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<document ID-DOI="http://dx.doi.org/10.3897/zookeys.520.10134" ID-GBIF-Dataset="117bbf8b-77ab-452c-9ed9-2c0f4ff43f8e" ID-PMC="PMC4591724" ID-Pensoft-Pub="1313-2970-520-109" ID-PubMed="26448715" ID-ZBK="A3F77DC0CA0F48149F097FE1A6042447" ModsDocAuthor="" ModsDocDate="2015" ModsDocID="1313-2970-520-109" ModsDocOrigin="ZooKeys 520" ModsDocTitle="Male secondary sexual structures and the systematics of the Thereusoppia species group (Lepidoptera, Lycaenidae, Eumaeini)" checkinTime="1451243986484" checkinUser="pensoft" docAuthor="Robbins, Robert K., Heredia, Maria Dolores &amp; Busby, Robert C." docDate="2015" docId="E145D79DC7E173C36933A94284834BCF" docLanguage="en" docName="ZooKeys 520: 109-130" docOrigin="ZooKeys 520" docSource="http://dx.doi.org/10.3897/zookeys.520.10134" docTitle="Thereus Huebner 1819" docType="treatment" docVersion="3" lastPageNumber="113" masterDocId="FF9D895BFFF3BC5E281FFF8CFFA8FFD9" masterDocTitle="Male secondary sexual structures and the systematics of the Thereusoppia species group (Lepidoptera, Lycaenidae, Eumaeini)" masterLastPageNumber="130" masterPageNumber="109" pageNumber="112" updateTime="1668160661356" updateUser="ExternalLinkService">
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<mods:title>Male secondary sexual structures and the systematics of the Thereusoppia species group (Lepidoptera, Lycaenidae, Eumaeini)</mods:title>
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<mods:name type="personal">
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Robbins, Robert K.</mods:namePart>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Heredia, Maria Dolores</mods:namePart>
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<mods:name type="personal">
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Busby, Robert C.</mods:namePart>
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<mods:title>ZooKeys</mods:title>
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<mods:date>2015</mods:date>
<mods:detail type="volume">
<mods:number>520</mods:number>
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<mods:start>109</mods:start>
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<mods:url>http://dx.doi.org/10.3897/zookeys.520.10134</mods:url>
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<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">http://dx.doi.org/10.3897/zookeys.520.10134</mods:identifier>
<mods:identifier type="Pensoft-Pub">1313-2970-520-109</mods:identifier>
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<treatment ID-GBIF-Taxon="152062294" LSID="urn:lsid:plazi:treatment:E145D79DC7E173C36933A94284834BCF" httpUri="http://treatment.plazi.org/id/E145D79DC7E173C36933A94284834BCF" lastPageId="4" lastPageNumber="113" pageId="3" pageNumber="112">
<subSubSection pageId="3" pageNumber="112" type="multiple">
<paragraph pageId="3" pageNumber="112">Taxon classification Animalia Lepidoptera Lycaenidae</paragraph>
</subSubSection>
<subSubSection pageId="3" pageNumber="112" type="nomenclature">
<paragraph pageId="3" pageNumber="112">
<taxonomicName authority="Huebner, [1819]" class="Insecta" family="Lycaenidae" genus="Thereus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Thereus" order="Lepidoptera" pageId="3" pageNumber="112" phylum="Arthropoda" rank="genus">
Thereus
<normalizedToken originalValue="Hübner">Huebner</normalizedToken>
, [1819]
</taxonomicName>
</paragraph>
</subSubSection>
<subSubSection pageId="3" pageNumber="112" type="type species">
<paragraph pageId="3" pageNumber="112">Type species.</paragraph>
<paragraph pageId="3" pageNumber="112">
<taxonomicName class="Insecta" family="Papilionidae" genus="Papilio" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Papilio lausus" order="Lepidoptera" pageId="3" pageNumber="112" phylum="Arthropoda" rank="species" species="lausus">Papilio lausus</taxonomicName>
Cramer
</paragraph>
</subSubSection>
<subSubSection pageId="3" pageNumber="112" type="diagnosis">
<paragraph pageId="3" pageNumber="112">Diagnosis.</paragraph>
<paragraph pageId="3" pageNumber="112">
<bibRefCitation author="Robbins, RK" journalOrPublisher="Smithsonian Contribution to Zoology" pageId="19" pageNumber="128" pagination="1 - 64" title="Evolution, comparative morphology, and identification of the eumaeine butterfly genus Rekoa Kaye (Lycaenidae: Theclinae)." url="10.5479/si.00810282.498" volume="498" year="1991">Robbins (1991)</bibRefCitation>
characterized
<taxonomicName class="Insecta" family="Lycaenidae" genus="Thereus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Thereus" order="Lepidoptera" pageId="3" pageNumber="112" phylum="Arthropoda" rank="genus">Thereus</taxonomicName>
by (1) a pair of sclerotized invaginations on the membrane attached to the ventro-lateral sides of the papillae anales (Figs 23-24, figured in
<bibRefCitation author="Robbins, RK" journalOrPublisher="Proceedings Entomological Society of Washington" pageId="19" pageNumber="128" pagination="162 - 169" title="The New World hairstreak genus Arawacus Kaye (Lepidoptera: Lycaenidae: Theclinae: Eumaeini)." volume="102" year="2000">Robbins 2000</bibRefCitation>
), (2) a pair of ventro-lateral brush organs (often inconspicuous) in addition to the pair of dorsal ones (Figs 15-18), and (3) the number of antennal nudum segments (as defined in the methods section) is sexually dimorphic, being greater in females than that in males by five or more segments (Fig. 25). The second trait is lacking in
<taxonomicName class="Insecta" family="Lycaenidae" genus="Thereus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Thereus pseudarcula" order="Lepidoptera" pageId="3" pageNumber="112" phylum="Arthropoda" rank="species" species="pseudarcula">Thereus pseudarcula</taxonomicName>
(Giacomelli), suggesting that this species is the phylogenetic sister to the remainder of the genus. Interestingly, it is the only
<taxonomicName class="Insecta" family="Lycaenidae" genus="Thereus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Thereus" order="Lepidoptera" pageId="3" pageNumber="112" phylum="Arthropoda" rank="genus">Thereus</taxonomicName>
species that appears to be a subtropical endemic. During this study, we found an instance in which sexual dimorphism of antennal nudum length was four segments-not five or greater-so the generic diagnosis is modified accordingly.
</paragraph>
<caption pageId="3" pageNumber="112">
<paragraph pageId="3" pageNumber="112">
Figure 25. Male (top) and female antennae of
<taxonomicName class="Insecta" family="Lycaenidae" genus="Thereus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Thereus lomalarga" order="Lepidoptera" pageId="3" pageNumber="112" phylum="Arthropoda" rank="species" species="lomalarga">Thereus lomalarga</taxonomicName>
in ventral aspect showing nudum extent. The male has 14 nudum segments in contrast to 21 nudum segments in the female. Scale bar: 2 mm.
</paragraph>
</caption>
<paragraph pageId="3" pageNumber="112">
<bibRefCitation author="Robbins, RK" journalOrPublisher="Smithsonian Contribution to Zoology" pageId="19" pageNumber="128" pagination="1 - 64" title="Evolution, comparative morphology, and identification of the eumaeine butterfly genus Rekoa Kaye (Lycaenidae: Theclinae)." url="10.5479/si.00810282.498" volume="498" year="1991">Robbins (1991)</bibRefCitation>
noted that
<taxonomicName class="Insecta" family="Lycaenidae" genus="Thereus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Thereus" order="Lepidoptera" pageId="3" pageNumber="112" phylum="Arthropoda" rank="genus">Thereus</taxonomicName>
contained 27 species, but more than a decade later, 31 were listed, of which nine were undescribed and two had questionable generic placement (
<bibRefCitation author="Robbins, RK" editor="Lamas, G" journalOrPublisher="Association for Tropical Lepidoptera and Scientific Publishers, Gainesville, FL" pageId="19" pageNumber="128" pagination="118 - 137" title="Lycaenidae. Theclinae. Tribe Eumaeini." volumeTitle="Checklist: Part 4 A. Hesperioidea - Papilionoidea" year="2004">Robbins 2004</bibRefCitation>
).
<bibRefCitation author="Balint, Z" journalOrPublisher="Universidad de Caldas" pageId="18" pageNumber="127" pagination="278 - 294" title="Systematics and taxonomic notes on Neotropical hairstreak lycaenids in connection to the genus Megathecla (Lycaenidae: Theclinae: Eumaeini). Boletin cientifico. Museo de Historia Natural." volume="9" year="2005">
<normalizedToken originalValue="Bálint">Balint</normalizedToken>
(2005)
</bibRefCitation>
proposed a new specific name, and we now recognize 35 species, of which eleven are undescribed and two have questionable generic placement. This paper begins the task of recognizing species groups, which facilitates the description of new species, and of determining the generic placement of the species with questionable generic placement.
</paragraph>
</subSubSection>
<subSubSection pageId="3" pageNumber="112" type="nomenclature">
<paragraph pageId="3" pageNumber="112">Nomenclature.</paragraph>
<paragraph pageId="3" pageNumber="112">
<bibRefCitation author="Robbins, RK" journalOrPublisher="Smithsonian Contribution to Zoology" pageId="19" pageNumber="128" pagination="1 - 64" title="Evolution, comparative morphology, and identification of the eumaeine butterfly genus Rekoa Kaye (Lycaenidae: Theclinae)." url="10.5479/si.00810282.498" volume="498" year="1991">Robbins (1991)</bibRefCitation>
accorded
<taxonomicName class="Insecta" family="Lycaenidae" genus="Thereus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Thereus" order="Lepidoptera" pageId="3" pageNumber="112" phylum="Arthropoda" rank="genus">Thereus</taxonomicName>
<normalizedToken originalValue="Hübner">Huebner</normalizedToken>
priority over
<taxonomicName class="Insecta" family="Lycaenidae" genus="Molus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Molus" order="Lepidoptera" pageId="3" pageNumber="112" phylum="Arthropoda" rank="genus">Molus</taxonomicName>
<normalizedToken originalValue="Hübner">Huebner</normalizedToken>
and synonymized
<taxonomicName class="Insecta" family="Lycaenidae" genus="Noreena" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Noreena" order="Lepidoptera" pageId="3" pageNumber="112" phylum="Arthropoda" rank="genus">Noreena</taxonomicName>
K. Johnson, MacPherson &amp; Ingraham. Subsequently proposed names
<taxonomicName class="Insecta" family="Lycaenidae" genus="Solanorum" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Solanorum" order="Lepidoptera" pageId="3" pageNumber="112" phylum="Arthropoda" rank="genus">Solanorum</taxonomicName>
Johnson,
<taxonomicName class="Insecta" family="Lycaenidae" genus="Timokla" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Timokla" order="Lepidoptera" pageId="3" pageNumber="112" phylum="Arthropoda" rank="genus">Timokla</taxonomicName>
Johnson, Kruse &amp; Kroenlein, and
<taxonomicName genus="Pedusa" lsidName="Pedusa" pageId="3" pageNumber="112" rank="genus">Pedusa</taxonomicName>
<normalizedToken originalValue="dAbrera">d'Abrera</normalizedToken>
were synonymized later (
<bibRefCitation author="Robbins, RK" editor="Lamas, G" journalOrPublisher="Association for Tropical Lepidoptera and Scientific Publishers, Gainesville, FL" pageId="19" pageNumber="128" pagination="118 - 137" title="Lycaenidae. Theclinae. Tribe Eumaeini." volumeTitle="Checklist: Part 4 A. Hesperioidea - Papilionoidea" year="2004">Robbins 2004</bibRefCitation>
).
</paragraph>
</subSubSection>
<subSubSection pageId="3" pageNumber="112" type="distribution">
<paragraph pageId="3" pageNumber="112">Distribution.</paragraph>
<paragraph pageId="3" pageNumber="112">
<taxonomicName class="Insecta" family="Lycaenidae" genus="Thereus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Thereus" order="Lepidoptera" pageId="3" pageNumber="112" phylum="Arthropoda" rank="genus">Thereus</taxonomicName>
occurs throughout the Neotropics from northern Mexico to Uruguay and Argentina.
<taxonomicName class="Insecta" family="Lycaenidae" genus="Thereus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Thereus lausus" order="Lepidoptera" pageId="3" pageNumber="112" phylum="Arthropoda" rank="species" species="lausus">Thereus lausus</taxonomicName>
,
<taxonomicName class="Insecta" family="Lycaenidae" genus="Thereus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Thereus cithonius" order="Lepidoptera" pageId="3" pageNumber="112" phylum="Arthropoda" rank="species" species="cithonius">Thereus cithonius</taxonomicName>
, and
<taxonomicName class="Insecta" family="Lycaenidae" genus="Thereus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Thereus ortalus" order="Lepidoptera" pageId="3" pageNumber="112" phylum="Arthropoda" rank="species" species="ortalus">Thereus ortalus</taxonomicName>
range widely from Mexico to southern Brazil, but distributions of species are otherwise more restricted. Approximately 2/3 of the species in the genus occur in the Amazon Region, as demarcated by
<bibRefCitation author="Brown, KS" journalOrPublisher="American Zoologist" pageId="18" pageNumber="127" pagination="453 - 471" title="Historical and ecological factors in the biogeography of aposematic neotropical butterflies." url="10.1093/icb/22.2.453" volume="22" year="1982">Brown (1982)</bibRefCitation>
.
</paragraph>
</subSubSection>
<subSubSection pageId="4" pageNumber="113" type="habitat">
<paragraph pageId="4" pageNumber="113">
<pageBreakToken pageId="4" pageNumber="113" start="start">Habitat</pageBreakToken>
.
</paragraph>
<paragraph pageId="4" pageNumber="113">
Most species inhabit wet lowland forest, with only a few exceptions.
<taxonomicName class="Insecta" family="Lycaenidae" genus="Thereus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Thereus gabathana" order="Lepidoptera" pageId="4" pageNumber="113" phylum="Arthropoda" rank="species" species="gabathana">Thereus gabathana</taxonomicName>
(Strand),
<taxonomicName class="Insecta" family="Lycaenidae" genus="Thereus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Thereus wojtusiaki" order="Lepidoptera" pageId="4" pageNumber="113" phylum="Arthropoda" rank="species" species="wojtusiaki">Thereus wojtusiaki</taxonomicName>
<normalizedToken originalValue="Bálint">Balint</normalizedToken>
,
<taxonomicName class="Insecta" family="Lycaenidae" genus="Thereus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Thereus orasus" order="Lepidoptera" pageId="4" pageNumber="113" phylum="Arthropoda" rank="species" species="orasus">Thereus orasus</taxonomicName>
(Godman &amp; Salvin), and an undescribed species are montane endemics, and
<taxonomicName class="Insecta" family="Lycaenidae" genus="Thereus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Thereus pseudarcula" order="Lepidoptera" pageId="4" pageNumber="113" phylum="Arthropoda" rank="species" species="pseudarcula">Thereus pseudarcula</taxonomicName>
is subtropical. The widespread
<taxonomicName class="Insecta" family="Lycaenidae" genus="Thereus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Thereus cithonius" order="Lepidoptera" pageId="4" pageNumber="113" phylum="Arthropoda" rank="species" species="cithonius">Thereus cithonius</taxonomicName>
occurs in a great variety of habitats, from wet forest to very dry deciduous forest and from sea level to 2,000 m elevation.
</paragraph>
</subSubSection>
<subSubSection pageId="4" pageNumber="113" type="biology">
<paragraph pageId="4" pageNumber="113">Biology.</paragraph>
<paragraph pageId="4" pageNumber="113">
Heredia and Robbins (in prep.) summarize the food plant records for the genus, almost all of which belong to the plant family
<taxonomicName family="Loranthaceae" lsidName="" pageId="4" pageNumber="113" rank="family">Loranthaceae</taxonomicName>
(mistletoe). As noted in this paper, males of some species set up mating territories in the morning before 09:30 hours while others set up territories in the early afternoon.
</paragraph>
</subSubSection>
</treatment>
</document>