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10.3897_zookeys.101.1182.xml
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10.3897_zookeys.101.1182.xml
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<!DOCTYPE article PUBLIC "-//TaxonX//DTD Taxonomic Treatment Publishing DTD v0 20100105//EN" "tax-treatment-NS0.dtd"><article article-type="research-article" xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" xmlns:tp="http://www.plazi.org/taxpub">
<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">ZooKeys</journal-id>
<journal-title-group>
<journal-title>ZooKeys</journal-title>
</journal-title-group>
<issn pub-type="ppub">1313-2989</issn>
<issn pub-type="epub">1313-2970</issn>
<publisher>
<publisher-name>Pensoft Publishers</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.3897/zookeys.101.1182</article-id>
<title-group>
<article-title>Alocanthedon, a new subgenus of <italic>Chalicodoma</italic> from Southeast Asia (Hymenoptera, Megachilidae)</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" xlink:type="simple">
<name name-style="western">
<surname>Engel</surname>
<given-names>Michael S.</given-names>
</name>
<xref ref-type="aff" rid="A1">1</xref>
<uri content-type="lsid" xlink:type="simple">urn:lsid:zoobank.org:author:3714A7FF-E19E-495A-AAF9-98D2F597B757</uri></contrib>
<contrib contrib-type="author" xlink:type="simple">
<name name-style="western">
<surname>Gonzalez</surname>
<given-names>Victor H.</given-names>
</name>
<xref ref-type="aff" rid="A2">2</xref>
<uri content-type="lsid" xlink:type="simple">urn:lsid:zoobank.org:author:EC898C32-6E54-49A1-9A65-4A3F5B00E284</uri></contrib>
</contrib-group>
<aff id="A1"><label>1</label>Division of Entomology, Natural History Museum, and Department of Ecology & Evolutionary Biology, 1501 Crestline Drive – Suite 140, University of Kansas, Lawrence, Kansas 66049–2811, USA</aff>
<aff id="A2"><label>2</label>Department of Ecology & Evolutionary Biology, 1200 Sunnyside Avenue, Haworth Hall, University of Kansas, Lawrence, Kansas 66045; Current address: USDA-ARS Bee Biology & Systematics Laboratory, Utah State University, Logan, Utah 84322–5310, USA</aff>
<author-notes>
<fn fn-type="corresp">
<p>Corresponding author: Michael S. Engel (<email xlink:type="simple">msengel@ku.edu</email>), Victor H. Gonzalez (<email xlink:type="simple">victorgonzab@gmail.com</email>).</p>
</fn>
<fn fn-type="edited-by">
<p>Academic editor: Michael Ohl</p>
</fn>
</author-notes>
<pub-date pub-type="collection">
<year>2011</year>
</pub-date>
<pub-date pub-type="epub">
<day>31</day>
<month>5</month>
<year>2011</year>
</pub-date>
<issue>101</issue>
<fpage>51</fpage>
<lpage>80</lpage>
<history>
<date date-type="received">
<day>27</day>
<month>2</month>
<year>2011</year>
</date>
<date date-type="accepted">
<day>18</day>
<month>4</month>
<year>2011</year>
</date>
</history>
<permissions>
<copyright-statement>Michael S. Engel, Victor H. Gonzalez</copyright-statement>
<license license-type="creative-commons-attribution" xlink:href="http://creativecommons.org/licenses/by/3.0" xlink:type="simple">
<license-p>This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.</license-p>
</license>
</permissions>
<self-uri content-type="lsid" xlink:type="simple">urn:lsid:zoobank.org:pub:</self-uri>
<abstract>
<label>Abstract</label>
<p>A new subgenus, <italic><tp:taxon-name>Alocanthedon</tp:taxon-name></italic> Engel and Gonzalez <bold>subgen. n.</bold>, is described for five species of unusual Southeast Asian bees in the genus <italic><tp:taxon-name>Chalicodoma</tp:taxon-name></italic> Lepeletier de Saint Fargeau (<tp:taxon-name>Megachilinae</tp:taxon-name>: <tp:taxon-name>Megachilini</tp:taxon-name>). The subgenus is most noteworthy for the deep postgenal depression or furrow in males (bordered outwardly near the base of the mandible by a protuberant, thick lamella) and the presence of a dense patch of black setae posteriorly in the forewing medial cell (except in one species) [resembling the dense patch of setae among the submarginal cells of <italic><tp:taxon-name>Thrinchostoma</tp:taxon-name></italic> Saussure (<tp:taxon-name>Halictidae</tp:taxon-name>: <tp:taxon-name>Halictinae</tp:taxon-name>: <tp:taxon-name>Halictini</tp:taxon-name>)]. The subgenus is characterized and distinguished from the related <italic><tp:taxon-name>Callomegachile</tp:taxon-name></italic> Michener. A key to the following five species presently included in the subgenus is provided: <italic><tp:taxon-name>Chalicodoma aterrimum</tp:taxon-name></italic> (Smith), <italic><tp:taxon-name>Chalicodoma atratiforme</tp:taxon-name></italic> (Meade-Waldo) <bold>comb. n.</bold>, <italic><tp:taxon-name>Chalicodoma memecylonae</tp:taxon-name></italic> Engel <bold>sp. n.</bold>, <italic><tp:taxon-name>Chalicodoma odontophorum</tp:taxon-name></italic> Engel <bold>sp. n.</bold>, and <italic><tp:taxon-name>Chalicodoma apoicola</tp:taxon-name></italic> Engel <bold>sp. n.</bold> <italic><tp:taxon-name>Chalicodoma (Callomegachile) atratiforme sininsulae</tp:taxon-name></italic> (Cockerell) is newly placed in synonymy with <italic><tp:taxon-name>C</tp:taxon-name></italic>. (<italic><tp:taxon-name>C</tp:taxon-name></italic>.) <italic>fulvipenne</italic> (Smith). Species have been collected from <tp:taxon-name>Memecylaceae</tp:taxon-name> (Myrtales) and <tp:taxon-name>Fabaceae</tp:taxon-name> (Fabales). The phylogenetic relationships of <italic><tp:taxon-name>Alocanthedon</tp:taxon-name></italic> among other <tp:taxon-name>Megachilini</tp:taxon-name> are briefly elaborated upon.</p>
</abstract>
<kwd-group>
<label>Keywords</label>
<kwd>Apoidea</kwd>
<kwd>Anthophila</kwd>
<kwd>Megachilini</kwd>
<kwd><italic>Chalicodoma</italic></kwd>
<kwd>taxonomy</kwd>
<kwd>Southeast Asia</kwd>
<kwd>Megachilinae</kwd>
<kwd>phylogeny</kwd>
</kwd-group>
</article-meta>
</front>
<body>
<sec sec-type="Introduction"><title>Introduction</title>
<p>The Southeast Asian fauna of megachiline bees is particularly diverse but simultaneously poorly documented, underrepresented in collections, and with many species largely confused (especially in the diverse subgenus <italic><tp:taxon-name>Eutricharaea</tp:taxon-name></italic> Thomson of <italic><tp:taxon-name>Megachile</tp:taxon-name></italic> Latreille). In recent years several rather remarkable groups of species have been identified among this fauna (<xref ref-type="bibr" rid="B3">Baker and Engel 2006</xref>; <xref ref-type="bibr" rid="B7">Engel and Baker 2006</xref>). The purpose of the present contribution is to put on record yet another new group of peculiar Southeast Asian megachiline bees so that their names may be available for forthcoming works on the phylogeny and classification of the tribe and to bring them to the attention of melittologists working with this fauna. Herein, we propose a new subgeneric name, <italic><tp:taxon-name>Alocanthedon</tp:taxon-name></italic> Engel and Gonzalez subgen. n., for five unusual species in the Old World genus <italic><tp:taxon-name>Chalicodoma</tp:taxon-name></italic> Lepeletier de Saint Fargeau (<tp:taxon-name>Megachilinae</tp:taxon-name>: <tp:taxon-name>Megachilini</tp:taxon-name>) that are easily recognized by a unique combination of morphological characters in both sexes, especially in the male. Unlike all other <italic><tp:taxon-name>Chalicodoma</tp:taxon-name></italic>, males in the new subgenus have a deep postgenal depression or furrow bordered outwardly by a protuberant, thick lamella and a dense patch of black setae posteriorly in the forewing medial cell (except in one species). To date, the only other group of bees known to have a similar patch of black setae on the forewing is the sweat bee genus <italic><tp:taxon-name>Thrinchostoma</tp:taxon-name></italic> Saussure (<tp:taxon-name>Halictidae</tp:taxon-name>: <tp:taxon-name>Halictinae</tp:taxon-name>: <tp:taxon-name>Halictini</tp:taxon-name>). However, such a patch of setae is found among the submarginal cells of the forewing in both sexes of <italic><tp:taxon-name>Thrinchostoma</tp:taxon-name></italic>.</p>
<p>Following the classificatory proposal of <xref ref-type="bibr" rid="B9">Gonzalez (2008)</xref>, <italic><tp:taxon-name>Chalicodoma</tp:taxon-name></italic> is herein recognized in a narrower sense than that of (<xref ref-type="bibr" rid="B12">Michener (1962</xref>, 1965). As here understood, it includes all subgenera of Group 2 of <italic><tp:taxon-name>Megachile</tp:taxon-name></italic> s.l. <italic>sensu</italic> <xref ref-type="bibr" rid="B14">Michener (2007)</xref>, except for <italic><tp:taxon-name>Matangapis</tp:taxon-name></italic> Baker and Engel and those taxa with heriadiform or hoplitiform bodies with sparse pubescence (i.e., <italic><tp:taxon-name>Chelostomoda</tp:taxon-name></italic> Michener and related groups). The relationships among the seven subgenera presently recognized in <italic><tp:taxon-name>Chalicodoma</tp:taxon-name></italic> (<italic><tp:taxon-name>Callomegachile</tp:taxon-name></italic> Michener, <italic><tp:taxon-name>Cestella</tp:taxon-name></italic> Pasteels, <italic><tp:taxon-name>Chalicodoma</tp:taxon-name></italic> s.str., <italic><tp:taxon-name>Cuspidella</tp:taxon-name></italic> Pasteels, <italic><tp:taxon-name>Gronoceras</tp:taxon-name></italic> Cockerell, <italic><tp:taxon-name>Largella</tp:taxon-name></italic> Pasteels, and <italic><tp:taxon-name>Pseudomegachile</tp:taxon-name></italic> Friese) need to be studied in detail. Such subgenera are highly diverse, morphologically heterogeneous, and as distinct as many genera of bees. Future work may show that generic status might be warranted for some or all.</p>
<p>We also briefly discuss the phylogenetic relationships of <italic><tp:taxon-name>Alocanthedon</tp:taxon-name></italic> among the subgenera of <italic><tp:taxon-name>Chalicodoma</tp:taxon-name></italic> and provide an overview of the morphological diversity of the related <italic><tp:taxon-name>Callomegachile</tp:taxon-name></italic> as well as taxonomic notes on other rare Southeast Asian species.<!--PageBreak--></p>
</sec>
<sec sec-type="materials|methods"><title>Materials and Methods</title>
<p>Morphological terminology follows that of <xref ref-type="bibr" rid="B6">Engel (2001)</xref> and <xref ref-type="bibr" rid="B14">Michener (2007)</xref> while the format for the descriptions loosely follows those of <xref ref-type="bibr" rid="B13">Michener (1965)</xref>, <xref ref-type="bibr" rid="B3">Baker and Engel (2006)</xref>, <xref ref-type="bibr" rid="B7">Engel and Baker (2006)</xref>, and <xref ref-type="bibr" rid="B10">Gonzalez et al. (2010)</xref> for megachiline bees. Institutional acronyms used herein are: SEMC, Snow Entomological Collection, Division of Entomology, University of Kansas Natural History Museum, Lawrence, Kansas, USA; NHML, Department of Entomology, The Natural History Museum, London, UK; and NSMT, National Science Museum (Natural History Museum), Tokyo, Japan.</p>
<p>To explore the phylogenetic relationships of these rare bees, we used the morphological characters and data set of <xref ref-type="bibr" rid="B9">Gonzalez (2008)</xref> for the phylogeny of <tp:taxon-name>Megachilini</tp:taxon-name> (six outgroup species, 107 ingroup species, and 231 characters). This data set is available from the authors or can be obtained from the unpublished dissertation, which is freely accessed throughout the University of Kansas libraries (http://kuscholarworks.ku.edu/dspace/handle/1808/4187). Because the specimens were not dissected, 31 characters, corresponding to some traits of the labiomaxillary complex, sixth sternum of females, sting apparatus, and male hidden sterna could not be coded. Parsimony analyses were performed in an Intel® CoreTM i3 processor using Tree analysis using New Technology (TNT; <xref ref-type="bibr" rid="B8">Goloboff et al. 2003</xref>). All characters were treated as unweighted, unordered, and nonadditive to allow characters to reverse freely and examine possible hypotheses of evolutionary relationships. Tree search in TNT was done by implementing sectorial searches (SS) with tree drifting (TD) and tree fusing (TF) and ratchet runs with TD and TF. We used the following search: keep a maximum of 10000 random trees, 500 random addition sequences, and 1000 ratchet iterations, including 100 cycles of TD and 100 rounds of TF per iteration. Trees were visualized and printed using Winclada (<xref ref-type="bibr" rid="B16">Nixon 1999</xref>).</p>
</sec>
<sec sec-type="Systematics"><title>Systematics</title>
<p><bold>Tribe <tp:taxon-name>Megachilini</tp:taxon-name> Latreille</bold></p>
<p><bold>Genus</bold> <italic><tp:taxon-name>Chalicodoma</tp:taxon-name></italic> <bold>Lepeletier de Saint Fargeau</bold></p>
<tp:taxon-treatment>
<tp:nomenclature>
<tp:taxon-name>
<tp:taxon-name-part taxon-name-part-type="genus">Alocanthedon</tp:taxon-name-part>
<object-id xlink:type="simple">urn:lsid:zoobank.org:act:42B75DAA-6A32-4AAF-8209-4E1BD5E56B22</object-id>
<object-id xlink:type="simple">http://species-id.net/wiki/Alocanthedon</object-id>
</tp:taxon-name>
<tp:taxon-authority>Engel and Gonzalez</tp:taxon-authority>
<tp:taxon-status xlink:type="simple">subgen. n.</tp:taxon-status>
</tp:nomenclature>
<tp:treatment-sec sec-type="Type species"><title>Type species.</title>
<p><italic><tp:taxon-name>Chalicodoma (Alocanthedon) odontophorum</tp:taxon-name></italic> Engel, sp. n.</p>
</tp:treatment-sec>
<tp:treatment-sec sec-type="Diagnosis"><title>Diagnosisn.</title>
<p>Large (ca. 20–25 mm), black, parallel-sided megachilines resembling some large, black species of <italic><tp:taxon-name>Callomegachile</tp:taxon-name></italic> such as those of the Eumegachilana group (<xref ref-type="fig" rid="F1">Figs. 1</xref>, <xref ref-type="fig" rid="F2">7</xref>, <xref ref-type="fig" rid="F3">10</xref>, <xref ref-type="fig" rid="F5">17</xref>, <xref ref-type="fig" rid="F6">23</xref>, <xref ref-type="fig" rid="F8">35</xref>, <xref ref-type="fig" rid="F9">41</xref>, <xref ref-type="fig" rid="F10">44</xref>) but in males with juxtamandibular flange or lamella and deep postgenal depression (<xref ref-type="fig" rid="F4">Figs. 15</xref>, <xref ref-type="fig" rid="F4">16</xref>, <xref ref-type="fig" rid="F5">19</xref>); with oblique carina or lamella medially on disc of procoxa; with modified pro- (<xref ref-type="fig" rid="F1">Figs. 5</xref>, <xref ref-type="fig" rid="F3">14</xref>, <xref ref-type="fig" rid="F5">21</xref>, <xref ref-type="fig" rid="F8">38</xref>) and mesotarsi; with <!--PageBreak-->dense cluster of short, black setae forming a conspicuous spot in the posterior half of the forewing medial cell (<xref ref-type="fig" rid="F1">Figs. 4</xref>, <xref ref-type="fig" rid="F3">11</xref>, <xref ref-type="fig" rid="F8">39</xref>, except in <italic><tp:taxon-name>Chalicodoma memecylonae</tp:taxon-name></italic>); with five exposed metasomal sterna and with apical margin of sternum six typically exposed; with basally large gonocoxae (in comparison with those of the large species of Eumegachilana group where they are disproportionately small for the size of the bee) that are attenuate, divergent, upcurved at apices, greatly exceeding apices of penis valves, and without setae apically (<xref ref-type="fig" rid="F7">Figs. 26–34</xref>); with penis valves slightly expanded apically; and in females with broad, not porrect, 4-toothed mandibles, with the outer surface dull, minutely roughened, and punctate; with clypeus slightly concave to V-shaped epistomal sulcus basally; with the pretarsal claws simple, basally with short, stout seta; and with the metasoma parallel-sided.</p>
</tp:treatment-sec>
<tp:treatment-sec sec-type="Description"><title>Description.</title>
<p><italic>Male</italic>: Mandible tridentate, with median inferior swelling or protuberance, basal projection absent; torulus with distinct lamella on upper half of inner margin; first flagellomere wider than long, length much less than one-half length of second flagellomere. Preoccipital carina distinct, continuing from vertex to gena; postgena bordering hypostoma with deep depression, posteriorly bordered by dense brush of black setae and long, finer patch of white setae, depression with bordering juxtamandibular flange or lamella near anterior mandibular base and bordering compound eye (<xref ref-type="fig" rid="F4">Figs. 15, 16</xref>). Forewing with dense cluster of short, black, simple, lanceolate setae forming a conspicuous spot in posterior half of medial cell (<xref ref-type="fig" rid="F1">Figs. 4</xref>, <xref ref-type="fig" rid="F3">11</xref>, <xref ref-type="fig" rid="F8">39</xref>: except in <italic><tp:taxon-name>Chalicodoma memecylonae</tp:taxon-name></italic>, <xref ref-type="fig" rid="F5">Fig. 20</xref>). Pronotal lobe with strong carina; procoxa with apical, anteriorly-directed spine, with oblique carina or lamella medially on procoxal surface, anterior surface without rufescent bristles; protarsus greatly modified, flattened and expanded in species-specific forms, with variegated fringes; meso- and metafemora slightly swollen; mesotibial spur present; mesotarsi flattened with concave inner basal surfaces, posterior border variously modified; pretarsal claws symmetrical, cleft; mesobasitarsus weakly to strongly arched basally, with variously developed basal ventral concavity; metatarsi unmodified, slender; metasomal tergum VI with preapical carina gently concave medially (depth of concavity varies dramatically across species), without teeth; metasomal sternum V exposed, densely pubescent; apex of metasomal sternum VI normally exposed, densely pubescent; gonocoxae relatively large (by comparison with those of the large species of Eumegachilana group where they are disproportionately small for the size of the bee), attenuate, divergent, apices upcurved, greatly exceeding apices of penis valves, without setae apically (<xref ref-type="fig" rid="F7">Figs. 26–34</xref>); volsella pointed, articulate, distinguished as separated sclerite; penis valves slightly expanded apically.</p>
<p><italic>Female</italic>: Mandible broad, not porrect or elongate (similar to <italic><tp:taxon-name>Callomegachile</tp:taxon-name></italic>) except somewhat elongate in <italic><tp:taxon-name>Chalicodoma odontophorum</tp:taxon-name></italic>, 4-toothed (third tooth reduced in <italic><tp:taxon-name>Chalicodoma aterrimum</tp:taxon-name></italic>), without cutting edge, outer surface dull, minutely roughened and coarsely and shallowly punctate. Clypeus not protuberant, not covering labral base; first flagellomere wider than long, about one-half length of second flagellomere. Pretarsal claws simple, symmetrical, basally with short, stout seta. Metasoma parallel-sided; tergum VI very weakly concave in profile, with pubescence as on preceding terga; sternum VI with scopal setae as on preceding sterna, without bare rim; sterna without apical pubescent bands.<!--PageBreak--></p>
</tp:treatment-sec>
<tp:treatment-sec sec-type="Etymology"><title>Etymology.</title>
<p>The new genus-group name is a combination of the Greek words <italic>alokos</italic> (meaning, “furrow”) and <italic>anthedon</italic> (meaning, “bee”), and is a reference to the deep postgenal furrow universally in males of this lineage. The name is feminine.</p>
</tp:treatment-sec>
<tp:treatment-sec sec-type="Included species"><title>Included species.</title>
<p>In addition to the type species the subgenus includes the following taxa: <italic><tp:taxon-name>Chalicodoma aterrimum</tp:taxon-name></italic> (<xref ref-type="bibr" rid="B17">Smith 1862</xref>), <italic><tp:taxon-name>Chalicodoma atratiforme</tp:taxon-name></italic> (Meade-Waldo, 1914), <italic><tp:taxon-name>Chalicodoma memecylonae</tp:taxon-name></italic> Engel sp. n., and <italic><tp:taxon-name>Chalicodoma apoicola</tp:taxon-name></italic> Engel sp. n. (<xref ref-type="table" rid="T1">Table 1</xref>).</p>
</tp:treatment-sec>
<tp:treatment-sec sec-type="Comments"><title>Comments.</title>
<p>The five currently included species are superficially quite similar, but for the hyaline or dark fuscous, rather than yellow, wings in <italic><tp:taxon-name>Chalicodoma aterrimum</tp:taxon-name></italic> and <italic><tp:taxon-name>Chalicodoma apoicola</tp:taxon-name></italic>, accordingly the new species are described in reference to the type species rather than repeat largely identical blocks of text. Owing the presence of species of <italic><tp:taxon-name>Alocanthedon</tp:taxon-name></italic> in the Philippines as well as across Wallace’s Line in Sulawesi it is likely that as of yet unrecognized taxa for the subgenus may occur in places throughout Indonesia and Malaysian Borneo, and perhaps as far East as Irian Jaya. Additional collecting of bees, for all groups, is needed across all of these islands.</p>
</tp:treatment-sec>
<tp:taxon-treatment>
<tp:nomenclature>
<tp:taxon-name>
<tp:taxon-name-part taxon-name-part-type="genus">Chalicodoma</tp:taxon-name-part>
<tp:taxon-name-part taxon-name-part-type="subgenus"> (Alocanthedon) </tp:taxon-name-part>
<tp:taxon-name-part taxon-name-part-type="species">odontophorum</tp:taxon-name-part>
<object-id xlink:type="simple">urn:lsid:zoobank.org:act:1DA0608B-A874-4338-BDAE-FECE7D9E7FEA</object-id>
<object-id xlink:type="simple">http://species-id.net/wiki/Chalicodoma_(Alocanthedon)_odontophorum</object-id>
</tp:taxon-name>
<tp:taxon-authority>Engel</tp:taxon-authority>
<tp:taxon-status xlink:type="simple">sp. n.</tp:taxon-status>
<xref ref-type="fig" rid="F1">Figs 1</xref>
<xref ref-type="fig" rid="F2">9</xref>
<xref ref-type="fig" rid="F7">26–28</xref>
</tp:nomenclature>
<tp:treatment-sec sec-type="Holotype"><title>Holotype.</title>
<p>Thailand: ♂, Sakaerat DDF [Nakhon Ratchasima Province, Sakaerat Environmental Research Area, ca. 40 km South Nakhon Ratchasima], 20 June 1995 (SEMC).</p>
</tp:treatment-sec>
<tp:treatment-sec sec-type="Paratypes"><title>Paratypes.</title>
<p>Thailand: ♀, Sakaerat DDF [Nakhon Ratchasima Province, Sakaerat Environmental Research Area, ca. 40 km South Nakhon Ratchasima], 17 June 1995 (SEMC); 1♂, Siam (SEMC).</p>
<p>Myanmar: 1♀, Middle Tenasserim, Thaungyin Valley, 5.93 [May 1893], C.T. Bingham (NHML).</p>
</tp:treatment-sec>
<tp:treatment-sec sec-type="Diagnosis"><title>Diagnosis.</title>
<p>Both sexes of this species have yellow forewings with grayish hyaline apex. The male can be easily distinguished by the clypeus densely covered by long, appressed, apically-directed setae obscuring integument (<xref ref-type="fig" rid="F1">Fig. 2</xref>) and the shape of the modified protarsi (<xref ref-type="fig" rid="F1">Fig. 5</xref>). The female of this species is recognized by the clypeus with a pronounced, erect, medioapical tubercle (<xref ref-type="fig" rid="F2">Fig. 8</xref>), the elongate mandibles and the labrum, with apical margin medially convex and apical fringe of erect setae separated from labral apical margin by at least one median ocellar diameter or slightly more.</p>
</tp:treatment-sec>
<tp:treatment-sec sec-type="Description"><title>Description.</title>
<p>As for the subgenus with the following additions: <italic>Male</italic>: Total body length 20 mm; forewing length 13.3 mm. Head broader than long (width 5.3 mm, length 4.0 mm); inner orbits of compound eyes slightly divergent below; intertorular distance 1.6 times torulorbital distance; interocellar distance 1.8 times median ocellar diameter, slightly shorter than ocellocular distance; ocelloccipital distance 4.2 times median ocellar diameter; compound eye about 2.2 times longer than wide, 1.2 times wider than gena in profile. Mandible with three teeth, with prominent, broad median inferior protuberance bearing dense, short, black setae. Juxtamandibular flange about <!--PageBreak-->twice as long as posterior height. Labrum rectangular, with apical row of stiff, erect, long setae. Clypeus broad, width more than three times medial length. Scape length about 2.5 times width; first flagellomere short, nearly one-third length of second flagellomere; remaining flagellomeres all much longer than wide, apicalmost flagellomere with broadly rounded apex, not tapering. Mesoscutum with distinct notauli and parapsidal lines. Procoxal spine elongate, with weak depression between spine and medial, transverse carina of procoxa, posterobasally setose on spine, anterior surface not setose; protibia with strong, outer, posterior carina running along apical three-quarters of length, apically produced into small posteriorly-directed spine before continuing transversely across apex to outer, anterior border where it forms definite ridge but not carinate, apical anterior surface faintly depressed in profile view; protarsus modified as in <xref ref-type="fig" rid="F1">figure 5</xref>; meso- and metafemora somewhat swollen; mesotibial spur curved, with bluntly rounded apex; mesobasitarsus with inner surface concave basally, posterior border along concavity relatively straight; outer metatibial spur blunt at apex (not tapering to acute apex); pretarsal claws long, curved, apically cleft. Postgradular depressions deeper than in female; terga II–V with apical transverse ridge (caudad postgradular depression), somewhat sinuate laterally, weak medially on terga II–IV; preapical carina of sixth tergum produced, weakly and broadly concave medially (<xref ref-type="fig" rid="F1">Fig. 6</xref>). Genitalia as in figures <xref ref-type="fig" rid="F7">26–28</xref>.</p>
<p>Integument black throughout except tegula, legs, and metasomal sterna largely dark reddish brown (nearly black in many areas), and expansion of protarsi more translucent brown. Wings orange-yellow except apical margin of forewing and apical and posterior margins of hind wing grayish hyaline (<xref ref-type="fig" rid="F1">Fig. 1</xref>); venation ferruginous to orange-yellow.</p>
<p>Mandible with outer surface dull, irregularly punctate and microreticulate; labrum strongly imbricate and impunctate; clypeus with small, contiguous punctures, with thin mediolongitudinal impunctate area; supraclypeal area and face below ocelli with small, contiguous punctures, punctures becoming more irregular at level of median ocellus; area between ocelli with small, contiguous punctures; ocellocular area with<!--PageBreak--> somewhat larger, coarser punctures separated by areas of finely imbricate integument; vertex with coarse, shallow punctures separated by a puncture width or less, integument between finely imbricate, punctures becoming more shallow and faint toward preoccipital carina; upper gena with irregular, elongate punctures separated by finely imbricate integument, remainder of gena and posterior postgena with more regular <!--PageBreak-->punctures separated by a puncture width or less, integument otherwise finely imbricate; postgenal surface inside of deep postgenal depression with scattered minute punctures separated by faintly imbricate to smooth integument; outer surface of juxtamandibular lamella with irregular punctures and imbricate integument. Pronotum imbricate, with small punctures separated by a puncture width or less; mesoscutum anteriorly and medially transversely wrinkled with irregular punctures, such integument blending laterally outside of parapsidal lines and posteriorly to coarsely punctate, punctures separated by a puncture width or less, those outside of parapsidal lines somewhat smaller and more regularly defined than those posteriorly, integument between punctures finely imbricate; tegula finely imbricate and minutely punctate, punctures separated by less than a puncture width, except along outer rim impunctate; axillae and mesoscutellum coarsely and contiguously punctate except mediobasally on mesoscutellum with punctures smaller and gradually becoming separated by a puncture width or less; metanotum imbricate with small punctures separated by a puncture width or less; pleura coarsely and contiguously punctate, those punctures along omaular ridge and ventrally somewhat dorsoventrally elongate, giving ventral surface a somewhat dorsoventrally rugulose appearance; declivitous basal area of propodeum with single row of coarse alveolae along extreme basal border, row interrupted medially, otherwise surface imbricate and impunctate; lateral surface imbricate with small punctures separated by less than a puncture width, gradually becoming more widely spaced posteriorly and on posterior surface. Anterior-facing surface of first metasomal tergum finely imbricate, dorsal-facing surface imbricate with small punctures separated by a puncture width or less, nearly contiguous in most areas; remaining terga sculptured as on dorsal-facing surface of first metasomal tergum; terga with narrow impunctate apical rims; sterna smooth to finely imbricate, with small punctures separated by less than a puncture width.</p>
<p>Pubescence generally dark fuscous to black except as follows: clypeus, supraclypeal area, and face outside of antennal toruli with dense, long, minutely-branched, tawny to white setae, largely obscuring the integument, those on clypeus more strongly tawny and largely appressed and apically directed; thin fringe of short, fine, silvery white setae along outer border of patch of black setae on median inferior protuberance and running proximally to mandibular condyle; dense patch of white pubescence immediately posterior to dense patch of somewhat shorter black setae at posterior end of postgenal depression, white patch largely occupying area at meeting of postgena and ventral area of gena; white, long setae on pronotal lateral surfaces, outer borders of propleura, outer base of procoxa, and preomaular area; white and black setae arranged on protarsus and apex of protibia as in <xref ref-type="fig" rid="F1">figure 5</xref>; and white setae on propodeal lateral surface, most dense ventrally, near metacoxa; wing setae generally yellow or tawny yellow except dense cluster of short, black setae forming a conspicuous spot in posterior half of forewing medial cell (<xref ref-type="fig" rid="F1">Fig. 4</xref>).</p>
<p><italic>Female</italic>: As for the male except in usual sexual differences and as follows: Total body length 19–21 mm; forewing length 14.9 mm. Head broader than long (width 6.1 mm, length 4.5 mm); intertorular distance about as long as torulorbital distance;<!--PageBreak--> interocellar distance 2.4 times median ocellar diameter, 0.8 times ocellocular distance; ocelloccipital distance 6.9 times median ocellar diameter; compound eye about twice as long as wide, slightly narrower than gena in profile. Mandible with four teeth; body of mandible elongate, basal section distinctly longer than apical, dentate margin (<xref ref-type="fig" rid="F2">Fig. 8</xref>). Labrum rectangular, with apical row of stiff, erect, long setae separated from apical margin by at least one median ocellar diameter; apical margin medially convex. Clypeus with pronounced, medioapical tubercle (all species have a minute point here, but not distinctly tuberculate as in this species) (<xref ref-type="fig" rid="F2">Fig. 8</xref>). Scape length more than three times width; first flagellomere short, about one-half length of second flagellomere; re<!--PageBreak-->maining flagellomeres all about twice as long as wide. Procoxae, tibiae, tarsi, and spurs unmodified; pretarsal claws long, curved, simple. Postgradular depressions faint; terga without transverse ridges, with slight lateral swellings on terga II and III.</p>
<p>Clypeus imbricate with coarse, shallow punctures separated by less than a puncture width in basal half except medially such punctures restricted to basal border; supraclypeal area with smaller coarse punctures than those on clypeus, punctures separated by less than a puncture width; face below ocelli with small, contiguous punctures, punctures becoming more irregular at level of median ocellus; ocellocular area with somewhat larger, coarser punctures separated by areas of finely imbricate integument; vertex with coarse, shallow punctures separated by 0.5–1.5 times a puncture width, integument between finely imbricate, punctures becoming more shallow and faint toward preoccipital carina; postgena strongly rugulose; hypostoma imbricate. Mesoscutum anteriorly and medially transversely wrinkled, more weakly so than in male, with irregular punctures, such integument blending laterally and posteriorly to faintly coarsely punctate, punctures separated by less than a puncture width, those outside of parapsidal lines sparse, integument between punctures finely imbricate; axillae and mesoscutellum strongly coarsely and contiguously punctate except mediobasally on mesoscutellum with punctures smaller and gradually becoming separated by a puncture width or less. Dorsal-facing surface of first metasomal tergum imbricate with small punctures separated by a puncture width or less medially and nearly contiguous laterally; remaining terga sculptured as on lateral areas of dorsal-facing surface of first metasomal tergum.</p>
<p>Usual sex differences in setation; pubescence dark fuscous to black except microtrichia on inner surface of mandible dark golden and small dirty white patch on lateral surface of propodeum near metacoxa; clypeus and supraclypeal area not obscured by dense pubescence; ventral surfaces of mesepisternum, coxae, trochanters, base of femora, anterior margins of metatibia and metafemur, and sternal scopa with capitate setae.</p>
<table-wrap position="float" orientation="portrait" id="T1">
<label>Table 1.</label>
<caption><p>Summary of currently included species in subgenus <italic><tp:taxon-name>Alocanthedon</tp:taxon-name></italic>.</p></caption>
<table>
<tbody>
<tr>
<th rowspan="1" colspan="1">Species</th>
<th rowspan="1" colspan="1">General distribution</th>
</tr>
<tr>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Chalicodoma apoicola</tp:taxon-name></italic> Engel sp. n.</td>
<td rowspan="1" colspan="1">Philippines: Mindanao: Davao del Sur</td>
</tr>
<tr>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Chalicodoma aterrimum</tp:taxon-name></italic> (Smith, 1862)</td>
<td rowspan="1" colspan="1">Sulawesi</td>
</tr>
<tr>
<td rowspan="3" colspan="1"><italic><tp:taxon-name>Chalicodoma atratiforme</tp:taxon-name></italic> (Meade-Waldo, 1914), comb. n.</td>
<td rowspan="1" colspan="1">Myanmar: Tanintharyi Division (= Tenasserim)</td>
</tr>
<tr>
<td rowspan="1" colspan="1">Thailand: Uthai Thani Province</td>
</tr>
<tr>
<td rowspan="1" colspan="1">Malaysia: Pahang State, Negeri Sembilan State</td>
</tr>
<tr>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Chalicodoma memecylonae</tp:taxon-name></italic> Engel sp. n.</td>
<td rowspan="1" colspan="1">Malaysia: Penang State, Perak State, Selangor State, Pahang State</td>
</tr>
<tr>
<td rowspan="2" colspan="1"><italic><tp:taxon-name>Chalicodoma odontophorum</tp:taxon-name></italic> Engel sp. n.</td>
<td rowspan="1" colspan="1">Thailand: Nakhon Ratchasima Province</td>
</tr>
<tr>
<td rowspan="1" colspan="1">Myanmar: Tanintharyi Division</td>
</tr>
</tbody>
</table>
</table-wrap>
<fig id="F1" position="float" orientation="portrait">
<label>Figures 1–6.</label>
<caption><p>Photomicrographs of male of <italic><tp:taxon-name>Chalicodoma</tp:taxon-name></italic> (<italic><tp:taxon-name>Alocanthedon</tp:taxon-name></italic>) <italic>odontophorum</italic> Engel, sp. n. <bold>1</bold> Dorsal habitus <bold>2</bold> Facial aspect <bold>3</bold> Dorsal aspect of mesoscutum and mesoscutellum <bold>4</bold> Detail of forewing medial cell <bold>5</bold> Protarsus, pro-pretarsus, and protibia <bold>6</bold> Dorsal aspect of metasomal terga V and VI.</p></caption>
<graphic xlink:href="ZooKeys-101-051-g001.jpg" position="float" orientation="portrait" xlink:type="simple"/>
</fig>
<fig id="F2" position="float" orientation="portrait">
<label>Figures 7–9.</label>
<caption><p>Photomicrographs of female of <italic><tp:taxon-name>Chalicodoma</tp:taxon-name></italic> (<italic><tp:taxon-name>Alocanthedon</tp:taxon-name></italic>) <italic>odontophorum</italic> Engel, sp. n. <bold>7</bold> Dorsal habitus <bold>8</bold> Facial aspect (with expanded detail of clypeus, arrow indicating medioapical tubercle) <bold>9</bold> Dorsal aspect of mesoscutum and mesoscutellum.</p></caption>
<graphic xlink:href="ZooKeys-101-051-g002.jpg" position="float" orientation="portrait" xlink:type="simple"/>
</fig>
</tp:treatment-sec>
<tp:treatment-sec sec-type="Etymology"><title>Etymology.</title>
<p>The specific epithet is a combination of the Greek word <italic>odontos</italic> (meaning, “teeth”) and suffix –<italic>phor</italic> (meaning, “carry”).</p>
</tp:treatment-sec>
<tp:treatment-sec sec-type="Floral records"><title>Floral records.</title>
<p>The holotype and paratype from Sakaerat were captured at flowers of <italic><tp:taxon-name>Sindora siamensis</tp:taxon-name></italic> Teijsman & Miquel (Fabales: <tp:taxon-name>Fabaceae</tp:taxon-name>: <tp:taxon-name>Caesalpinioideae</tp:taxon-name>: Detarieae).</p>
</tp:treatment-sec>
</tp:taxon-treatment>
<tp:taxon-treatment>
<tp:nomenclature>
<tp:taxon-name>
<tp:taxon-name-part taxon-name-part-type="genus">Chalicodoma</tp:taxon-name-part>
<tp:taxon-name-part taxon-name-part-type="subgenus"> (Alocanthedon) </tp:taxon-name-part>
<tp:taxon-name-part taxon-name-part-type="species">apoicola</tp:taxon-name-part>
<object-id xlink:type="simple">urn:lsid:zoobank.org:act:A7C422CE-C4EB-4C1A-A8B4-541DB0B1E39C</object-id>
<object-id xlink:type="simple">http://species-id.net/wiki/Chalicodoma_(Alocanthedon)_apoicola</object-id>
</tp:taxon-name>
<tp:taxon-authority>Engel</tp:taxon-authority>
<tp:taxon-status xlink:type="simple">sp. n.</tp:taxon-status>
<xref ref-type="fig" rid="F3">Figs 10</xref>
<xref ref-type="fig" rid="F4">16</xref>
</tp:nomenclature>
<tp:treatment-sec sec-type="Holotype"><title>Holotype.</title>
<p>Philippines: ♂, Mindanao, Tagurano [Davao del Sur, near Mt. Apo and Mt. Apo National Park], Davao City, 25–26.vi.1977 [25–26 June 1977], Y. Kurosawa (NSMT).</p>
</tp:treatment-sec>
<tp:treatment-sec sec-type="Diagnosis"><title>Diagnosisn.</title>
<p>The new species is most similar to <italic><tp:taxon-name>Chalicodoma aterrimum</tp:taxon-name></italic>, but can be distinguished from this and other <italic><tp:taxon-name>Alocanthedon</tp:taxon-name></italic> by the following combination of traits: clypeus covered with dense, long, appressed, reddish setae obscuring integument (<xref ref-type="bibr" rid="B3">Fig. 12</xref>)<!--PageBreak-->; forewing dark fuscous with black venation; dense black setal patch present posteriorly in forewing medial cell (<xref ref-type="fig" rid="F3">Fig. 11</xref>); dorsal-facing surface of first metasomal tergum with large, anterobasal areas of impunctate and imbricate integument; terga with strong transverse ridges on non-depressed, postgradular discs, carinate on terga II and III; sixth metasomal tergum deeply concave medioapically; medioapical apical margin of second metasomal sternum convex as short, broad extension; protibial apical outer surface distinctly depressed; outer anterior margin of protibia with dense fringe of long black setae; and unique protarsal shape and setation (<xref ref-type="fig" rid="F3">Fig. 14</xref>).</p>
</tp:treatment-sec>
<tp:treatment-sec sec-type="Description"><title>Description.</title>
<p>As described for <italic><tp:taxon-name>Chalicodoma odontophorum</tp:taxon-name></italic> (<italic>vide supra</italic>) except as follows: <italic>Male</italic>: Total body length 24.6 mm; forewing length 19.3 mm. Head broader than long (width 7.4 mm, length 5.5 mm); intertorular distance 1.9 times torulorbital distance; interocellar distance 1.5 times median ocellar width, 1.1 times ocellocular distance; ocelloccipital distance 3.5 times median ocellar width; compound eye about twice as long as wide, about as wide as gena in profile. Protibia with outer posterior carina run<!--PageBreak-->ning along apical three-quarters of length, apically produced into small posteriorly-directed spine before continuing transversely across apex to outer, anterior border where it forms a carinate ridge for short distance along depression, apical anterior surface distinctly depressed; protarsus modified as in <xref ref-type="fig" rid="F3">figure 14</xref>; mesotibial spur relatively straight; mesobasitarsus with inner surface deeply concave basally, posterior border along concavity notched, such that there is a posterior protuberance bordering the concavity. Metasomal terga II–V with apical transverse ridge (caudad postgradular depression), somewhat sinuate laterally, distinctly carinate on terga II–III except medially, strongly ridged on tergum IV, weak on tergum V; carina of sixth tergum produced, medioapical margin of carina of sixth metasomal tergum strongly and deeply concave (<xref ref-type="fig" rid="F3">Fig. 10</xref>).</p>
<p>Wings dark fuscous, infumate (<xref ref-type="fig" rid="F3">Fig. 10</xref>); venation black.</p>
<p>Mesoscutum anteriorly and medially transversely somewhat wrinkled (not as strongly so as in <italic><tp:taxon-name>Chalicodoma odontophorum</tp:taxon-name></italic>) with irregular punctures, such integument blending laterally outside of parapsidal lines and posteriorly to coarsely punctate, punctures separated by less than a puncture width, nearly contiguous in many areas, those punctures outside of parapsidal lines somewhat smaller and more regularly defined than those posteriorly, integument between punctures finely imbricate; lateral surface of propodeum imbricate with small punctures separated by a puncture width or less, gradually becoming more widely spaced posteriorly and on posterior surface. Dorsal-facing surface of first metasomal tergum imbricate with small punctures separated by a puncture width or less, nearly contiguous in most areas, with large laterobasal areas of impunctate (and asetose) and more distinctly imbricate integument; remaining terga sculptured as on dorsal-facing surface of first metasomal tergum, although punctures typically more tightly packed.</p>
<p>Pubescence generally dark fuscous to black except as follows: clypeus and face outside of antennal toruli with dense, long, minutely-branched, reddish setae, largely obscuring the integument, those on clypeus more strongly reddish and largely appressed and apically directed; supraclypeal area with similar setae to those on face except more tawny in color (<xref ref-type="fig" rid="F3">Fig. 12</xref>); wing setae generally fine and black, dense cluster of short, black setae forming a conspicuous spot in posterior half of forewing medial cell (<xref ref-type="fig" rid="F3">Fig. 11</xref>).</p>
<p><italic>Female</italic>: Unknown.</p>
</tp:treatment-sec>
<tp:treatment-sec sec-type="Etymology"><title>Etymology.</title>
<p>The specific epithet is a combination of Mount Apo and the Latin suffix –<italic>cola</italic>, meaning “dweller”. The name is treated as a noun in apposition.</p>
</tp:treatment-sec>
<tp:treatment-sec sec-type="Comments"><title>Comments.</title>
<p>Permission to dissect the holotype and only known specimen was not provided and thus the genitalia remain unknown for this distinctive species. Nonetheless, genitalic variation is relatively minor across species of <italic><tp:taxon-name>Alocanthedon</tp:taxon-name></italic> and the structure of the head, tarsi, metasomal terga and sterna, and integumental sculpturing will sufficiently serve to identify future material of this species.</p>
<p><italic><tp:taxon-name>Chalicodoma apoicola</tp:taxon-name></italic>, herein described from the male alone, occurs in the same region as <italic><tp:taxon-name>Callomegachile (Callomegachile) davaonensis</tp:taxon-name></italic> (<xref ref-type="bibr" rid="B4">Cockerell 1918</xref>), described from the female sex and from a nearby area. Since the most distinguishing features of the subgenus are in the male it is possible that the latter species belongs to <italic><tp:taxon-name>Alocanthedon</tp:taxon-name></italic> and may be closely allied to <italic><tp:taxon-name>Chalicodoma apoicola</tp:taxon-name></italic>. It is tempting to speculate that <italic><tp:taxon-name>Chalicodoma apoicola</tp:taxon-name></italic> represents the<!--PageBreak--> unknown male for <italic><tp:taxon-name>Chalicodoma davaonensis</tp:taxon-name></italic>. From the original description, however, this seems unlikely given the significantly larger size of <italic><tp:taxon-name>Capoicola</tp:taxon-name></italic> (<italic>ca</italic>. 15 mm in <italic><tp:taxon-name>Chalicodoma davaonensis</tp:taxon-name></italic>, total length smaller than the forewing length in <italic><tp:taxon-name>Chalicodoma apoicola</tp:taxon-name></italic>) and the reddish translucent wings and ferruginous pterostigma and veins of <italic><tp:taxon-name>Chalicodoma davaonensis</tp:taxon-name></italic> (in this regard more similar to some of the Malaysian, Thai, and Burmese species of <italic><tp:taxon-name>Alocanthedon</tp:taxon-name></italic>). As already noted, significant collecting efforts for bees in Mindanao are needed so as to more accurately characterize these species and to more fully understand the fauna.</p>
<fig id="F3" position="float" orientation="portrait">
<label>Figures 10–14.</label>
<caption><p>Photomicrographs of male of <italic><tp:taxon-name>Chalicodoma</tp:taxon-name></italic> (<italic><tp:taxon-name>Alocanthedon</tp:taxon-name></italic>) <italic>apoicola</italic> Engel, sp. n. <bold>10</bold> Dorsal habitus <bold>11</bold> Detail of forewing medial cell <bold>12</bold> Facial aspect <bold>13</bold> Dorsal aspect of mesoscutum and mesoscutellum <bold>14</bold> Protarsus, pro-pretarsus, and apex of protibia.</p></caption>
<graphic xlink:href="ZooKeys-101-051-g003.jpg" position="float" orientation="portrait" xlink:type="simple"/>
</fig>
<fig id="F4" position="float" orientation="portrait">
<label>Figures 15–16.</label>
<caption><p>Photomicrographs of male head of <italic><tp:taxon-name>Chalicodoma</tp:taxon-name></italic> (<italic><tp:taxon-name>Alocanthedon</tp:taxon-name></italic>) <italic>apoicola</italic> Engel, sp. n. depicting juxtamandibular flange/lamella and postgenal depression <bold>15</bold> Right lateral aspect of apical portion of head <bold>16</bold> Ventral aspect of right side of head.</p></caption>
<graphic xlink:href="ZooKeys-101-051-g004.jpg" position="float" orientation="portrait" xlink:type="simple"/>
</fig>
</tp:treatment-sec>
</tp:taxon-treatment>
<tp:taxon-treatment>
<tp:nomenclature>
<tp:taxon-name>
<tp:taxon-name-part taxon-name-part-type="genus">Chalicodoma</tp:taxon-name-part>
<tp:taxon-name-part taxon-name-part-type="subgenus"> (Alocanthedon) </tp:taxon-name-part>
<tp:taxon-name-part taxon-name-part-type="species">memecylonae</tp:taxon-name-part>
<object-id xlink:type="simple">urn:lsid:zoobank.org:act:066C973E-81A4-496A-BA57-F7701C49DC33</object-id>
<object-id xlink:type="simple">http://species-id.net/wiki/Chalicodoma_(Alocanthedon)_memecylonae</object-id>
</tp:taxon-name>
<tp:taxon-authority>Engel</tp:taxon-authority>
<tp:taxon-status xlink:type="simple">sp. n.</tp:taxon-status>
<xref ref-type="fig" rid="F5">Figs 17</xref>
<xref ref-type="fig" rid="F6">25</xref>
<xref ref-type="fig" rid="F7">29–31</xref>
</tp:nomenclature>
<tp:treatment-sec sec-type="Holotype"><title>Holotype.</title>
<p>Malaysia (Peninsular): ♂, Malaya, Penang, Batu Feringgi, 17 November 1963, H.T. Pagden (NHML).</p>
</tp:treatment-sec>
<tp:treatment-sec sec-type="Paratypes"><title>Paratypes.</title>
<p>Malaysia (Peninsular): 2♂♂, Malaya, Pangkor Island, 5.i.1958 [5 January 1958], H.T. Pagden (NHML); 1♂, Malaya, Penang, Botanical Gardens at flowers <italic><tp:taxon-name>Duranta</tp:taxon-name></italic> [an introduced ornamental of <tp:taxon-name>Verbenaceae</tp:taxon-name>, native to the Americas, and accordingly not considered a host plant record herein], 11.xi.1958 [11 November 1958], H.T. Pagden (NHML); 1♂, Malaya, Penang, Batu Feringgi, 17 November 1963, H.T. Pagden (NHML); 1♀, Malaya, Penang, Mt. Erskine Road, at <italic><tp:taxon-name>Memecylon</tp:taxon-name></italic>, 18 July 1955, H.T. Pagden (NHML); 1♀, Malaya, Penang, Mt. Erskine Road, 3 June <!--PageBreak-->1955, H.T. Pagden (NHML); 1♀, Malaya, Penang, Mt. Erskine Road, 9 August 1955, H.T. Pagden (SEMC); 1♀, Malaya, Kuala Lumpur, 10.9.1933 [10 September 1933], H.M. Pendlebury, Ex F.M.S. [Federated Malay States] Museum (NHML); 1♀, Malay Penin. [Peninsula], West Coast, Langkawi Is. [Island], 19 April 1928, H.M. Pendlebury, Ex F.M.S. [Federated Malay States] Museum (NHML); 1♀, Selangor, Serdang, 10.xii.1928 [10 December 1928], H.T. Pagden (NHML); 1♀, Serdang, in Memecy. [<italic><tp:taxon-name>Memecylon</tp:taxon-name></italic>], 10.xii.1928 [10 December 1928], H.T. Pagden (NHML).</p>
</tp:treatment-sec>
<tp:treatment-sec sec-type="Diagnosis"><title>Diagnosisn.</title>
<p>Both sexes of this species have yellow forewings with grayish hyaline apex. The male can be easily recognized by the clypeus with the disc not densely covered by setae (<xref ref-type="fig" rid="F5">Fig. 18</xref>) and the absence of a patch of dense setae in the medial cell of the forewing (<xref ref-type="fig" rid="F5">Fig. 20</xref>). The female is most similar to <italic><tp:taxon-name>Chalicodoma atratiforme</tp:taxon-name></italic> from which it can be separated by the mesoscutum with more pronounced transverse wrinkling on disc, posteriorly with well-defined coarse, irregular punctures separated by a less than a puncture width, not loosely arranged in transverse series like those in wrinkles (<xref ref-type="fig" rid="F6">Fig. 25</xref>). Also, the hypostomal area is more coarsely punctate than in <italic><tp:taxon-name>Chalicodoma atratiforme</tp:taxon-name></italic>.</p>
</tp:treatment-sec>
<tp:treatment-sec sec-type="Description"><title>Description.</title>
<p>As described for <italic><tp:taxon-name>Chalicodoma odontophorum</tp:taxon-name></italic> (<italic>vide supra</italic>) except as follows: Total body length 18.0 mm; forewing length 12.7 mm. Head broader than long (width 5.0 mm, length 3.7 mm); intertorular distance 1.6 times torulorbital distance; interocellar distance 1.9 times median ocellar diameter, 1.2 times ocellocular distance; ocelloccipital distance 3.4 times median ocellar diameter; compound eye about twice as long as wide, slightly broader than gena in profile. Procoxal spine shorter and broader than that of <italic><tp:taxon-name>Chalicodoma odontophorum</tp:taxon-name></italic>, with weak depression between spine and oblique lamella of procoxa, lamella short, posterobasally setose on spine, anterior surface not setose; protibia with strong, outer, posterior carina running along apical three-quarters of length, apically produced into small posteriorly-directed spine, not carinate along transverse apex of protibia, anterior border ridged but not carinate, apical anterior surface not depressed; protarsus modified as in <xref ref-type="fig" rid="F5">figure 21</xref>; meso- and metafemora somewhat swollen; mesotibial spur relatively straight, with bluntly rounded apex. Postgradular depressions deeper and broader than in female; terga II–IV with apical transverse ridge (caudad postgradular depression), ridges somewhat sinuate laterally, weak medially on terga II–III, entirely weak on tergum IV; preapical carina of sixth tergum produced, weakly and narrowly concave medially (<xref ref-type="fig" rid="F5">Fig. 22</xref>). Genitalia as in figures <xref ref-type="fig" rid="F7">29–31</xref>.</p>
<p>Integument black throughout except tegula, legs, and metasomal sterna largely dark reddish brown (nearly black in many areas), and expansions of protarsi dark brown. Wings orange-yellow except apical margin of forewing and apical and posterior margins of hind wing grayish hyaline (<xref ref-type="fig" rid="F5">Fig. 17</xref>); venation ferruginous to orange-yellow.</p>
<p>Mandible with outer surface shiny, irregularly punctate; vertex with coarse, shallow punctures separated by less than a puncture width, integument between finely imbricate, punctures becoming more shallow toward preoccipital carina; upper gena with irregular punctures separated by finely imbricate integument, remainder of gena and posterior postgena with more regular punctures separated by a puncture width or frequently less, integument otherwise finely imbricate. Mesoscutum anteriorly and medially transversely wrinkled with irregular punctures, such integument blending<!--PageBreak--> laterally outside of parapsidal lines and posteriorly to coarsely and contiguously punctate, integument between (where evident) finely imbricate; axillae and mesoscutellum coarsely and contiguously punctate throughout. Dorsal-facing surface of first metaso<!--PageBreak-->mal tergum imbricate with small punctures nearly contiguous; remaining terga sculptured as on dorsal-facing surface of first metasomal tergum.</p>
<p>Pubescence generally dark fuscous to black except as follows: clypeus apically with dense fringe of long, apically-directed reddish setae (<xref ref-type="fig" rid="F5">Figs. 18, 19</xref>); supraclypeal area with long, numerous, tawny setae, but not entirely obscuring integument; white and black setae arranged on protarsus and apex of protibia as in <xref ref-type="fig" rid="F5">figure 21</xref>; wing setae generally yellow or tawny yellow, without dense cluster of setae forming spot in medial cell (<xref ref-type="fig" rid="F5">Figs. 17, 20</xref>).</p>
<p><italic>Female</italic>: As for the male except in usual sexual differences and as follows: Total body length 20 mm; forewing length 13.2 mm. Head broader than long (width 5.2 mm, length 3.7 mm); intertorular distance 1.2 torulorbital distance; interocellar distance 2.4 times median ocellar diameter, 1.3 times ocellocular distance; ocelloccipital distance 4.4 times median ocellar diameter; compound eye about twice as long as wide, slightly narrower than gena in profile. Mandible with four teeth; body of mandible short, basal section about as long as or slightly shorter than apical, dentate margin. Labrum rectangular, with apical row of stiff, erect, long setae running along apical margin; apical margin relatively straight. Clypeus without pronounced, medioapical tubercle, margin relatively straight. First flagellomere short, about two-thirds length of second flagellomere; remaining flagellomeres all about twice as long as wide. Procoxae, tibiae, tarsi, and spurs unmodified; pretarsal claws long, curved, simple. Postgradular depressions weaker than in male; terga without transverse ridges, with slight lateral swellings on terga II–IV.</p>
<p>Mandible with outer surface dull, irregularly punctate and microreticulate; clypeus imbricate with coarse, shallow punctures separated by less than a puncture width except along apical border. Mesoscutum anteriorly and medially transversely wrinkled, more weakly so than in male, with irregular punctures, such integument blending laterally and posteriorly to coarsely punctate, punctures separated by less than a puncture width, integument between punctures finely imbricate; axillae and mesoscutellum strongly coarsely punctate, punctures separated by less than a puncture width throughout.</p>
<p>Usual sex differences in setation; pubescence dark fuscous to black except microtrichia on inner surface of mandible dark golden and small dirty white patch on lateral surface of propodeum near metacoxa; clypeus and supraclypeal area not obscured by dense pubescence.</p>
</tp:treatment-sec>
<tp:treatment-sec sec-type="Etymology"><title>Etymology.</title>
<p>The specific epithet is based on the plant generic name <italic><tp:taxon-name>Memecylon</tp:taxon-name></italic> (neuter), and at which the type series was captured.</p>
</tp:treatment-sec>
<tp:treatment-sec sec-type="Floral records"><title>Floral records.</title>
<p>The paratype females were captured at flowers of “<italic><tp:taxon-name>Memecylon umbellatum</tp:taxon-name></italic>” Wall. (Myrtales: <tp:taxon-name>Memecylaceae</tp:taxon-name>), a <italic>nomen nudum</italic> for what is today recognized by the accepted name <italic><tp:taxon-name>Memecylon lilacinum</tp:taxon-name></italic> Zoll. & Moritz, and not to be confused with <italic><tp:taxon-name>Memecylon umbellatum</tp:taxon-name></italic> Burm.f. in Peninsular India and Sri Lanka.<!--PageBreak--><!--PageBreak--></p>
<fig id="F5" position="float" orientation="portrait">
<label>Figures 17–22.</label>
<caption><p>Photomicrographs of male of <italic><tp:taxon-name>Chalicodoma</tp:taxon-name></italic> (<italic><tp:taxon-name>Alocanthedon</tp:taxon-name></italic>) <italic>memecylonae</italic> Engel, sp. n. <bold>17</bold> Dorsal habitus <bold>18</bold> Facial aspect <bold>19</bold> Lateral aspect of head, arrow indicates postgenal flange, note also broad, medial inferior protuberance of male mandible <bold>20</bold> Detail of forewing medial cell <bold>21</bold> Protarsus, pro-pretarsus, and apex of protibia <bold>22</bold> Dorsal aspect of metasomal terga V and VI.</p></caption>
<graphic xlink:href="ZooKeys-101-051-g005.jpg" position="float" orientation="portrait" xlink:type="simple"/>
</fig>
<fig id="F6" position="float" orientation="portrait">
<label>Figures 23–25.</label>
<caption><p>Photomicrographs of female of <italic><tp:taxon-name>Chalicodoma</tp:taxon-name></italic> (<italic><tp:taxon-name>Alocanthedon</tp:taxon-name></italic>) <italic>memecylonae</italic> Engel, sp. n. <bold>23</bold> Dorsal habitus <bold>24</bold> Facial aspect <bold>25</bold> Dorsal aspect of mesoscutum and mesoscutellum.</p></caption>
<graphic xlink:href="ZooKeys-101-051-g006.jpg" position="float" orientation="portrait" xlink:type="simple"/>
</fig>
<fig id="F7" position="float" orientation="portrait">
<label>Figures 26–34.</label>
<caption><p>Photomicrographs of representative male genitalia for species of <italic><tp:taxon-name>Alocanthedon</tp:taxon-name></italic> Engel and Gonzalez, subgen. n. in ventral, lateral and dorsal aspects <bold>26–28</bold> <italic><tp:taxon-name>Chalicodoma</tp:taxon-name></italic> (<italic><tp:taxon-name>Alocanthedon</tp:taxon-name></italic>) <italic>odontophorum</italic> Engel, sp. n. <bold>29–31</bold> <italic>C</italic>. (<italic>A</italic>.) <italic>memecylonae</italic> Engel, sp. n. <bold>32–34</bold> <italic>C</italic>. (<italic>A</italic>.) <italic>atratiforme</italic> (Meade-Waldo).</p></caption>
<graphic xlink:href="ZooKeys-101-051-g007.jpg" position="float" orientation="portrait" xlink:type="simple"/>
</fig>
</tp:treatment-sec>
</tp:taxon-treatment>
<tp:taxon-treatment>
<tp:nomenclature>
<tp:taxon-name>
<tp:taxon-name-part taxon-name-part-type="genus">Chalicodoma</tp:taxon-name-part>
<tp:taxon-name-part taxon-name-part-type="subgenus"> (Alocanthedon) </tp:taxon-name-part>
<tp:taxon-name-part taxon-name-part-type="species">aterrimum</tp:taxon-name-part>
<object-id xlink:type="simple">http://species-id.net/wiki/Chalicodoma_(Alocanthedon)_aterrimum</object-id>
</tp:taxon-name>
<tp:taxon-authority>(Smith)</tp:taxon-authority>
<xref ref-type="fig" rid="F8">Figs 35</xref>
<xref ref-type="fig" rid="F9">43</xref>
<tp:nomenclature-citation-list>
<tp:nomenclature-citation><tp:taxon-name>Megachile aterrima</tp:taxon-name>
<comment><xref ref-type="bibr" rid="B17">Smith 1862</xref>: 60.
</comment>
</tp:nomenclature-citation>
<tp:nomenclature-citation><tp:taxon-name>Chalicodoma aterrimum</tp:taxon-name><comment>(Smith); <xref ref-type="bibr" rid="B2">Baker 1993</xref>: 223.
</comment>
</tp:nomenclature-citation>
</tp:nomenclature-citation-list>
</tp:nomenclature>
<tp:treatment-sec sec-type="Additional material"><title>Additional material.</title>
<p>Indonesia (Sulawesi): 1♂, 1♀, Central Sulawesi, Palolo nr. Palu, vii.1995 [July 1995] (SEMC); 1♂, Central Sulawesi, Sintuwu, SE of Palu, 1.xii.2000 [1 December 2000], I. Steffan-Dewenter (SEMC).</p>
</tp:treatment-sec>
<tp:treatment-sec sec-type="Diagnosis"><title>Diagnosisn.</title>
<p>Both sexes of this species are easily recognized by the largely hyaline forewings with fuscous apex and dark brown to black venation (<xref ref-type="fig" rid="F8">Figs. 35</xref>, <xref ref-type="fig" rid="F9">41</xref>). The male resembles that of <italic><tp:taxon-name>Chalicodoma apoicola</tp:taxon-name></italic> in the presence of a dense cluster of setae forming a spot in the medial cell of the forewing (<xref ref-type="fig" rid="F8">Fig. 39</xref>), the outer surface of the protibia distinctly depressed distally (as seen in profile view), and the distinct median emargination of the preapical carina of tergum VI (<xref ref-type="fig" rid="F8">Fig. 40</xref>). It can be separated by the smaller body size (17–19 mm vs. 24.6 mm), the disc of clypeus largely exposed, not covered by dense, appressed setae (<xref ref-type="fig" rid="F8">Fig. 36</xref>), and the shape of protarsi and setation (compare <xref ref-type="fig" rid="F3">Figs. 14</xref> and <xref ref-type="fig" rid="F8">38</xref>). As in <italic><tp:taxon-name>Chalicodoma memecylonae</tp:taxon-name></italic> and <italic><tp:taxon-name>Chalicodoma atratiforme</tp:taxon-name></italic>, the clypeal margin of the female is relatively straight, without a medioapical tubercle. In addition to the forewing color (yellow with grayish hyaline apex in those species), the female of <italic><tp:taxon-name>Chalicodoma aterrimum</tp:taxon-name></italic> differs from those species by the sparser and larger punctures on the basal two terga contrasting with the smaller, denser punctures of the remaining segments. In <italic><tp:taxon-name>Chalicodoma memecylonae</tp:taxon-name></italic> and <italic><tp:taxon-name>Chalicodoma atratiforme</tp:taxon-name></italic> the punctures are small and tightly packed on all terga.</p>
</tp:treatment-sec>
</tp:taxon-treatment>
<tp:taxon-treatment>
<tp:nomenclature>
<tp:taxon-name>
<tp:taxon-name-part taxon-name-part-type="genus">Chalicodoma</tp:taxon-name-part>
<tp:taxon-name-part taxon-name-part-type="subgenus"> (Alocanthedon) </tp:taxon-name-part>
<tp:taxon-name-part taxon-name-part-type="species">atratiforme</tp:taxon-name-part>
<object-id xlink:type="simple">http://species-id.net/wiki/Chalicodoma_(Alocanthedon)_atratiforme</object-id>
</tp:taxon-name>
<tp:taxon-authority>(Meade-Waldo)</tp:taxon-authority>
<tp:taxon-status xlink:type="simple">comb. n.</tp:taxon-status>
<xref ref-type="fig" rid="F7">Figs 32–34</xref>
<xref ref-type="fig" rid="F10">44–46</xref>
<tp:nomenclature-citation-list>
<tp:nomenclature-citation><tp:taxon-name>Megachile atratiformis</tp:taxon-name>
<comment><xref ref-type="bibr" rid="B11">Meade-Waldo 1914</xref>: 456.
</comment>
</tp:nomenclature-citation>
</tp:nomenclature-citation-list>
</tp:nomenclature>
<tp:treatment-sec sec-type="Additional material"><title>Additional material.</title>
<p>Malaysia (Peninsular): 3♀♀, Pahang, Batu Balai Estate, 18 March 1927, E. Seimund, Ex F.M.S. [Federated Malay States] Museum (NHML); 1♂, 1♀, Negri Sembilan, Gunong Angsi, 2000–2790’ [feet], April 1918, Ex F.M.S. [Federated Malay States] Museum (NHML).</p>
<p>Thailand: 1♀, Uthai Thani Province [western Thailand], Huay Kha Khaeng Wildlife Sanctuary, 15º36’ N, 99º20’ E, 1.xi.1995 [1 November 1995], J. Gazhoul, captured at <italic><tp:taxon-name>Dipterocarpus obtusifolius</tp:taxon-name></italic> Teijsman & Miquel (<tp:taxon-name>Dipterocarpaceae</tp:taxon-name>), D.B. & M.W. Baker Collection (SEMC); 1♀, 150 n.w. Bangkok, Huay Kha Khaeng, 5.xi.1995 [5 November 1995], J. Ghazoul (NHML).</p>
</tp:treatment-sec>
<tp:treatment-sec sec-type="Diagnosis"><title>Diagnosisn.</title>
<p>The female of this species can be recognized by the clypeus lacking a medioapical tubercle, short mandibles (<xref ref-type="fig" rid="F10">Fig. 45</xref>), labrum with an apical fringe of erect setae running along relatively straight apical margin, and the yellow wings (<xref ref-type="fig" rid="F10">Fig. 44</xref>). The male can be recognized by the dense setal patch in forewing medial cell, tergum <!--PageBreak--><!--PageBreak-->VI with shallow medioapical concavity on preapical carina, protibia with outer, apical surface not depressed, and genitalia as in <xref ref-type="fig" rid="F7">figures 32–34</xref>.</p>
</tp:treatment-sec>
<tp:treatment-sec sec-type="Remarks"><title>Remarks.</title>
<p>It should be noted that Meade-Waldo’s (1914) material from “Middle Tenasserim” is not conspecific with his type (B.M. Type Hym.17a2037) for <italic><tp:taxon-name>Chalicodoma atratiforme</tp:taxon-name></italic> and are actually specimens of <italic><tp:taxon-name>Chalicodoma odontophorum</tp:taxon-name></italic>. Additionally, despite <xref ref-type="bibr" rid="B5">Cockerell’s (1927)</xref> assignment of the subspecies <italic><tp:taxon-name>Chalicodoma atratiforme sininsulae</tp:taxon-name></italic> (Cockerell) (Type USNM 70455; ♀; Turtle Island [Koh Tao], Gulf of Siam; <italic>visum</italic>) to Meade-Waldo’s species, his female has nothing whatever to do with <italic><tp:taxon-name>Chalicodoma atratiforme</tp:taxon-name></italic> and is instead a synonym of <italic><tp:taxon-name>Callomegachile (Callomegachile) fulvipenne</tp:taxon-name></italic> (<xref ref-type="bibr" rid="B18">Smith 1879</xref>) (<bold>syn. n.</bold>).</p>
<fig id="F8" position="float" orientation="portrait">
<label>Figures 35–40.</label>
<caption><p>Photomicrographs of male of <italic><tp:taxon-name>Chalicodoma</tp:taxon-name></italic> (<italic><tp:taxon-name>Alocanthedon</tp:taxon-name></italic>) <italic>aterrimum</italic> (Smith) <bold>35</bold> Dorsal habitus <bold>36</bold> Facial aspect <bold>37</bold> Dorsal aspect of mesoscutum and mesoscutellum (small portion of first metasomal tergum also visible) <bold>38</bold> Protarsus, pro-pretarsus, and apex of protibia <bold>39</bold> Detail of forewing medial cell <bold>40</bold> Dorsal aspect of metasomal terga V, VI, and apical portion of IV.</p></caption>
<graphic xlink:href="ZooKeys-101-051-g008.jpg" position="float" orientation="portrait" xlink:type="simple"/>
</fig>
<fig id="F9" position="float" orientation="portrait">
<label>Figures 41–43.</label>
<caption><p>Photomicrographs of female of <italic><tp:taxon-name>Chalicodoma</tp:taxon-name></italic> (<italic><tp:taxon-name>Alocanthedon</tp:taxon-name></italic>) <italic>aterrimum</italic> (Smith) <bold>41</bold> Dorsal habitus <bold>42</bold> Facial aspect <bold>43</bold> Dorsal aspect of mesoscutum and mesoscutellum.</p></caption>
<graphic xlink:href="ZooKeys-101-051-g009.jpg" position="float" orientation="portrait" xlink:type="simple"/>
</fig>
</tp:treatment-sec>
<tp:treatment-sec>
<title><bold>Key to Species of</bold> Alocanthedon</title>
<p><bold>Note:</bold> The female of <italic><tp:taxon-name>Chalicodoma apoicola</tp:taxon-name></italic> is unknown.</p>
<table-wrap content-type="key" position="anchor" orientation="portrait">
<table>
<tbody>
<tr>
<td rowspan="1" colspan="1">1</td>
<td rowspan="1" colspan="1">Forewing either largely hyaline with fuscous apex (<xref ref-type="fig" rid="F8">Figs. 35</xref>, <xref ref-type="fig" rid="F9">41</xref>) or entirely dark fuscous (<xref ref-type="fig" rid="F3">Fig. 10</xref>), with dark brown to black venation; preapical carina of male tergum VI with deep medioapical concavity (<xref ref-type="fig" rid="F3">Figs. 10</xref>, <xref ref-type="fig" rid="F8">40</xref>); outer, apical surface of male protibia distinctly depressed</td>
<td rowspan="1" colspan="1">2</td>
</tr>
<tr>
<td rowspan="1" colspan="1">–</td>
<td rowspan="1" colspan="1">Forewing yellow, like parchment, with smoky fuscous or grayish hyaline apex (<xref ref-type="fig" rid="F1">Figs. 1</xref>, <xref ref-type="fig" rid="F2">7</xref>, <xref ref-type="fig" rid="F5">17</xref>, <xref ref-type="fig" rid="F6">23</xref>, <xref ref-type="fig" rid="F10">44</xref>), with ferruginous to orange-yellow venation; preapical carina of male tergum VI with shallow medioapical concavity (<xref ref-type="fig" rid="F1">Figs. 6</xref>, <xref ref-type="fig" rid="F5">22</xref>); outer, apical surface of male protibia faintly or not depressed</td>
<td rowspan="1" colspan="1">3</td>
</tr>
<tr>
<td rowspan="1" colspan="1">2(1)</td>
<td rowspan="1" colspan="1">Male clypeal disc largely exposed, not obscured by dense, appressed setae (<xref ref-type="fig" rid="F8">Fig. 36</xref>); procoxal spines broad, oblique procoxal lamella long, distance from lamella to outer basal corner of procoxa shorter than lamella length; anterior border of outer surface of protibia with thin fringe of short, erect, black setae; protarsi as in <xref ref-type="fig" rid="F8">figure 38</xref>; terga with faint to absent transverse ridges on non-depressed, postgradular discs; apical margin of second metasomal sternum straight, not produced medially; size moderate (17–22 mm) [Sulawesi]</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Chalicodoma aterrimum</tp:taxon-name></italic> (Smith)</td>
</tr>
<tr>
<td rowspan="1" colspan="1">–</td>
<td rowspan="1" colspan="1">Male clypeal surface largely obscured by dense, long, appressed, reddish setae (<xref ref-type="fig" rid="F3">Fig. 12</xref>); procoxal spines more slender, elongate, oblique procoxal lamella short, distance from lamella to outer basal corner of procoxa slightly longer than lamella length; anterior border of outer surface of protibia with dense fringe of long, erect, slightly wavy, black setae; protarsi as in <xref ref-type="fig" rid="F3">figure 14</xref>; terga with strong transverse carinae or ridges on non-depressed, postgradular discs; apical margin of second metasomal sternum with broad, short, medioapical extension; size very large (nearly 25 mm) [Mindanao, Philippines]</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Chalicodoma apoicola</tp:taxon-name></italic> Engel sp. n.</td>
</tr>
<tr>
<td rowspan="1" colspan="1">3(1)</td>
<td rowspan="1" colspan="1">Female clypeus with pronounced, erect, medioapical tubercle (<xref ref-type="fig" rid="F2">Fig. 8</xref>); female mandibles elongate (<xref ref-type="fig" rid="F2">Fig. 8</xref>); apical fringe of erect setae on labrum separated from labral apical margin by at least one median ocellar diameter or slightly more, apical margin of labrum medially convex; male clypeus densely covered by long, appressed, apically-directed setae, obscuring integument, face with dense setae outside of antennal toruli tawny or white (<xref ref-type="fig" rid="F1">Fig. 2</xref>); male protarsus as in <xref ref-type="fig" rid="F1">figure 5</xref> [Thailand, Myanmar]</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Chalicodoma odontophorum</tp:taxon-name></italic> Engel sp. n.</td>
</tr>
<tr>
<td rowspan="1" colspan="1">–</td>
<td rowspan="1" colspan="1">Female clypeus without medioapical tubercle; female mandibles short (<xref ref-type="fig" rid="F6">Figs. 25</xref>, <xref ref-type="fig" rid="F10">45</xref>); apical fringe of erect setae on female labrum running along labral apical margin (separated by less than a median ocellar diameter), apical margin relatively straight; male clypeal disc largely exposed, setae on face outside of antennal toruli largely black or tawny to white only near clypeus (<xref ref-type="fig" rid="F5">Fig. 18</xref>); male protarsus not as in <xref ref-type="fig" rid="F1">figure 5</xref></td>
<td rowspan="1" colspan="1">4</td>
</tr>
<tr>
<td rowspan="1" colspan="1">4(3)</td>
<td rowspan="1" colspan="1">Female mesoscutum with pronounced transverse wrinkling on disc, posteriorly with well-defined coarse punctures separated by a less than a puncture width, punctures irregular, not loosely arranged in transverse series like those in wrinkles (<xref ref-type="fig" rid="F6">Fig. 25</xref>); male forewing without patch of dense setae in forewing medial cell (<xref ref-type="fig" rid="F5">Figs. 17, 20</xref>); male genitalia as in <xref ref-type="fig" rid="F7">figures 29–31</xref> [Peninsular Malaysia]</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Chalicodoma memecylonae</tp:taxon-name></italic> Engel sp. n.</td>
</tr>
<tr>
<td rowspan="1" colspan="1">–</td>
<td rowspan="1" colspan="1">Female mesoscutum with central wrinkling of integument less pronounced, posteriorly with ill-defined, somewhat transverse punctures separated by a puncture width or more and loosely in transverse rows like weak wrinkles on disc (<xref ref-type="fig" rid="F10">Fig. 46</xref>); male forewing with dense patch of black setae in forewing medial cell; male genitalia as in <xref ref-type="fig" rid="F7">figures 32–34</xref> [Peninsular Malaysia, Thailand]</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Chalicodoma atratiforme</tp:taxon-name></italic> (Meade-Waldo)</td>
</tr>
</tbody>
</table>
</table-wrap>
</tp:treatment-sec>
</tp:taxon-treatment>
</tp:taxon-treatment>
</sec>
<sec sec-type="Cladistics"><title>Cladistics</title>
<p>A total of 157 most parsimonious trees (Length = 2275, Consistency Index = 12, Retention Index = 53) were obtained when including <italic><tp:taxon-name>Chalicodoma (A.) memecylonae</tp:taxon-name></italic> in the analysis of the data set of <xref ref-type="bibr" rid="B9">Gonzalez (2008)</xref>; 58 nodes collapsed in the consensus tree. <italic><tp:taxon-name>Chalicodoma memecylonae</tp:taxon-name></italic> was included in a clade, sister to <italic><tp:taxon-name>Cuspidella</tp:taxon-name></italic>, containing four of the six species of <italic><tp:taxon-name>Callomegachile</tp:taxon-name></italic> used in the analysis: (<italic><tp:taxon-name>Chalicodoma mystaceana</tp:taxon-name></italic> + <italic><tp:taxon-name>Chalicodoma biseta</tp:taxon-name></italic>) + [<italic><tp:taxon-name>Chalicodoma memecylonae</tp:taxon-name></italic> (<italic><tp:taxon-name>Chalicodoma sculpturalis</tp:taxon-name></italic> + <italic><tp:taxon-name>Chalicodoma clotho</tp:taxon-name></italic>)]. The same topology was obtained when deactivating five characters related to parasitism, or excluding one of the outgroup species (i.e., <italic><tp:taxon-name>Dioxys</tp:taxon-name></italic> Lepeletier de Saint Fargeau and Audinet-Serville) or species with missing characters as an attempt to explore other hypotheses of relationships. However, no synapomorphies or high bootstrap values (> 50%) support the placement of <italic><tp:taxon-name>Chalicodoma memecylonae</tp:taxon-name></italic> within <italic><tp:taxon-name>Callomegachile</tp:taxon-name></italic>.</p>
<p><italic><tp:taxon-name>Alocanthedon</tp:taxon-name></italic> is supported by significant autapomorphies as outlined in the subgeneric description above (<italic>vide supra</italic>). Most features shared between <italic><tp:taxon-name>Alocanthedon</tp:taxon-name></italic> and other <italic><tp:taxon-name>Chalicodoma</tp:taxon-name></italic> subgenera and groups (many of which may be elevated suitably to subgeneric status) are plesiomorphic and are summarized here. The following plesiomorphies are shared between <italic><tp:taxon-name>Chalicodoma memecylonae</tp:taxon-name></italic> and <italic><tp:taxon-name>Callomegachile</tp:taxon-name></italic> in the analyses: first flagellomere shorter than second flagellomere; outer surface of mandible dull, micro<!--PageBreak-->reticulate to finely punctuate; mesepisternum coarsely punctuate, forming strong rows with distinct shining ridges among them; clypeus of male with sparse setae on basal half, distal half densely covered by setae (integument not visible) [completely covered in two other <italic><tp:taxon-name>Alocanthedon</tp:taxon-name></italic> species as in <xref ref-type="fig" rid="F1">Fig. 2</xref>]; T6 of male without lateral spine on apical margin; male genitalia with rounded or pointed volsella, and penis valve distinctly curved or arched inward. Likewise, <italic><tp:taxon-name>Chalicodoma memecylonae</tp:taxon-name></italic> shares the following plesiomorphies with <italic><tp:taxon-name>Chalicodoma sculpturalis</tp:taxon-name></italic> and <italic><tp:taxon-name>Chalicodoma clotho</tp:taxon-name></italic>, both species of the ‘Eumegachilana’ group: compound eyes strongly diverging ventrally; clypeoantennal distance short, equal to or shorter than vertical diameter of antennal torulus; antennal scape long, at least 1.2 times longer than torulocellar distance; clypeus short, ≥ 3x wider than long; labrum with two types of setae on disc, minute, yellowish, appressed setae, and long (≥ 1 median ocellar diameter); mandible with outer premarginal fimbria reduced or absent;<!--PageBreak--> mandible of male with broad, subtriangular, posteriorly-directed process on basal third of inferior border.</p>
<fig id="F10" position="float" orientation="portrait">
<label>Figures 44–46.</label>
<caption><p>Photomicrographs of female of <italic><tp:taxon-name>Chalicodoma</tp:taxon-name></italic> (<italic><tp:taxon-name>Alocanthedon</tp:taxon-name></italic>) <italic>atratiforme</italic> (Meade-Waldo), comb. n. <bold>44</bold> Dorsal habitus <bold>45</bold> Facial aspect <bold>46</bold> Dorsal aspect of mesoscutum and mesoscutellum.</p></caption>
<graphic xlink:href="ZooKeys-101-051-g010.jpg" position="float" orientation="portrait" xlink:type="simple"/>
</fig>
</sec>
<sec sec-type="Discussion"><title>Discussion</title>
<p>The presence of a dense patch of black setae on the forewing of males (except in one species), resembling the dense patch of setae among the submarginal cells of <italic><tp:taxon-name>Thrinchostoma</tp:taxon-name></italic> (<tp:taxon-name>Halictidae</tp:taxon-name>), is unique among <tp:taxon-name>Megachilidae</tp:taxon-name>. Likewise, the deep postgenal depression in males having a distinctly protuberant, thick lamella next to the mandible, procoxa with oblique carina or lamella medially on disc, mesobasitarsus with the inner surface concave basally, and the distinctly long, narrow gonostyli of the genitalia are apomorphic characters within <italic><tp:taxon-name>Chalicodoma</tp:taxon-name></italic>. Such characters are distinctive enough to support the separation of those species into a new subgenus. Also, <italic><tp:taxon-name>Alocanthedon</tp:taxon-name></italic> females can be reliably separated from any <italic><tp:taxon-name>Chalicodoma</tp:taxon-name></italic>, particularly those of the subgenus <italic><tp:taxon-name>Callomegachile</tp:taxon-name></italic>, by the clypeal shape (concave to V-shaped on epistomal sulcus basally and medially-projected on distal margin), and the presence of a short, stout seta on the pretarsal claw basally. Females of a few Oriental <italic><tp:taxon-name>Callomegachile</tp:taxon-name></italic> species, such as <italic><tp:taxon-name>Chalicodoma terminalis</tp:taxon-name></italic> (Smith), also have a similar modification in the clypeus but the setae on the pretarsal claws are either normal or they have a different combination of characters (e.g., presence of omaular carina, different shape of labrum, punctation, &c).</p>
<p>Based on the phylogenetic analysis, it may seems best to regard <italic><tp:taxon-name>Chalicodoma memecylonae</tp:taxon-name></italic> as a species group of <italic><tp:taxon-name>Callomegachile</tp:taxon-name></italic> rather than separating it as a new subgenus, despite that no synapomorphies or high bootstrap values (> 50%) supported such relationship in the analysis. Although emphasizing differences is sometimes useful in morphologically homogeneous groups, such as <tp:taxon-name>Augochlorini</tp:taxon-name>, <tp:taxon-name>Euglossini</tp:taxon-name>, or <tp:taxon-name>Meliponini</tp:taxon-name>, the high morphological variation present across <tp:taxon-name>Megachilini</tp:taxon-name> may lead to an excessive splitting, thus conveying little information regarding relationships. Female characters are usually less variable than male characters in <tp:taxon-name>Megachilini</tp:taxon-name>, as in the other aforementioned tribes. Given such variation in the male, some species have been separated subgenerically entirely on male secondary sexual characters (such as those found in the male <italic><tp:taxon-name>Chalicodoma memecylonae</tp:taxon-name></italic>) when females are clearly associated with an existing subgenus. For example, the female of <italic><tp:taxon-name>Megachile laeta</tp:taxon-name></italic> Smith, placed in the subgenus <italic><tp:taxon-name>Leptorachina</tp:taxon-name></italic> by <xref ref-type="bibr" rid="B15">Mitchell (1980)</xref>, is easily assigned to <italic><tp:taxon-name>Megachile</tp:taxon-name></italic> subgenus <italic><tp:taxon-name>Leptorachis</tp:taxon-name></italic> Mitchell based on the mandibular structure and distinctive pubescence of the S6; the male, however, is unlike any other <italic><tp:taxon-name>Leptorachis</tp:taxon-name></italic> in having highly modified front legs that are used to hold the female during mating. Thus, to avoid excessive splitting, it seems more convenient to emphasize the similarities rather than the differences among groups and, although male characters might provide useful phylogenetic information, there is no need to isolate a species into its own subgenus solely on the male morphology.</p>
<p>Female characters (e.g., mandible with outer surface of dull, microreticulate to finely punctuate integument, without cutting edges, mesepisternum coarsely punctuate, forming strong rows with distinct shining ridges among them, and body parallel-<!--PageBreak-->sided) also clearly associate <italic><tp:taxon-name>Chalicodoma memecylonae</tp:taxon-name></italic> with <italic><tp:taxon-name>Callomegachile</tp:taxon-name></italic> and its recognition as a separated subgenus may render <italic><tp:taxon-name>Callomegachile</tp:taxon-name></italic> paraphyletic. However, given our limited understanding on the phylogenetic relationships of <italic><tp:taxon-name>Chalicodoma</tp:taxon-name></italic> subgenera as well as of the species groups within <italic><tp:taxon-name>Callomegachile</tp:taxon-name></italic>, such a subgeneric recognition might be desirable for the time being to highlight those autopomorphic characters not found elsewhere in <tp:taxon-name>Megachilidae</tp:taxon-name>.</p>
<p>A preliminary morphological analysis of <tp:taxon-name>Megachilini</tp:taxon-name> suggested that <italic><tp:taxon-name>Callomegachile</tp:taxon-name></italic> is likely not monophyletic (<xref ref-type="bibr" rid="B9">Gonzalez 2008</xref>). Two of the six species of <italic><tp:taxon-name>Callomegachile</tp:taxon-name></italic> used in the analysis, <italic><tp:taxon-name>Chalicodoma torrida</tp:taxon-name></italic> and <italic><tp:taxon-name>Chalicodoma decemsignata</tp:taxon-name></italic> Radoszkowski, never grouped with the other species of the subgenus in that analysis or when we reanalyzed the same data set including <italic><tp:taxon-name>Chalicodoma memecylonae</tp:taxon-name></italic>. These two distinctive species, as well as <italic><tp:taxon-name>Chalicodoma biseta</tp:taxon-name></italic> and <italic><tp:taxon-name>Chalicodoma clotho</tp:taxon-name></italic> have been subgenerically separated into <italic><tp:taxon-name>Carinula</tp:taxon-name></italic> Michener et al., <italic><tp:taxon-name>Morphella</tp:taxon-name></italic><!--PageBreak--><!--PageBreak--><!--PageBreak--> Pasteels, and <italic><tp:taxon-name>Eumegachilana</tp:taxon-name></italic> Michener, respectively (<xref ref-type="bibr" rid="B14">Michener 2007</xref>). However, they have been treated as species groups because of the variation among species within each group and intergradations in some of the main characters that separate them from other <italic><tp:taxon-name>Callomegachile</tp:taxon-name></italic> (<xref ref-type="bibr" rid="B14">Michener 2007</xref>). With nearly 100 species described (<xref ref-type="bibr" rid="B1">Ascher and Pickering 2011</xref>), <italic><tp:taxon-name>Callomegachile</tp:taxon-name></italic> is the most diverse, morphologically heterogeneous, and widely distributed of all subgenera of <italic><tp:taxon-name>Chalicodoma</tp:taxon-name></italic>. The subgenus is largely tropical, occurring in Sub-Saharan Africa, southern Palaearctic, Australia, and Southeast Asia (<xref ref-type="bibr" rid="B14">Michener 2007</xref>). Species vary greatly not only in body size (~8 to 39 mm) but also in the pubescence, the shape of the mandible and labrum, punctation, presence of preoccipital and omaular carinae in both sexes, as well as in the shape of the hidden sterna and genitalia of the male. Without a doubt, given the number of species and astonishing morphological diversity, a detailed phylogenetic analysis of <italic><tp:taxon-name>Callomegachile</tp:taxon-name></italic> is needed. To document the variation in the aforementioned characters, we examined 50 <italic><tp:taxon-name>Callomegachile</tp:taxon-name></italic> species occurring across the distribution range of the subgenus. The variation found among species with unique combination of characters (<italic>n</italic> = 26) is summarized in <xref ref-type="table" rid="T2">table 2</xref>. The list is not exhaustive as several characters of the hidden sterna and male genitalia were not included. However, we hope to draw more attention to and encourage future phylogenetic studies using these characters. Some of them might prove useful in recognizing natural species groups or inferring floral hosts, such as the presence of capitate hairs on the ventral surfaces of the meso- and metasoma.</p>
<table-wrap position="float" orientation="portrait" id="T2">
<label>Table 2.</label>
<caption>
<p>Some morphological characters of certain species of <italic><tp:taxon-name>Chalicodoma</tp:taxon-name></italic> subgenus <italic><tp:taxon-name>Callomegachile</tp:taxon-name></italic> s.l. Female: <bold>1</bold> = labrum triangular; <bold>2</bold> = labrum with distinct fringe of setae basally; <bold>3</bold> = number of mandibular teeth; <bold>4</bold> = elongated mandible; <bold>5</bold> = clypeal carina; <bold>6</bold> = clypeus with slightly concave to V-shaped epistomal sulcus basally; <bold>7</bold> = vertex with a fine, shining longitudinal line from ocelli to posterior margin of vertex; <bold>8</bold> = preoccipital carina; <bold>9</bold> = omaular carina (species with asterisks have omaular carina ventrally only); <bold>10</bold> = pronotal lobe strongly carinate or lamellate; <bold>11</bold> = mesepisternum coarsely punctate, forming strong rows with distinct shining ridges among them; <bold>12</bold> = mesoscutum coarsely punctate, forming strong rows with distinct shining ridges among them; <bold>13</bold> = capitate hairs on ventral surface of mesepisternum and coxae; <bold>14</bold> = pretarsal claws with a seta conspicuously shorter and stouter than the other. Male: <bold>15</bold> = genal concavity; <bold>16</bold> = low juxtamandibular border (not strongly projecting as a lamella as in <italic><tp:taxon-name>Alocanthedon</tp:taxon-name></italic>); <bold>17</bold> = procoxal spine; <bold>18</bold> = modified protarsi; <bold>19</bold> = T6 with strong preapical carina; <bold>20</bold> = T6 with distinct depression above preapical carina; <bold>21</bold> = sterna densely covered by short appressed setae; <bold>22</bold> = volsella. <bold>Dist</bold> = Distribution: O = Oriental, E = Ethiopian; A = Australian; S = Sub-Saharan Africa. Plus (+) and dash (-) symbols indicate presence and absence of a character.</p></caption>
<table>
<tbody>
<tr>
<th rowspan="1" colspan="24"><italic>Characters</italic></th>
</tr>
<tr>
<th rowspan="1" colspan="1"><italic>Species</italic></th>
<th rowspan="1" colspan="1"><italic>1</italic></th>
<th rowspan="1" colspan="1"><italic>2</italic></th>
<th rowspan="1" colspan="1"><italic>3</italic></th>
<th rowspan="1" colspan="1"><italic>4</italic></th>
<th rowspan="1" colspan="1"><italic>5</italic></th>
<th rowspan="1" colspan="1"><italic>6</italic></th>
<th rowspan="1" colspan="1"><italic>7</italic></th>
<th rowspan="1" colspan="1"><italic>8</italic></th>
<th rowspan="1" colspan="1"><italic>9</italic></th>
<th rowspan="1" colspan="1"><italic>10</italic></th>
<th rowspan="1" colspan="1"><italic>11</italic></th>
<th rowspan="1" colspan="1"><italic>12</italic></th>
<th rowspan="1" colspan="1"><italic>13</italic></th>
<th rowspan="1" colspan="1"><italic>14</italic></th>
<th rowspan="1" colspan="1"><italic>15</italic></th>
<th rowspan="1" colspan="1"><italic>16</italic></th>
<th rowspan="1" colspan="1"><italic>17</italic></th>
<th rowspan="1" colspan="1"><italic>18</italic></th>
<th rowspan="1" colspan="1"><italic>19</italic></th>
<th rowspan="1" colspan="1"><italic>20</italic></th>
<th rowspan="1" colspan="1"><italic>21</italic></th>
<th rowspan="1" colspan="1"><italic>22</italic></th>
<th rowspan="1" colspan="1"><italic>Dist</italic></th>
</tr>
<tr>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Chalicodoma antinorii</tp:taxon-name></italic> (Gribodo, 1879)</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">6</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">E</td>
</tr>
<tr>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Chalicodoma biseta</tp:taxon-name></italic> Vachal, 1903</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">4</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">E, S</td>
</tr>
<tr>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Chalicodoma cephalotes</tp:taxon-name></italic> (Smith, 1853)</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">3</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+*</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">O</td>
</tr>
<tr>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Chalicodoma chrysorrhoea</tp:taxon-name></italic> (Gerstäcker, 1857)</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">4</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+*</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">E</td>
</tr>
<tr>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Chalicodoma devexa</tp:taxon-name></italic> (Vachal, 1903<italic>)</italic></td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">4</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">E</td>
</tr>
<tr>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Chalicodoma disjuncta</tp:taxon-name></italic> (Fabricius, 1781)</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">4</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+*</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">O</td>
</tr>
<tr>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Chalicodoma excavata</tp:taxon-name></italic> (Cockerell, 1937)</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">3</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">E</td>
</tr>
<tr>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Chalicodoma fulvipenne</tp:taxon-name></italic> (Smith, 1879)</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">5</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">O</td>
</tr>
<tr>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Chalicodoma grandiceps</tp:taxon-name></italic> (Friese, 1903)</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">3</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+*</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">E</td>
</tr>
<tr>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Chalicodoma incisa</tp:taxon-name></italic> (Smith, 1858)</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">5</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">O</td>
</tr>
<tr>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Chalicodoma lerma</tp:taxon-name></italic> (Cameron, 1908) <italic>comb. n.</italic></td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">4</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">O</td>
</tr>
<tr>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Chalicodoma mephistolica</tp:taxon-name></italic> (Gribodo, 1894)</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">4</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+*</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">E</td>
</tr>
<tr>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Chalicodoma mystaceana</tp:taxon-name></italic> Michener, 1962</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">5</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">A</td>
</tr>
<tr>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Chalicodoma perniciosa</tp:taxon-name></italic> (Friese, 1903)</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">3</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+*</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">E</td>
</tr>
<tr>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Chalicodoma rambutwan</tp:taxon-name></italic> (Cheesman, 1936)</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">4</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">O</td>
</tr>
<tr>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Chalicodoma rufipes</tp:taxon-name></italic> (Fabricius, 1781)</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">4</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+*</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">E</td>
</tr>
<tr>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Chalicodoma rufiventris</tp:taxon-name></italic> (Guérin-Méneville, 1834)</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">3</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+*</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">E</td>
</tr>
<tr>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Chalicodoma simonyi</tp:taxon-name></italic> (Friese, 1903)</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">4</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+*</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">E</td>
</tr>
<tr>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Chalicodoma terminalis</tp:taxon-name></italic> (Smith, 1858) <italic>n. comb.</italic></td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">5</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">O</td>
</tr>
<tr>
<td rowspan="1" colspan="1">‘Carinula’ group</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"/>