{"id":4,"date":"2009-01-21T21:41:39","date_gmt":"2009-01-21T20:41:39","guid":{"rendered":"https:\/\/wpethzprd.ethz.ch\/osmiini\/?page_id=4"},"modified":"2014-11-08T21:44:05","modified_gmt":"2014-11-08T20:44:05","slug":"phylogeny-and-classification","status":"publish","type":"page","link":"https:\/\/blogs.ethz.ch\/osmiini\/phylogeny-and-classification\/","title":{"rendered":"Phylogeny and Classification"},"content":{"rendered":"<p><a href=\"https:\/\/blogs.ethz.ch\/osmiini\/files\/2009\/09\/hoplitisloti_web.jpg\"><img loading=\"lazy\" decoding=\"async\" class=\"alignnone size-full wp-image-2820\" src=\"https:\/\/blogs.ethz.ch\/osmiini\/files\/2009\/09\/hoplitisloti_web.jpg\" alt=\"\" width=\"433\" height=\"296\" \/><\/a><\/p>\n<h5 style=\"text-align: left\"><span style=\"font-weight: normal\">Exposed brood cell of <\/span><em><span style=\"font-weight: normal\">Hoplitis (Hoplitis) loti<\/span><\/em><span style=\"font-weight: normal\"> (Morawitz, 1867) on the surface of a rock; the cell consists of small pebbles cemented together with mud. Foto A. M\u00fcller.<\/span><\/h5>\n<h3 style=\"text-align: left\">Phylogeny<\/h3>\n<p style=\"text-align: left\">In a recent molecular phylogenetic study based on three nuclear genes and 95 osmiine bee species (Praz et al., 2008b), the core osmiine genera were found to form a well-supported monophyletic group (Figure 1). However, three small genera, <em>Noteriades, Afroheriades <\/em>and<em> Pseudoheriades<\/em>, which were formerly included in the Osmiini (Michener, 2007), do not appear to belong within this clade. Gonzalez et al. (2012) reached a similar conclusion for <em>Noteriades <\/em>being a member of the Megachilini rather than the Osmiini. The phylogenetic position of the genera <em>Ochreriades<\/em> and <em>Bekilia<\/em> remains unsolved. Depending on the analysis, <em>Ochreriades<\/em> turned out to be either sister to the Osmiini or sister to the clade Megachilini+Anthidiini+Osmiini. <em>Bekilia<\/em> was not included in the analysis. It is a monotypic genus with uncertain affinities due to the lost type material and may belong to the Anthidiini close to <em>Afrostelis<\/em> rather than to the Osmiini (Griswold &amp; Michener, 1998; Michener, 2007).<\/p>\n<h3>Classification<\/h3>\n<p style=\"text-align: left\">The molecular phylogeny of Praz et al. (2008b) results in four taxonomic changes in comparison to the classification of Michener (2007): the former genera <em>Stenosmia<\/em> and <em>Hoplosmia<\/em> are reduced to subgeneric rank in <em>Hoplitis<\/em> and <em>Osmia<\/em>, respectively, <em>Micreriades<\/em> is recognized as a subgenus of <em>Hoplitis, <\/em>and the subgenus <em>Nasutosmia<\/em> is transferred from <em>Hoplitis<\/em> to <em>Osmia<\/em>. It further enables a clearer suprageneric subdivision of the Osmiini than that suggested by Michener (2007): the genus <em>Chelostoma<\/em> clearly emerged as the sister group of all other osmiine bees (Figure 1) and thus deserves the same rank as the <em>Heriades<\/em> group and the <em>Osmia<\/em> group, often recognized as subtribes Heriadina and Osmiina, respectively (Engel, 2005; Ungricht et al., 2008).<\/p>\n<p><!--StartFragment--><\/p>\n<p style=\"text-align: left\">New suprageneric grouping of the Osmiini (Praz et al., 2008b):<\/p>\n<p><strong><em>Chelostoma<\/em> group<\/strong><\/p>\n<p style=\"padding-left: 30px\"><em>Chelostoma <\/em>Latreille<\/p>\n<p><strong><em>Heriades<\/em> group<\/strong><\/p>\n<p style=\"padding-left: 30px\"><em>Heriades <\/em>Spinola<\/p>\n<p style=\"padding-left: 30px\"><em>Hofferia <\/em>Tkalcu<\/p>\n<p style=\"padding-left: 30px\"><em>Othinosmia <\/em>Michener<\/p>\n<p style=\"padding-left: 30px\"><em>Protosmia <\/em>Ducke<\/p>\n<p style=\"padding-left: 30px\"><em>Stenoheriades <\/em>Tkalcu<\/p>\n<p style=\"padding-left: 30px\"><em>Xeroheriades <\/em>Griswold<\/p>\n<p><strong><em>Osmia<\/em> group<\/strong><\/p>\n<p style=\"padding-left: 30px\"><em>Ashmeadiella <\/em>Cockerell<\/p>\n<p style=\"padding-left: 30px\"><em>Atoposmia <\/em>Cockerell<\/p>\n<p style=\"padding-left: 30px\"><em>Haetosmia <\/em>Popov<\/p>\n<p style=\"padding-left: 30px\"><em>Hoplitis <\/em>(including <em>Stenosmia) <\/em>Klug<\/p>\n<p style=\"padding-left: 30px\"><em>Osmia <\/em>(including <em>Hoplosmia) <\/em>Panzer<\/p>\n<p style=\"padding-left: 30px\"><em>Wainia <\/em>Tkalcu<\/p>\n<p style=\"text-align: left\">A recent phylogenetic analysis of the genus <em>Chelostoma<\/em> on a worldwide scale (Sedivy et al. 2008) revealed a close relationship between the North American <em>C<\/em><em>helostoma (Prochelostoma) philadelphi <\/em>and the eastern Palaearctic <em>Chelostoma (Ceraheriades) petersi <\/em>(= <em>lamellum<\/em>) with respect to morphology, DNA sequences and flower preferences. Thus, their inclusion in two different subgenera is no longer justified and <em>Ceraheriades<\/em> is treated here as a junior synonym of <em>Prochelostoma <\/em>(A. M\u00fcller, unpublished).<\/p>\n<p style=\"text-align: left\"><em>Hoplitis oxypyga<\/em>, the only representative of the subgenus <em>Exanthocopa<\/em>, was known so far only in the male sex. Recently, females of this rare North African species were detected (A. M\u00fcller, unpublished). The females are morphologically so close to the females of <em>Hoplitis (Anthocopa) bisulca <\/em>that they can hardly be distinguished. Therefore, the subgenus <em>Exanthocopa<\/em> is synonymized here with the subgenus <em>Anthocopa<\/em> (A. M\u00fcller, unpublished).<\/p>\n<p style=\"text-align: left\">The species of the subgenus <em>Tergosmia<\/em> and and the only species of the subgenus <em>Ozbekosmia<\/em>, <em>Osmia avosetta<\/em>, share many morphological characteristics (Warncke, 1988b). In addition, the construction of their brood cells is unique in that the cells are composed of two layers of flower petals that sandwich a thin middle layer of mud (Rozen et al., 2010). Furthermore, all <em>Tergosmia<\/em> species as well as <em>Osmia avosetta<\/em> exhibit a strong or exclusive preference for Fabaceae as pollen hosts (see species accounts on this website). For these reasons, we propose to merge the subgenera <em>Tergosmia<\/em> and <em>Ozbekosmia<\/em> into one subgenus <em>Tergosmia<\/em> comb. nov. (A. M\u00fcller, unpublished).<\/p>\n<p style=\"text-align: left\">A recent study (Sedivy et al., 2013c) proposed to unite all <em>Hoplitis<\/em> species of the subgenera <em>Annosmia<\/em>, <em>Bytinskia<\/em>, <em>Coloplitis<\/em> and <em>Hoplitis<\/em> (the \u201e<em>Annosmia-Hoplitis<\/em> group\u201c) in a single subgenus <em>Hoplitis<\/em> because i) <em>Annosmia<\/em> is polyphyletic in its current circumscription, ii) the cleptoparasitic <em>Bytinskia<\/em> species have evolved from the same lineage as their <em>Annosmia<\/em> hosts, iii) the most basal clade of <em>Hoplitis<\/em> is morphologically and biologically intermediate between <em>Annosmia<\/em> and <em>Hoplitis<\/em>, and iv) <em>Coloplitis<\/em> is morphologically very close to <em>Annosmia<\/em>. An alternative solution to eliminate the polyphyly of <em>Annosmia<\/em> would be the establishment of a monotypic subgenus for the most basal species of the <em>Annosmia-Hoplitis<\/em> group, <em>Hoplitis bassana<\/em>, which, however, appears unjustified owing to the morphological similarity of <em>H. bassana<\/em> with <em>Annosmia<\/em>. Merging all four subgenera into a single large subgenus results in a monophylectic taxon, which is morphologically distinctive due to a conspicuous yellowish membrane below the lateral extremity of the labrum that is present in all species.<\/p>\n<p style=\"text-align: left\">The same study (Sedivy et al., 2013c) proposed to merge all <em>Hoplitis<\/em> species of the subgenera <em>Alcidamea<\/em>, <em>Cyrtosmia<\/em>, <em>Dasyosmia<\/em>, <em>Megalosmia<\/em>, <em>Monumetha<\/em> and <em>Prionohoplitis<\/em> (the \u201e<em>Alcidamea<\/em>\u201c group) into a single subgenus <em>Alcidamea <\/em>for the following reasons: i) <em>Alcidamea<\/em> is clearly polyphyletic in its current circumscription with all the other subgenera emerging from within this subgenus, and ii) the monophyly of <em>Prionohoplitis<\/em> is not supported. Uniting all six subgenera of the <em>Alcidamea<\/em> group results in a phylogenetically strongly supported clade, which, however, is characterized by only a few morphological characters such as the shape of the female clypus and the shapes of the male sterna 4-7.<\/p>\n<p style=\"text-align: left\">Sedivy et al. (2013c) also proposed to unite all <em>Hoplitis<\/em> species of the exclusively nearctic subgenera <em>Acrosmia, Hoplitina, Penteriades<\/em> and <em>Proteriades<\/em> (the \u201e<em>Proteriades<\/em> group\u201c) in a single subgenus <em>Proteriades<\/em> because i) neither <em>Proteriades<\/em> nor <em>Hoplitina<\/em> appear to be monophyletic taxa, ii) the distinction between the <em>Hoplitis<\/em> subgenera <em>Penteriades<\/em> and <em>Acrosmia<\/em> based on the presence or absence of hooked hairs on the female proboscis is not justified as two of the five <em>Acrosmia <\/em>species also possess such hooked hairs, and iii) the four subgenera closely resemble each other morphologically, especially in the female sex. Merging all four subgenera of the <em>Proteriades<\/em> group into a single large subgenus results in a morphologically and biologically well characterized taxon, whose monophyly is strongly supported by numerous morphological characters.<\/p>\n<p style=\"text-align: left\">A recent study (Haider et al., 2013) on the phylogeny and floral hosts of <em>Osmia<\/em> bees of the subgenera <em>Monosmia, Osmia<\/em> and <em>Orientosmia<\/em> revealed paraphyly of the largest subgenus <em>Osmia<\/em>. Given this paraphyly in conjunction with the pronounced morphological resemblance among the species of all three subgenera, the authors propose to to merge <em>Monosmia<\/em> (1 species),<em> Osmia<\/em> (25 species) and <em>Orientosmia<\/em> (3 species) into one single large subgenus <em>Osmia<\/em>.<\/p>\n<p style=\"text-align: left\">Based on a new molecular phylogeny of the bee genus <em>Osmia <\/em>with emphasis on North American <em>Melanosmia<\/em>, Rightmyer et al. (2013) erected a new Nearctic subgenus of <em>Osmia<\/em>, <em>Hapsidosmia<\/em>, for a single species formerly treated as a member of the subgenus <em>Melanosmia<\/em>. Furthermore, they synonymized the two Nearctic subgenera <em>Mystacosmia<\/em> and <em>Acanthosmioides<\/em> under <em>Melanosmia<\/em>. This study also clarified the phylogenetic relationships among the different subgenera of <em>Osmia<\/em>, e.g. <em>Osmia (Erythrosmia)<\/em> is newly supported as sister to <em>Osmia (Tergosmia).<\/em><\/p>\n<h5 style=\"text-align: left\"><a href=\"https:\/\/blogs.ethz.ch\/osmiini\/files\/2009\/09\/phylogenie13.jpg\"><img decoding=\"async\" class=\"alignnone size-full wp-image-2918\" src=\"https:\/\/blogs.ethz.ch\/osmiini\/files\/2009\/09\/phylogenie13.jpg\" alt=\"\" \/><\/a><\/h5>\n<h5 style=\"text-align: left\"><strong>Figure 1: <\/strong><span style=\"font-weight: normal\">Parsimony bootstrap consensus tree of the osmiine bees. All nodes with less than 50% bootstrap support were collapsed. After Praz et al., Molecular Phylogenetics and Evolution, 49, 185-197, 2008. With permission from Elsevier (<a href=\"http:\/\/www.sciencedirect.com\/science\/journal\/10557903\" target=\"_blank\">http:\/\/www.sciencedirect.com\/science\/journal\/10557903<\/a>).<\/span><\/h5>\n<p style=\"text-align: left\"><em><br \/>\n<\/em><\/p>\n","protected":false},"excerpt":{"rendered":"<p>Exposed brood cell of Hoplitis (Hoplitis) loti (Morawitz, 1867) on the surface of a rock; the cell consists of small pebbles cemented together with mud. Foto A. M\u00fcller. Phylogeny In a recent molecular phylogenetic study based on three nuclear genes and 95 osmiine bee species (Praz et al., 2008b), the core osmiine genera were found [&hellip;]<\/p>\n","protected":false},"author":1092,"featured_media":0,"parent":0,"menu_order":2,"comment_status":"open","ping_status":"closed","template":"","meta":{"footnotes":""},"class_list":["post-4","page","type-page","status-publish","hentry"],"_links":{"self":[{"href":"https:\/\/blogs.ethz.ch\/osmiini\/wp-json\/wp\/v2\/pages\/4","targetHints":{"allow":["GET"]}}],"collection":[{"href":"https:\/\/blogs.ethz.ch\/osmiini\/wp-json\/wp\/v2\/pages"}],"about":[{"href":"https:\/\/blogs.ethz.ch\/osmiini\/wp-json\/wp\/v2\/types\/page"}],"author":[{"embeddable":true,"href":"https:\/\/blogs.ethz.ch\/osmiini\/wp-json\/wp\/v2\/users\/1092"}],"replies":[{"embeddable":true,"href":"https:\/\/blogs.ethz.ch\/osmiini\/wp-json\/wp\/v2\/comments?post=4"}],"version-history":[{"count":0,"href":"https:\/\/blogs.ethz.ch\/osmiini\/wp-json\/wp\/v2\/pages\/4\/revisions"}],"wp:attachment":[{"href":"https:\/\/blogs.ethz.ch\/osmiini\/wp-json\/wp\/v2\/media?parent=4"}],"curies":[{"name":"wp","href":"https:\/\/api.w.org\/{rel}","templated":true}]}}