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Cladistic analysis is provided to study the advancement of 29 biological personalities concerned nesting behavior, colony provisioning, oviposition, cocoon spinning, defecation, and life cycle (nest characters) in 11 species of Osmia bees (Hymenoptera: Megachilidae) and also two outgrsoimg.org genera. A molecular phylogeny based upon 38 allozyme loci-as-personalities and a merged character phylogeny are created to compare through the colony character phylogeny. All phylogenetic trees assistance the monophyly of the genus Osmia and also the subgenus Osmia (Osmia) through Osmia ribifloris Cockerell basal to the other species in this subgenus. The subgenus Helicosmia (=Chalcosmia) is refixed as a sister clade to O. (Osmia) in the molecular tree and also as sister to the subgenus Cephalosmia in the colony character and also merged trees. Infinish organic indevelopment from added species of all 3 subgenera is provided to support the consistency of swarm personalities at the subgeneric level and their usage in the facility of subgeneric phylogenies in the family members Megachilidae.
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The concept that behavior evolves in fundamentally the exact same fashion as morphology justifies the use of behavioral characters both on their very own and also unified via morphological or molecular personalities in the inference of phylogenies (Wenzel 1992, de Queiroz and also Wimberger 1993, Proctor 1996). This possibility has long been recognized by ethologists (Whitguy 1899, Heinroth 1911, Tinbergen 1959), yet phylogenetic studies making use of behavior characters are still scarce (Wenzel 1992, de Queiroz and Wimberger 1993, Proctor 1996). It has been said that behavioral personalities are hard to homologize and are more evolutionarily labile (subject to convergence and also reversal) than morphological personalities (Atz 1970, Baroni Urbani 1989), but some research studies have actually shown equivalent levels of homoplasy between behavioral and morphological or molecular data sets (McLennan et al. 1988, Arntzen and Sparreboom 1989, Coddington 1990, Prum 1990, de Queiroz and Wimberger 1993, Proctor 1996). The restricted use of behavior characters in phylohereditary research studies may be attributed to the challenge of collecting behavioral data, a task that frequently requires big quantities of observation time. Sometimes, behavior data have the right to be derived through the study of frameworks that are the result of certain habits. In nesting Aculeate Hymenoptera (ants, bees, wasps), a significant amount of behavior (nest structure, provisioning, oviplace, cocoon spinning) and physiological (life cycle, defecation) indevelopment can be derived via swarm analysis. Thus, behavior characters, regularly regarded nesting tasks and also colony design, have been used in aculeate Hymenoptera phylogenetic researches, greatly in combination via larger numbers of morphological characters (Carpenter 1982, 1987, 1988,,; Carpenter and also Cumming 1985; McGinley and also Rozen 1987; Baroni Urbani 1989, 1993,; Alexander 1990, 1991,; Rozen 1991, Wenzel 1993). Other Aculeate research studies have traced the evolution of behavior traits on phylogenies acquired from morphological or molecular personalities (Packer 1991, Carpenter et al. 1993, Chavarria and also Carpenter 1994, Engel and also Schultz 1997).
Bee (Apoidea) nesting behavior may be classified as burrowing (carpenters or miners) and also nonburrowing, according to whether species execute or perform not excavate their own colonies (Malyshev 1935, Stephen et al. 1969, O’Toole and Raw 1991). Amongst the latter, some species usage preestablished cavities, and also others build their cells in more or much less exposed instances. The primitive condition in the household Megachilidae is to nest in burrows, excavated in either soil (Fideliinae - Rozen 1970, 1973, 1977,,; McGinley and also Rozen 1987; - TrachusaMalyshev 1935, Michener 1941,, Westwealthy 1989) or lumber (Lithurgini - Malyshev 1935, Cros 1939, Brach 1978). However before, three or more forms of nests (including colonies in wood cavities, in snail shells, in cracks in rocks, burrows in the ground, burrows in stems, and also exposed nests) are uncovered in a number of acquired genera (Anthidium, Osmia, Hoplitis, Megachile subgenera Litomegachile and Delomegachile) (Malyshev 1935, Stephen et al. 1969, Eickwort et al. 1981, Westrich 1989). The absence of secreted substances in swarm construction and the constant incorporation of outside products, in cavity-nesters and also burrowing species, is characteristic of the Megachilidae (Stephen et al. 1969). The usage of particular nesting products has actually appeared numerous times independently throughout the advancement of the family. Soil use is found in species of Osmia, Hoplitis, Chelostoma, Megachile subgenera Eumegachile and also Chalicodoma, resin in species of Heriades, Chalicodoma, Anthidium, Trachusa, and Hoplitis, and masticated leaf in species of Osmia, Ashmeadiella, Hoplitis, and also Anthidium (Malyshev 1935, Krombein 1967, Stephen et al. 1969, Rust 1980, Westwealthy 1989, Bosch et al. 1993). Pollen specialization is one more plastic character. Many Megachilidae are polylectic, however oligolecty appears to be regular in the primitive Fideliinae and also Lithurgini (Rozen 1970, 1977,; Brach 1978; Roberts 1978; Parker and Potter 1973; Yáñez 1997) and also is common in Trachusa, Anthidium, Chelostoma, Heriades, Osmia, and also Hoplitis (Rust 1974, Westrich 1989, Cane 1996, Müller 1996). Regardless of this apparently high level of homoplasy, Torchio (1989) confirmed that organic characters can be offered to characterize Osmia subgenera.
In this work, we supplied a cladistic evaluation to examine the development of 29 personalities pertained to swarm building, colony provisioning, oviposition, life cycle, defecation, and also cocoon spinning (swarm characters) in 11 species of Osmia, representing 3 different subgenera. Two outteam species, Hoplitis adunca (Panzer) and also Megachile (Chalicodoma) angelarum (Cockerell), are consisted of in the analyses. We provide a molecular phylogeny based on 38 allozyme loci-as-personalities that we compare via the behavioral phylogeny and also the linked phylogeny of the 13 species. The two objectives of this article are as follows: (1) talk about the usefulness of nest personalities in the establishment of phylogenetic hypotheses in the Megachilidae and also (2) discuss the development of nesting habits and related biological attributes among the three Osmia subgenera thought about and also in relation to the remainder of the Megachilidae.
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Materials and also Methods
The genus Osmia has ≈500 species found in the Holarctic (Rust 1974). All Osmia are solitary, and also many kind of nest in preestabliburned cavities. Nest-traps (Krombein 1967) consisting of cavities of different diameters drilled in wooden blocks were put in various places in the USA, Japan, France, and Spain (Table 1). Bee colonies derived in the cavities were taken to the laboratory, where they were dissected. We collected information on nest style, nesting materials, provision structure and also oviposition, cocoon framework, shape and also place of fecal pwrite-ups, and also progeny developpsychological stages. Progeny were reared to the adult stage and also frozen for molecular analyses. We acquired colonies and adult bees for 11 species of Osmia in 3 various subgenera <Osmia, Helicosmia (=Chalcosmia), and also Cephalosmia> (Table 1). The subgenus Chalcosmia has actually been freshly homologized with the subgenus Helicosmia (Griswold and Michener 1997, Michener 2000). Megachile angelarum and Hoplitis adunca were offered as outgrsoimg.org species. Outgrsoimg.org standing was establiburned according to Roig-Alsina and also Michener (1993). Based on adult morphology, the Megachilidae are taken into consideration a monophyletic grsoimg.org split right into two subfamily members, the Fideliinae and also the Megachilinae, with the latter containing 4 people, the Lithurgini (basal), Anthidiini, Megachilini, and Osmiini. Although the condition of the last three tribes remains unrefixed as soon as larval characters are used, adult characters area Osmia and also Hoplitis within Osmiini, and also Megachilini as the sister tribe to Osmiini (Roig-Alsina and Michener 1993). Some authors take into consideration Hoplitis as a subgenus within Osmia (Westrich 1989).