the Anthidiini (Gonzalez et al., 2019). ...
... The suprageneric classification of Megachilinae, including the recognition and description of new taxa, has been reviewed in recent studies, supported by phylogenetic hypotheses (Praz et al., 2008;Litman et al., 2011;Gonzalez et al., 2012Gonzalez et al., , 2019. Some of these works investigated the relationships at tribal level and demonstrated the paraphyly of some lineages, such as the fideliine bees (Litman et al., 2011), the Osmiini (Praz et al., 2008) and the Anthidiini (Gonzalez et al., 2019). ...
Anthidiini comprise a large, diversified, and widely distributed tribe of megachiline bees. Recently, morphological and molecular analyses recovered five major monophyletic groups within the tribe. For this reason, we review the current classification of the tribe, giving status of subtribe to these lineages. A new subtribe, Epanthidiina (type genus: Epanthidium Moure), is proposed for a large group restricted to the Neotropical region. Morphological and molecular phylogenetic hypotheses support the taxonomic limits of the new subtribe. We also propose a new genus, Urbanthidium (type species: Anthodioctes gracilis Urban), in order to accommodate results from a previous study in which Anthodioctes Holmberg came out paraphyletic. Two species are transferred to the new genus: Urbanthidium gracile (Urban) comb. n., Urbanthidium psaenythioides (Holmberg) comb. n.
Keywords:
Apoidea; Classification; Megachilidae; Systematics; Taxonomy
