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<document id="65635F24257ABAB628D24DD2B973584D" ID-CLB-Dataset="273558" ID-DOI="10.5852/ejt.2023.892.2283" ID-GBIF-Dataset="f9b457af-ea60-4812-9bed-ed48423af33e" ID-ISSN="2118-9773" ID-Zenodo-Dep="8382675" ID-ZooBank="07D215E7-FB43-4640-BB5B-D1AF50269AF1" IM.bibliography_approvedBy="jonas" IM.illustrations_approvedBy="jonas" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_approvedBy="jonas" IM.taxonomicNames_approvedBy="felipe" IM.treatmentCitations_approvedBy="felipe" IM.treatments_approvedBy="jonas" checkinTime="1695805589820" checkinUser="plazi" docAuthor="Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li &amp; Orr, Michael C." docDate="2023" docId="1A438786790D04169EECFCC2FCA7FB24" docLanguage="en" docName="EJT.2023.892.1-65.pdf" docOrigin="European Journal of Taxonomy 892 (1)" docSource="https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851" docStyle="DocumentStyle:EF2B578F1D15862ADE45B0C07C620911.14:EJT.2018-.journal_article.type1" docStyleId="EF2B578F1D15862ADE45B0C07C620911" docStyleName="EJT.2018-.journal_article.type1" docStyleVersion="14" docTitle="Bombus (Alpigenobombus) rainai Williams 2022" docType="treatment" docVersion="9" lastPageNumber="36" masterDocId="E67AFFFE792F04359F0CFFE2FFE4FF9B" masterDocTitle="Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae)" masterLastPageNumber="65" masterPageNumber="1" pageNumber="35" updateTime="1699690410639" updateUser="plazi" zenodo-license-document="CC-BY-4.0" zenodo-license-figures="CC-BY-4.0" zenodo-license-treatments="UNSPECIFIED">
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<mods:title id="E2267898735EFE10564805FB4D2F8FA2">Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae)</mods:title>
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<mods:namePart id="8AC8A81306118D0C647AAE53D929C6C3">Williams, Paul H.</mods:namePart>
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<mods:namePart id="FA27929AB6B50DCFD5D59B08C5ADAB87">An, Jiandong</mods:namePart>
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<mods:affiliation id="22710B5BF6738791BB1DB3F1A569BAE0">Institute of Apicultural Research (Chinese Academy of Agricultural Sciences), 2 Yuanmingyuan West Road, Haidian, Beijing 100093, China.</mods:affiliation>
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<mods:affiliation id="451652AA23FA35B58B6A6FC3E6A9BDD3">Institute of Apicultural Research (Chinese Academy of Agricultural Sciences), 2 Yuanmingyuan West Road, Haidian, Beijing 100093, China.</mods:affiliation>
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<mods:affiliation id="1D97568DCDA42D7B02B96CA45FAE3BE5">Rajiv Gandhi University, Papum Pare, Arunachal Pradesh 791112, India. &amp; National Centre for Biological Sciences, Tata Institute of Fundamental Research, Bangalore 560065, India.</mods:affiliation>
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<mods:affiliation id="8C79F5CAF3E903AABC2DAFEC56659624">University of Vienna, Vienna, Austria.</mods:affiliation>
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<mods:affiliation id="4F03D1113535E690703431B16DB8DC6B">Institute of Zoology (Chinese Academy of Sciences), 1 Beichen West Road, Chaoyang, Beijing 100101, China. &amp; Staatliches Museum f ̧ r Naturkunde Stuttgart, Stuttgart, Germany.</mods:affiliation>
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<treatment id="1A438786790D04169EECFCC2FCA7FB24" ID-DOI="http://doi.org/10.5281/zenodo.10023245" ID-GBIF-Taxon="214226936" ID-Zenodo-Dep="10023245" LSID="urn:lsid:plazi:treatment:1A438786790D04169EECFCC2FCA7FB24" httpUri="http://treatment.plazi.org/id/1A438786790D04169EECFCC2FCA7FB24" lastPageId="35" lastPageNumber="36" pageId="34" pageNumber="35">
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4.
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<emphasis id="A09EEA82790D04179EF2FCC2FC7CFCA1" bold="true" box="[510,920,800,827]" pageId="34" pageNumber="35">
<emphasis id="A09EEA82790D04179EF2FCC2FD85FCA0" bold="true" box="[510,609,800,827]" italics="true" pageId="34" pageNumber="35">Bombus</emphasis>
(
<emphasis id="A09EEA82790D04179D7EFCC2FCDAFCA0" bold="true" box="[626,830,800,827]" italics="true" pageId="34" pageNumber="35">Alpigenobombus</emphasis>
)
<emphasis id="A09EEA82790D04179C43FCC2FC7CFCA1" bold="true" box="[847,920,800,826]" italics="true" pageId="34" pageNumber="35">rainai</emphasis>
</emphasis>
Williams, 2022
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<subSubSection id="DAF0651B790D04179DB7FCA1FC9DFCC5" box="[699,889,835,862]" pageId="34" pageNumber="35" type="description">
<paragraph id="92553690790D04179DB7FCA1FC9DFCC5" blockId="34.[480,1107,800,862]" box="[699,889,835,862]" pageId="34" pageNumber="35">
<figureCitation id="0AD12A15790D04179DB7FCA1FCA5FCC5" box="[699,833,835,862]" captionStart="Figs 19–105" captionStartId="28.[189,242,1857,1883]" captionTargetBox="[264,1318,295,2006]" captionTargetId="figure-11@27.[240,1348,265,2019]" captionTargetPageId="27" captionText="Figs 19–105 (see pages 28–29). Simplified diagrams for the colour patterns of the hair on the dorsum for particular female (f) and male (m) specimens of the species from Fig. 12. The dorsum is divided into regions, each of which shows only the predominant or most apparent colour for that region, using a simplified colour palette, with olive indicating a mixture of black and yellow hair, and grey indicating a mixture of black and white hair." figureDoi="http://doi.org/10.5281/zenodo.10015244" httpUri="https://zenodo.org/record/10015244/files/figure.png" pageId="34" pageNumber="35">Figs 49‒53</figureCitation>
,
<figureCitation id="0AD12A15790D04179C41FCA6FC9DFCC5" box="[845,889,836,862]" captionStart="Figs 106–115" captionStartId="29.[189,242,1752,1778]" captionTargetBox="[248,1333,302,1671]" captionTargetId="figure-11@29.[236,1352,265,1701]" captionTargetPageId="29" captionText="Figs 106–115. Morphology of the male genitalia for species of the subgenus Alpigenobombus Skorikov, 1914, from the dorsal aspect, anterior at the bottom of the image, posterior at the top. 106. Bombus wurflenii Radoszkowski, 1860, Turkey. 107. B mastrucatus Gerstaecker, 1869, Austria. 108. B. kashmirensis Friese, 1909, India-Kashmir. 109. B. rainai Williams, 2022, India-Kashmir. 110. B. sikkimi Friese, 1918, Nepal. 111. B. nobilis Friese, 1905, China-Sichuan. 112. B. validus Friese, 1905, China-Gansu. 113. B. genalis Friese, 1918, China-Yunnan. 114. B. breviceps Smith, 1852, ChinaSichuan. 115. B. grahami (Frison, 1933) China-Sichuan (left penis-valve recurved hook missing). Scale bars = 1 mm." figureDoi="http://doi.org/10.5281/zenodo.8382695" httpUri="https://zenodo.org/record/8382695/files/figure.png" pageId="34" pageNumber="35">109</figureCitation>
</paragraph>
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<taxonomicName id="55EA4D13790D04179FB1FC67FCB5FC04" authority="Williams, 2022 a: 132" authorityName="Williams" authorityPageNumber="132" authorityYear="2022" box="[189,849,901,927]" class="Insecta" family="Apidae" genus="Bombus" kingdom="Animalia" order="Hymenoptera" pageId="34" pageNumber="35" phylum="Arthropoda" rank="species" species="rainai" subGenus="Alpigenobombus">
<emphasis id="A09EEA82790D04179FB1FC67FEF8FC04" box="[189,284,901,927]" italics="true" pageId="34" pageNumber="35">Bombus</emphasis>
(
<emphasis id="A09EEA82790D04179E21FC67FE16FC04" box="[301,498,901,927]" italics="true" pageId="34" pageNumber="35">Alpigenobombus</emphasis>
)
<emphasis id="A09EEA82790D04179D0FFC64FDAFFC04" box="[515,587,902,927]" italics="true" pageId="34" pageNumber="35">rainai</emphasis>
Williams, 2022a: 132
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.
</paragraph>
<paragraph id="92553690790D04179FB1FC24FB30FC7B" blockId="34.[189,1236,966,992]" box="[189,1236,966,992]" pageId="34" pageNumber="35">
<treatmentCitationGroup id="B2FA11BE790D04179FB1FC24FB30FC7B" box="[189,1236,966,992]" pageId="34" pageNumber="35">
<taxonomicName id="55EA4D13790D04179FB1FC24FD7FFC7B" ID-CoL="MFLZ" authorityName="Friese" authorityYear="1909" box="[189,667,966,992]" class="Insecta" family="Apidae" genus="Bombus" kingdom="Animalia" order="Hymenoptera" pageId="34" pageNumber="35" phylum="Arthropoda" rank="species" species="kashmirensis" subGenus="Alpigenobombus">
<emphasis id="A09EEA82790D04179FB1FC24FEF8FC7B" box="[189,284,966,992]" italics="true" pageId="34" pageNumber="35">Bombus</emphasis>
(
<emphasis id="A09EEA82790D04179E21FC24FE16FC7B" box="[301,498,966,992]" italics="true" pageId="34" pageNumber="35">Alpigenobombus</emphasis>
)
<emphasis id="A09EEA82790D04179D0FFC24FD7FFC7B" box="[515,667,966,992]" italics="true" pageId="34" pageNumber="35">kashmirensis</emphasis>
</taxonomicName>
<emphasis id="A09EEA82790D04179DAEFC25FD55FC7B" box="[674,689,967,992]" italics="true" pageId="34" pageNumber="35">‒</emphasis>
<treatmentCitation id="134B1081790D04179DB4FC24FC70FC7B" author="Williams P. H." box="[696,916,966,992]" page="66" pageId="34" pageNumber="35" year="1991">
<bibRefCitation id="F67B4B61790D04179DB4FC24FC70FC7B" author="Williams P. H." box="[696,916,966,992]" pageId="34" pageNumber="35" pagination="1 - 204" refId="ref38198" refString="Williams P. H. 1991. The bumble bees of the Kashmir Himalaya (Hymenoptera: Apidae, Bombini). Bulletin of the British Museum (Natural History) (Entomology) 60: 1 - 204. Available from https: // www. biodiversitylibrary. org / page / 41055963 [accessed 13 Aug. 2023]." type="journal article" year="1991">Williams 1991: 66</bibRefCitation>
</treatmentCitation>
, (in part) misidentification.
</treatmentCitationGroup>
</paragraph>
</subSubSection>
<subSubSection id="DAF0651B790D04179FB1FBE5FDD0F9C9" pageId="34" pageNumber="35" type="discussion">
<paragraph id="92553690790D04179FB1FBE5FD9BFBBA" blockId="34.[189,639,1031,1058]" box="[189,639,1031,1058]" pageId="34" pageNumber="35">
<heading id="C91D81FC790D04179FB1FBE5FD9BFBBA" bold="true" box="[189,639,1031,1058]" fontSize="11" level="3" pageId="34" pageNumber="35" reason="3">
<emphasis id="A09EEA82790D04179FB1FBE5FD9BFBBA" bold="true" box="[189,639,1031,1058]" pageId="34" pageNumber="35">Species-taxon concept and variation</emphasis>
</heading>
</paragraph>
<paragraph id="92553690790D04179FB1FBD4FACBFB21" blockId="34.[189,1400,1078,1210]" pageId="34" pageNumber="35">
The taxon concept of the species
<taxonomicName id="55EA4D13790D04179D47FBD5FD50FBCB" authorityName="Williams" authorityYear="2022" box="[587,692,1079,1104]" class="Insecta" family="Apidae" genus="Bombus" kingdom="Animalia" order="Hymenoptera" pageId="34" pageNumber="35" phylum="Arthropoda" rank="species" species="rainai">
<emphasis id="A09EEA82790D04179D47FBD5FD50FBCB" box="[587,692,1079,1104]" italics="true" pageId="34" pageNumber="35">B. rainai</emphasis>
</taxonomicName>
here agrees with the recent interpretation (Williams 2022a) that it is separate from the taxon concept of the species
<taxonomicName id="55EA4D13790D04179C6CFBB8FBFFFBE8" authorityName="Friese" authorityYear="1909" box="[864,1051,1113,1139]" class="Insecta" family="Apidae" genus="Bombus" kingdom="Animalia" order="Hymenoptera" pageId="34" pageNumber="35" phylum="Arthropoda" rank="species" species="kashmirensis">
<emphasis id="A09EEA82790D04179C6CFBB8FBFFFBE8" box="[864,1051,1113,1139]" italics="true" pageId="34" pageNumber="35">B. kashmirensis</emphasis>
</taxonomicName>
(see the comments above on
<taxonomicName id="55EA4D13790D04179FB3FB9FFE92FB0C" authorityName="Friese" authorityYear="1909" box="[191,374,1149,1175]" class="Insecta" family="Apidae" genus="Bombus" kingdom="Animalia" order="Hymenoptera" pageId="34" pageNumber="35" phylum="Arthropoda" rank="species" species="kashmirensis">
<emphasis id="A09EEA82790D04179FB3FB9FFE92FB0C" box="[191,374,1149,1175]" italics="true" pageId="34" pageNumber="35">B. kashmirensis</emphasis>
</taxonomicName>
), based on: (1) our PTP analysis supports independent species-level coalescents in the COI gene (
<figureCitation id="0AD12A15790D04179E33FB42FE72FB21" box="[319,406,1184,1210]" captionStart="Fig" captionStartId="16.[189,232,1646,1672]" captionTargetBox="[246,1342,268,1600]" captionTargetId="figure-10@16.[246,1342,265,1603]" captionTargetPageId="16" captionText="Fig. 12. Interpretation of filtered MrBayes estimate of phylogeny as a metric tree (outgroup B. festivus Smith, 1861, not shown) for the subgenus Alpigenobombus Skorikov, 1914, from COI-like sequences from the sequences identified in black and orange in Fig. 11 together with Bayesian Poisson-tree-process (PTP) models re-applied for assessing support for species’ gene coalescents by maximum likelihood (for the 11 most likely candidate species). The scale bar is calibrated in substitutions per nucleotide site. Sequence labels and branch colours as in Fig. 10. Numbers above nodes are the Bayesian support values that all daughter haplotypes are parts of a single species. Grey spots show likely mitochondrial-to-nuclear transfers of the low-divergence numts accepted for estimating this tree. Asterisks mark sequences used as informal proxies for the type specimens of each of the taxon names in Table 2. To the right in grey are shown the interpretations of the PTP results as candidate species using the oldest available names for the species." figureDoi="http://doi.org/10.5281/zenodo.8382683" httpUri="https://zenodo.org/record/8382683/files/figure.png" pageId="34" pageNumber="35">Fig. 12</figureCitation>
); corroborated by (2) diagnostic morphological character states (see the keys).
</paragraph>
<paragraph id="92553690790D04179FB1FB03FCC8FA85" blockId="34.[189,1398,1249,1311]" pageId="34" pageNumber="35">
This taxon was discussed as divergent from
<taxonomicName id="55EA4D13790D04179DC4FB00FC65FB60" authorityName="Friese" authorityYear="1909" box="[712,897,1249,1275]" class="Insecta" family="Apidae" genus="Bombus" kingdom="Animalia" order="Hymenoptera" pageId="34" pageNumber="35" phylum="Arthropoda" rank="species" species="kashmirensis">
<emphasis id="A09EEA82790D04179DC4FB00FC65FB60" box="[712,897,1249,1275]" italics="true" pageId="34" pageNumber="35">B. kashmirensis</emphasis>
</taxonomicName>
by
<bibRefCitation id="F67B4B61790D04179CA1FB03FB24FB60" author="Williams P. H." box="[941,1216,1249,1275]" pageId="34" pageNumber="35" pagination="1 - 204" refId="ref38198" refString="Williams P. H. 1991. The bumble bees of the Kashmir Himalaya (Hymenoptera: Apidae, Bombini). Bulletin of the British Museum (Natural History) (Entomology) 60: 1 - 204. Available from https: // www. biodiversitylibrary. org / page / 41055963 [accessed 13 Aug. 2023]." type="journal article" year="1991">Williams (1991: 68‒69</bibRefCitation>
, figs 308‒310), although it was not then named as a separate species.
</paragraph>
<paragraph id="92553690790D04179FB1FAA4FD4DFA3C" blockId="34.[188,1399,1350,1447]" pageId="34" pageNumber="35">
Although the PTP and morphological results (
<figureCitation id="0AD12A15790D04179DD8FAA4FCCFFAFB" box="[724,811,1350,1376]" captionStart="Fig" captionStartId="16.[189,232,1646,1672]" captionTargetBox="[246,1342,268,1600]" captionTargetId="figure-10@16.[246,1342,265,1603]" captionTargetPageId="16" captionText="Fig. 12. Interpretation of filtered MrBayes estimate of phylogeny as a metric tree (outgroup B. festivus Smith, 1861, not shown) for the subgenus Alpigenobombus Skorikov, 1914, from COI-like sequences from the sequences identified in black and orange in Fig. 11 together with Bayesian Poisson-tree-process (PTP) models re-applied for assessing support for species’ gene coalescents by maximum likelihood (for the 11 most likely candidate species). The scale bar is calibrated in substitutions per nucleotide site. Sequence labels and branch colours as in Fig. 10. Numbers above nodes are the Bayesian support values that all daughter haplotypes are parts of a single species. Grey spots show likely mitochondrial-to-nuclear transfers of the low-divergence numts accepted for estimating this tree. Asterisks mark sequences used as informal proxies for the type specimens of each of the taxon names in Table 2. To the right in grey are shown the interpretations of the PTP results as candidate species using the oldest available names for the species." figureDoi="http://doi.org/10.5281/zenodo.8382683" httpUri="https://zenodo.org/record/8382683/files/figure.png" pageId="34" pageNumber="35">Fig. 12</figureCitation>
, keys) support the interpretation that
<taxonomicName id="55EA4D13790D04179BD1FAA5FAA1FAFB" authorityName="Williams" authorityYear="2022" box="[1245,1349,1351,1376]" class="Insecta" family="Apidae" genus="Bombus" kingdom="Animalia" order="Hymenoptera" pageId="34" pageNumber="35" phylum="Arthropoda" rank="species" species="rainai">
<emphasis id="A09EEA82790D04179BD1FAA5FAA1FAFB" box="[1245,1349,1351,1376]" italics="true" pageId="34" pageNumber="35">B. rainai</emphasis>
</taxonomicName>
and
<taxonomicName id="55EA4D13790D04179FB0FA88FE92FA18" authorityName="Friese" authorityYear="1909" box="[188,374,1385,1411]" class="Insecta" family="Apidae" genus="Bombus" kingdom="Animalia" order="Hymenoptera" pageId="34" pageNumber="35" phylum="Arthropoda" rank="species" species="kashmirensis">
<emphasis id="A09EEA82790D04179FB0FA88FE92FA18" box="[188,374,1385,1411]" italics="true" pageId="34" pageNumber="35">B. kashmirensis</emphasis>
</taxonomicName>
are separate species (Williams 2022a), the available COI-barcode-like sequences may all be low-divergence neonumts (
<figureCitation id="0AD12A15790D04179D49FA6EFD7FFA3C" box="[581,667,1420,1447]" captionStart="Fig" captionStartId="14.[189,232,1681,1707]" captionTargetBox="[270,1317,267,1631]" captionTargetId="figure-11@14.[270,1317,265,1633]" captionTargetPageId="14" captionText="Fig. 11. Re-interpretation of sequences from Fig. 10, re-drawn with the percentage of nucleotides A and T at codon position 3 (AT3%) for each sequence given after the sequence label. Sequences are classified: (a) in purple – earlier-diverging groups of sequences without unique morphology that often have lower AT3% and shorter sequences that appear to duplicate species as pseudospecies in Fig. 10 – these sequences are interpreted as older, more divergent numts; (b) red – remaining sequences with low AT3% (&lt;95%) – these are interpreted as younger, less divergent numts; (c) orange – sequences from the red group but with the highest AT3% available for some taxa, selected to retain these taxa in the analysis; (d) grey and brown – sequences that are duplicate haplotypes for the black and orange sequences above and often shorter; (e) black – likely orthologous sequences with high AT3% (≥ 95%) and unique haplotypes." figureDoi="http://doi.org/10.5281/zenodo.8382681" httpUri="https://zenodo.org/record/8382681/files/figure.png" pageId="34" pageNumber="35">Fig. 11</figureCitation>
).
</paragraph>
<paragraph id="92553690790D04179FB1FA2CFDD0F9C9" blockId="34.[189,1399,1486,1618]" pageId="34" pageNumber="35">
Variation of
<taxonomicName id="55EA4D13790D04179E4AFA2DFE4FFA73" authorityName="Williams" authorityYear="2022" box="[326,427,1487,1512]" class="Insecta" family="Apidae" genus="Bombus" kingdom="Animalia" order="Hymenoptera" pageId="34" pageNumber="35" phylum="Arthropoda" rank="species" species="rainai">
<emphasis id="A09EEA82790D04179E4AFA2DFE4FFA73" box="[326,427,1487,1512]" italics="true" pageId="34" pageNumber="35">B. rainai</emphasis>
</taxonomicName>
is shown in the colour-pattern diagrams in Figs 49‒53.
<taxonomicName id="55EA4D13790D04179B12FA2CFB2CFA73" authorityName="Williams" authorityYear="2022" box="[1054,1224,1486,1512]" class="Insecta" family="Apidae" genus="Bombus" kingdom="Animalia" order="Hymenoptera" pageId="34" pageNumber="35" phylum="Arthropoda" rank="species" species="rainai">
<emphasis id="A09EEA82790D04179B12FA2CFB2CFA73" box="[1054,1224,1486,1512]" italics="true" pageId="34" pageNumber="35">Bombus rainai</emphasis>
</taxonomicName>
, with its white-banded red-tailed colour pattern in the Himalaya, appears to mimic the abundant
<taxonomicName id="55EA4D13790D04179B86FA10FE28F9B5" authority="Smith, 1878" authorityName="Smith" authorityYear="1878" class="Insecta" family="Apidae" genus="Bombus" kingdom="Animalia" order="Hymenoptera" pageId="34" pageNumber="35" phylum="Arthropoda" rank="species" species="longiceps" subGenus="Sibiricobombus">
<emphasis id="A09EEA82790D04179B86FA10FB47F990" box="[1162,1187,1522,1547]" italics="true" pageId="34" pageNumber="35">B.</emphasis>
(
<emphasis id="A09EEA82790D04179BB9FA13FA89F990" box="[1205,1389,1521,1547]" italics="true" pageId="34" pageNumber="35">Sibiricobombus</emphasis>
)
<emphasis id="A09EEA82790D04179FB1F9F6FEC9F9B5" box="[189,301,1556,1582]" italics="true" pageId="34" pageNumber="35">longiceps</emphasis>
Smith, 1878
</taxonomicName>
(until recently regarded as part of
<taxonomicName id="55EA4D13790D04179C70F9F7FB33F9B4" authority="Morawitz, 1875" authorityName="Morawitz" authorityYear="1875" box="[892,1239,1556,1583]" class="Insecta" family="Apidae" genus="Bombus" kingdom="Animalia" order="Hymenoptera" pageId="34" pageNumber="35" phylum="Arthropoda" rank="species" species="asiaticus">
<emphasis id="A09EEA82790D04179C70F9F7FBEDF9B5" box="[892,1033,1557,1582]" italics="true" pageId="34" pageNumber="35">B. asiaticus</emphasis>
Morawitz, 1875
</taxonomicName>
), and similar species (
<bibRefCitation id="F67B4B61790D04179E2DF9D5FE30F9C9" author="Williams P. H." box="[289,468,1591,1618]" pageId="34" pageNumber="35" pagination="97 - 118" refId="ref38392" refString="Williams P. H. 2007. The distribution of bumblebee colour patterns world-wide: possible significance for thermoregulation, crypsis, and warning mimicry. Biological Journal of the Linnean Society 92: 97 - 118. https: // doi. org / 10.1111 / j. 1095 - 8312.2007.00878. x" type="journal article" year="2007">Williams 2007</bibRefCitation>
: fig. 5l).
</paragraph>
</subSubSection>
<subSubSection id="DAF0651B790D04179FB1F99BFDAEF97E" pageId="34" pageNumber="35" type="materials_examined">
<paragraph id="92553690790D04179FB1F99BFE88F908" blockId="34.[189,364,1657,1683]" box="[189,364,1657,1683]" pageId="34" pageNumber="35">
<heading id="C91D81FC790D04179FB1F99BFE88F908" bold="true" box="[189,364,1657,1683]" fontSize="11" level="3" pageId="34" pageNumber="35" reason="3">
<emphasis id="A09EEA82790D04179FB1F99BFE88F908" bold="true" box="[189,364,1657,1683]" pageId="34" pageNumber="35">
<typeStatus id="4D518832790D04179FB1F99BFF1EF908" box="[189,250,1657,1683]" pageId="34" pageNumber="35">Type</typeStatus>
material
</emphasis>
</heading>
</paragraph>
<paragraph id="92553690790D04179FB1F945FDAEF97E" blockId="34.[189,1398,1703,1765]" pageId="34" pageNumber="35">
<taxonomicName id="55EA4D13790D04179FB1F945FCB0F959" authority="Williams 2022 a: 132" authorityName="Williams" authorityPageNumber="132" authorityYear="2022" box="[189,852,1703,1730]" class="Insecta" family="Apidae" genus="Bombus" kingdom="Animalia" order="Hymenoptera" pageId="34" pageNumber="35" phylum="Arthropoda" rank="species" species="rainai" subGenus="Alpigenobombus">
<emphasis id="A09EEA82790D04179FB1F945FEF8F95A" box="[189,284,1703,1729]" italics="true" pageId="34" pageNumber="35">Bombus</emphasis>
(
<emphasis id="A09EEA82790D04179E23F945FE10F95A" box="[303,500,1703,1729]" italics="true" pageId="34" pageNumber="35">Alpigenobombus</emphasis>
)
<emphasis id="A09EEA82790D04179D0BF94AFDABF95A" box="[519,591,1704,1729]" italics="true" pageId="34" pageNumber="35">rainai</emphasis>
Williams 2022a: 132
</taxonomicName>
.
<materialsCitation id="22823CCD790D04179C6EF945FDA2F97E" ID-GBIF-Occurrence="4428536351" collectionCode="ZSC" country="India" location="Kashmir" pageId="34" pageNumber="35" specimenCount="1" specimenCount-female="1" typeStatus="holotype">
<typeStatus id="4D518832790D04179C6EF945FC2BF95A" box="[866,975,1703,1729]" pageId="34" pageNumber="35" type="holotype">Holotype</typeStatus>
by original designation:
<specimenCount id="84ECFD19790D04179BF1F94AFAF7F959" box="[1277,1299,1704,1730]" pageId="34" pageNumber="35" type="female">♀</specimenCount>
(queen)
<location id="9735604B790D04179FB1F929FEC0F97E" LSID="urn:lsid:plazi:treatment:1A438786790D04169EECFCC2FCA7FB24:9735604B790D04179FB1F929FEC0F97E" box="[189,292,1739,1765]" country="India" name="Kashmir" pageId="34" pageNumber="35">Kashmir</location>
,
<collectingCountry id="EAFD7600790D04179E23F929FE8FF97E" box="[303,363,1739,1765]" name="India" pageId="34" pageNumber="35">India</collectingCountry>
(
<collectionCode id="F4FBAE55790D04179E70F929FE5CF97E" box="[380,440,1739,1765]" pageId="34" pageNumber="35">ZSC</collectionCode>
). Examined
</materialsCitation>
.
</paragraph>
</subSubSection>
<subSubSection id="DAF0651B790D04169FB1F8EEFEF1FD39" lastPageId="35" lastPageNumber="36" pageId="34" pageNumber="35" type="diagnosis">
<paragraph id="92553690790D04179FB1F8EEFE14F8BD" blockId="34.[189,496,1804,1830]" box="[189,496,1804,1830]" pageId="34" pageNumber="35">
<heading id="C91D81FC790D04179FB1F8EEFE14F8BD" bold="true" box="[189,496,1804,1830]" fontSize="11" level="3" pageId="34" pageNumber="35" reason="3">
<emphasis id="A09EEA82790D04179FB1F8EEFE14F8BD" bold="true" box="[189,496,1804,1830]" pageId="34" pageNumber="35">Morphological diagnosis</emphasis>
</heading>
</paragraph>
<paragraph id="92553690790D04179FE6F8D9FEA1F8CE" blockId="34.[189,1401,1851,2018]" box="[234,325,1851,1877]" pageId="34" pageNumber="35">
<emphasis id="A09EEA82790D04179FE6F8D9FEA1F8CE" bold="true" box="[234,325,1851,1877]" pageId="34" pageNumber="35">Female</emphasis>
</paragraph>
<paragraph id="92553690790D04169FB1F8BCFAF3FEDD" blockId="34.[189,1401,1851,2018]" lastBlockId="35.[188,1398,265,327]" lastPageId="35" lastPageNumber="36" pageId="34" pageNumber="35">
Wings nearly clear with veins dark brown, hair medium length, oculo-malar area shorter than broad, clypeus adjacent to the labrum slightly raised in the centre so that the transverse anterior groove is interrupted, this central area with few small punctures and more shining than the surrounding areas, oculo-ocellar area in its outer half with only a few small shallow punctures, any larger punctures confined to the eye margin (cf.
<taxonomicName id="55EA4D13790C04169EBBFEE8FD94FEB8" authorityName="Friese" authorityYear="1909" box="[439,624,265,291]" class="Insecta" family="Apidae" genus="Bombus" kingdom="Animalia" order="Hymenoptera" pageId="35" pageNumber="36" phylum="Arthropoda" rank="species" species="kashmirensis">
<emphasis id="A09EEA82790C04169EBBFEE8FD94FEB8" box="[439,624,265,291]" italics="true" pageId="35" pageNumber="36">B. kashmirensis</emphasis>
</taxonomicName>
); hair of the side of the thorax in no more than its upper third white and the scutellum predominantly black, hair of T5 entirely orange without obviously paler tips.
</paragraph>
<paragraph id="92553690790C04169FE6FE8CFECEFE13" blockId="35.[188,1400,366,674]" box="[234,298,366,392]" pageId="35" pageNumber="36">
<emphasis id="A09EEA82790C04169FE6FE8CFECEFE13" bold="true" box="[234,298,366,392]" pageId="35" pageNumber="36">Male</emphasis>
</paragraph>
<paragraph id="92553690790C04169FB1FE73FEF1FD39" blockId="35.[188,1400,366,674]" pageId="35" pageNumber="36">
Wings nearly clear with veins dark brown, hair medium length, oculo-malar area shorter than broad; genitalia (
<figureCitation id="0AD12A15790C04169E38FE56FE73FE55" box="[308,407,436,462]" captionStart="Figs 106–115" captionStartId="29.[189,242,1752,1778]" captionTargetBox="[248,1333,302,1671]" captionTargetId="figure-11@29.[236,1352,265,1701]" captionTargetPageId="29" captionText="Figs 106–115. Morphology of the male genitalia for species of the subgenus Alpigenobombus Skorikov, 1914, from the dorsal aspect, anterior at the bottom of the image, posterior at the top. 106. Bombus wurflenii Radoszkowski, 1860, Turkey. 107. B mastrucatus Gerstaecker, 1869, Austria. 108. B. kashmirensis Friese, 1909, India-Kashmir. 109. B. rainai Williams, 2022, India-Kashmir. 110. B. sikkimi Friese, 1918, Nepal. 111. B. nobilis Friese, 1905, China-Sichuan. 112. B. validus Friese, 1905, China-Gansu. 113. B. genalis Friese, 1918, China-Yunnan. 114. B. breviceps Smith, 1852, China- Sichuan. 115. B. grahami (Frison, 1933) China-Sichuan (left penis-valve recurved hook missing). Scale bars = 1 mm." figureDoi="http://doi.org/10.5281/zenodo.8382695" httpUri="https://zenodo.org/record/8382695/files/figure.png" pageId="35" pageNumber="36">Fig. 109</figureCitation>
) with the gonostylus long and distally near its midline axis convexly rounded, length on its outer side about a quarter as long as on its inner side with the two inner corners of the distal lobe rounded, the margin between them convex, penis-valve head strongly recurved, the recurved hook much longer than broad and narrow, tapering gradually to the apex (cf.
<taxonomicName id="55EA4D13790C04169CC8FDFCFB99FDA3" authorityName="Friese" authorityYear="1909" box="[964,1149,542,568]" class="Insecta" family="Apidae" genus="Bombus" kingdom="Animalia" order="Hymenoptera" pageId="35" pageNumber="36" phylum="Arthropoda" rank="species" species="kashmirensis">
<emphasis id="A09EEA82790C04169CC8FDFCFB99FDA3" box="[964,1149,542,568]" italics="true" pageId="35" pageNumber="36">B. kashmirensis</emphasis>
</taxonomicName>
); male eye distinctly enlarged relative to female eye; hair of the side of the thorax with black and white intermixed, but often predominantly black in its lower half, hair of the sides of T5‒6 orange, hair of T3 often predominantly orange.
</paragraph>
</subSubSection>
<subSubSection id="DAF0651B790C04169FB1FD2AFCC5FCC3" pageId="35" pageNumber="36" type="materials_examined">
<paragraph id="92553690790C04169FB1FD2AFDCAFD79" blockId="35.[189,558,712,739]" box="[189,558,712,739]" pageId="35" pageNumber="36">
<heading id="C91D81FC790C04169FB1FD2AFDCAFD79" bold="true" box="[189,558,712,739]" fontSize="11" level="3" pageId="35" pageNumber="36" reason="3">
<emphasis id="A09EEA82790C04169FB1FD2AFDCAFD79" bold="true" box="[189,558,712,739]" pageId="35" pageNumber="36">
Material sequenced in
<figureCitation id="0AD12A15790C04169ED5FD2AFDCAFD79" box="[473,558,712,738]" captionStart="Fig" captionStartId="16.[189,232,1646,1672]" captionTargetBox="[246,1342,268,1600]" captionTargetId="figure-10@16.[246,1342,265,1603]" captionTargetPageId="16" captionText="Fig. 12. Interpretation of filtered MrBayes estimate of phylogeny as a metric tree (outgroup B. festivus Smith, 1861, not shown) for the subgenus Alpigenobombus Skorikov, 1914, from COI-like sequences from the sequences identified in black and orange in Fig. 11 together with Bayesian Poisson-tree-process (PTP) models re-applied for assessing support for species’ gene coalescents by maximum likelihood (for the 11 most likely candidate species). The scale bar is calibrated in substitutions per nucleotide site. Sequence labels and branch colours as in Fig. 10. Numbers above nodes are the Bayesian support values that all daughter haplotypes are parts of a single species. Grey spots show likely mitochondrial-to-nuclear transfers of the low-divergence numts accepted for estimating this tree. Asterisks mark sequences used as informal proxies for the type specimens of each of the taxon names in Table 2. To the right in grey are shown the interpretations of the PTP results as candidate species using the oldest available names for the species." figureDoi="http://doi.org/10.5281/zenodo.8382683" httpUri="https://zenodo.org/record/8382683/files/figure.png" pageId="35" pageNumber="36">Fig. 12</figureCitation>
</emphasis>
</heading>
</paragraph>
<paragraph id="92553690790C04169FB1FD15FCC5FCC3" blockId="35.[189,1399,759,856]" pageId="35" pageNumber="36">
<materialsCitation id="22823CCD790C04169FB1FD15FDF3FCAE" ID-GBIF-Occurrence="4428536386" collectingDate="2009-06-26" collectionCode="ZSC" collectorName="R. Raina" country="India" latitude="34.636" location="Kashmir" longLatPrecision="71" longitude="74.864" municipality="Achoora" pageId="35" pageNumber="36" specimenCode="AG#080" specimenCount="1" specimenCount-female="1">
<collectingCountry id="EAFD7600790C04169FB1FD15FEEAFC8A" box="[189,270,759,785]" name="India" pageId="35" pageNumber="36">INDIA</collectingCountry>
•
<specimenCount id="84ECFD19790C04169E27FD15FEBDFC89" box="[299,345,759,786]" pageId="35" pageNumber="36" type="female">1 ♀</specimenCount>
(queen);
<location id="9735604B790C04169EC2FD15FDD1FC8A" LSID="urn:lsid:plazi:treatment:1A438786790D04169EECFCC2FCA7FB24:9735604B790C04169EC2FD15FDD1FC8A" box="[462,565,759,785]" country="India" latitude="34.636" longLatPrecision="71" longitude="74.864" municipality="Achoora" name="Kashmir" pageId="35" pageNumber="36">Kashmir</location>
,
<collectingMunicipality id="7231ACEA790C04169D4DFD15FD4EFC8A" box="[577,682,759,785]" pageId="35" pageNumber="36">Achoora</collectingMunicipality>
;
<geoCoordinate id="F7DE5057790C04169DBBFD15FCD1FC89" box="[695,821,759,786]" degrees="34.636" direction="north" orientation="latitude" pageId="35" pageNumber="36" precision="55" value="34.636">34.636° N</geoCoordinate>
,
<geoCoordinate id="F7DE5057790C04169C4FFD15FC59FC89" box="[835,957,759,786]" degrees="74.864" direction="east" orientation="longitude" pageId="35" pageNumber="36" precision="55" value="74.864">74.864° E</geoCoordinate>
;
<date id="E6541050790C04169CC0FD15FB8EFC8A" box="[972,1130,759,786]" pageId="35" pageNumber="36" value="2009-06-26">
<collectingDate id="F610E9B8790C04169CC0FD15FB8EFC8A" box="[972,1130,759,786]" pageId="35" pageNumber="36" value="2009-06-26">26 Jun. 2009</collectingDate>
</date>
;
<collectorName id="3F1F5346790C04169B74FD1AFB04FC8A" box="[1144,1248,759,786]" pageId="35" pageNumber="36">R. Raina</collectorName>
leg.; BOLD seq: 1552E01;
<collectionCode id="F4FBAE55790C04169E61FCF9FE43FCAE" box="[365,423,795,821]" pageId="35" pageNumber="36">ZSC</collectionCode>
:
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</materialsCitation>
•
<materialsCitation id="22823CCD790C04169D3CFCF9FCF9FCC3" ID-GBIF-Occurrence="4428536317" collectingDate="2009-08-21" collectionCode="ZSC" collectorName="R. Raina" country="India" latitude="34.055" location="Kashmir" longLatPrecision="72" longitude="74.386" municipality="Apharwat" pageId="35" pageNumber="36" specimenCode="AG#081" specimenCount="1" specimenCount-female="1">
<specimenCount id="84ECFD19790C04169D3CFCF9FDB8FCAE" box="[560,604,795,821]" pageId="35" pageNumber="36" type="female">1 ♀</specimenCount>
(queen);
<location id="9735604B790C04169DC1FCF8FCD0FCAF" LSID="urn:lsid:plazi:treatment:1A438786790D04169EECFCC2FCA7FB24:9735604B790C04169DC1FCF8FCD0FCAF" box="[717,820,794,820]" country="India" latitude="34.055" longLatPrecision="72" longitude="74.386" municipality="Apharwat" name="Kashmir" pageId="35" pageNumber="36">Kashmir</location>
,
<collectingMunicipality id="7231ACEA790C04169C32FCF8FC52FCAF" box="[830,950,794,820]" pageId="35" pageNumber="36">Apharwat</collectingMunicipality>
;
<geoCoordinate id="F7DE5057790C04169CCDFCF8FBD9FCAE" box="[961,1085,794,821]" degrees="34.055" direction="north" orientation="latitude" pageId="35" pageNumber="36" precision="55" value="34.055">34.055° N</geoCoordinate>
,
<geoCoordinate id="F7DE5057790C04169B45FCF8FB26FCAE" box="[1097,1218,794,821]" degrees="74.386" direction="east" orientation="longitude" pageId="35" pageNumber="36" precision="55" value="74.386">74.386° E</geoCoordinate>
;
<date id="E6541050790C04169BC3FCF9FA96FCAE" box="[1231,1394,795,821]" pageId="35" pageNumber="36" value="2009-08-21">
<collectingDate id="F610E9B8790C04169BC3FCF9FA96FCAE" box="[1231,1394,795,821]" pageId="35" pageNumber="36" value="2009-08-21">21 Aug. 2009</collectingDate>
</date>
;
<collectorName id="3F1F5346790C04169FB1FCDCFEC7FCC3" box="[189,291,830,856]" pageId="35" pageNumber="36">R. Raina</collectorName>
leg.; BOLD seq: 1552D12;
<collectionCode id="F4FBAE55790C04169D7CFCDCFD4EFCC3" box="[624,682,830,856]" pageId="35" pageNumber="36">ZSC</collectionCode>
:
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.
</paragraph>
</subSubSection>
<subSubSection id="DAF0651B790C04169FB1FC9DFDF6FBB7" pageId="35" pageNumber="36" type="distribution">
<paragraph id="92553690790C04169FB1FC9DFE49FC02" blockId="35.[189,429,895,921]" box="[189,429,895,921]" pageId="35" pageNumber="36">
<heading id="C91D81FC790C04169FB1FC9DFE49FC02" bold="true" box="[189,429,895,921]" fontSize="11" level="3" pageId="35" pageNumber="36" reason="3">
<emphasis id="A09EEA82790C04169FB1FC9DFE49FC02" bold="true" box="[189,429,895,921]" pageId="35" pageNumber="36">Global distribution</emphasis>
</heading>
</paragraph>
<paragraph id="92553690790C04169FB1FC4CFC32FC53" blockId="35.[189,982,941,968]" box="[189,982,941,968]" pageId="35" pageNumber="36">
Western Himalaya:
<collectingCountry id="EAFD7600790C04169EA9FC4FFDEFFC5C" box="[421,523,941,967]" name="Pakistan" pageId="35" pageNumber="36">Pakistan</collectingCountry>
,
<collectingCountry id="EAFD7600790C04169D1BFC4FFDB7FC5C" box="[535,595,941,967]" name="India" pageId="35" pageNumber="36">India</collectingCountry>
(Kashmir): NHMUK, PW, ZSC.
</paragraph>
<paragraph id="92553690790C04169FB1FC0DFDF6FBB7" blockId="35.[189,1399,1007,1069]" pageId="35" pageNumber="36">
This species is recorded at elevations of
<quantity id="55129B75790C04169DACFC0DFCA2FB91" box="[672,838,1007,1034]" metricMagnitude="3" metricUnit="m" metricValue="3.371" metricValueMax="4.2" metricValueMin="2.542" pageId="35" pageNumber="36" unit="m" value="3371.0" valueMax="4200.0" valueMin="2542.0">2542‒4200 m</quantity>
, primarily above the tree line and high into the alpine zone (
<bibRefCitation id="F67B4B61790C04169E5FFBF0FDE1FBB7" author="Williams P. H." box="[339,517,1042,1068]" pageId="35" pageNumber="36" pagination="1 - 204" refId="ref38198" refString="Williams P. H. 1991. The bumble bees of the Kashmir Himalaya (Hymenoptera: Apidae, Bombini). Bulletin of the British Museum (Natural History) (Entomology) 60: 1 - 204. Available from https: // www. biodiversitylibrary. org / page / 41055963 [accessed 13 Aug. 2023]." type="journal article" year="1991">Williams 1991</bibRefCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="DAF0651B790C04169FB1FBB1FCA7FB24" pageId="35" pageNumber="36" type="biology_ecology">
<paragraph id="92553690790C04169FB1FBB1FEDBFBF6" blockId="35.[189,319,1107,1133]" box="[189,319,1107,1133]" pageId="35" pageNumber="36">
<heading id="C91D81FC790C04169FB1FBB1FEDBFBF6" bold="true" box="[189,319,1107,1133]" fontSize="11" level="3" pageId="35" pageNumber="36" reason="3">
<emphasis id="A09EEA82790C04169FB1FBB1FEDBFBF6" bold="true" box="[189,319,1107,1133]" pageId="35" pageNumber="36">Behaviour</emphasis>
</heading>
</paragraph>
<paragraph id="92553690790C04169FB1FB60FCA7FB24" blockId="35.[189,1398,1154,1216]" pageId="35" pageNumber="36">
Male eye distinctly enlarged relative to female eye: males are expected to show ‘cruising’ behaviour when searching for mates in Kashmir (
<bibRefCitation id="F67B4B61790C04169D8FFB47FCD2FB24" author="Williams P. H." box="[643,822,1189,1215]" pageId="35" pageNumber="36" pagination="1 - 204" refId="ref38198" refString="Williams P. H. 1991. The bumble bees of the Kashmir Himalaya (Hymenoptera: Apidae, Bombini). Bulletin of the British Museum (Natural History) (Entomology) 60: 1 - 204. Available from https: // www. biodiversitylibrary. org / page / 41055963 [accessed 13 Aug. 2023]." type="journal article" year="1991">Williams 1991</bibRefCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>
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