178 lines
23 KiB
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178 lines
23 KiB
XML
<document ID-DOI="http://dx.doi.org/10.3897/zookeys.149.2345" ID-GBIF-Dataset="ce66510c-5f50-4191-9097-7166355bf4a8" ID-PMC="PMC3234406" ID-Pensoft-Pub="1313-2970-149-17" ID-PubMed="22207791" ModsDocAuthor="" ModsDocDate="2011" ModsDocID="1313-2970-149-17" ModsDocOrigin="ZooKeys 149" ModsDocTitle="Two new species of the Hagnagora anicata complex (Geometridae, Larentiinae) from Costa Rica" checkinTime="1451249591128" checkinUser="pensoft" docAuthor="Sullivan, J. Bolling" docDate="2011" docId="31C665966F4556F5C670D96FA6707EB9" docLanguage="en" docName="ZooKeys 149: 17-29" docOrigin="ZooKeys 149" docSource="http://dx.doi.org/10.3897/zookeys.149.2345" docTitle="Hagnagora unnia Sullivan, sp. n." docType="treatment" docVersion="4" lastPageNumber="24" masterDocId="EF132A7D27270911FFBF8A6E7A34FF9B" masterDocTitle="Two new species of the Hagnagora anicata complex (Geometridae, Larentiinae) from Costa Rica" masterLastPageNumber="29" masterPageNumber="17" pageNumber="21" updateTime="1668152788704" updateUser="ExternalLinkService">
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<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
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<mods:titleInfo>
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<mods:title>Two new species of the Hagnagora anicata complex (Geometridae, Larentiinae) from Costa Rica</mods:title>
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</mods:titleInfo>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Sullivan, J. Bolling</mods:namePart>
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<mods:typeOfResource>text</mods:typeOfResource>
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<mods:title>ZooKeys</mods:title>
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</mods:titleInfo>
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<mods:part>
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<mods:date>2011</mods:date>
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<mods:detail type="volume">
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<mods:number>149</mods:number>
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<mods:extent unit="page">
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<mods:start>17</mods:start>
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<mods:end>29</mods:end>
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<mods:url>http://dx.doi.org/10.3897/zookeys.149.2345</mods:url>
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<mods:classification>journal article</mods:classification>
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<mods:identifier type="DOI">http://dx.doi.org/10.3897/zookeys.149.2345</mods:identifier>
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<mods:identifier type="Pensoft-Pub">1313-2970-149-17</mods:identifier>
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<treatment ID-GBIF-Taxon="152032294" LSID="urn:lsid:zoobank.org:act:6D9F2D75-F2BA-43D3-A056-249ED36135F6" httpUri="http://treatment.plazi.org/id/31C665966F4556F5C670D96FA6707EB9" lastPageId="7" lastPageNumber="24" pageId="4" pageNumber="21">
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<subSubSection pageId="4" pageNumber="21" type="nomenclature">
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<paragraph pageId="4" pageNumber="21">
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<taxonomicName LSID="urn:lsid:zoobank.org:act:6D9F2D75-F2BA-43D3-A056-249ED36135F6" authority="Sullivan" class="Insecta" family="Geometridae" genus="Hagnagora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hagnagora unnia" order="Lepidoptera" pageId="4" pageNumber="21" phylum="Arthropoda" rank="species" species="unnia">Hagnagora unnia Sullivan</taxonomicName>
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<taxonomicNameLabel pageId="4" pageNumber="21">sp. n.</taxonomicNameLabel>
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Figs 34
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</paragraph>
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</subSubSection>
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<subSubSection lastPageId="5" lastPageNumber="22" pageId="4" pageNumber="21" type="type material">
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<paragraph pageId="4" pageNumber="21">Type material.</paragraph>
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<paragraph lastPageId="5" lastPageNumber="22" pageId="4" pageNumber="21">
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Holotype male: Costa Rica, Tapanti National Park, Cartago Province, 1275 m, 12-17 February 2005, J. Bolling Sullivan, DNA voucher no. 06-CRBS-0049 (INBio) Paratypes 13 females: Costa Rica. Same data as holotype (1 female), DNA voucher no. 0305-CRBS-0011; Alajuela Province, Volcan Poas, 2500 m, 7-8 August 2007, J. Bolling Sullivan, (6 females), DNA voucher nos. 07-CRBS-1040, 07-CRBS-1051, 07-CRBS-384, 07-CRBS-385, 07-CRBS-386, 07-CRBS-390; Cartago Province, Villa Mills, 2841 m, J. Bolling Sullivan, (6 females), DNA voucher / GenBank
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<pageBreakToken pageId="5" pageNumber="22" start="start">accession</pageBreakToken>
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nos. 10-CRBS-1845 / JF856991, 10-CRBS-545 / HM879258, 10-CRBS-548 / HM879261, 10-CRBS-550 / HM879263, 10-CRBS-553 / HM879266, 10-CRBS-554 / HM879267. Paratypes deposited in INBio, USNM, JBS.
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</paragraph>
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</subSubSection>
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<subSubSection pageId="5" pageNumber="22" type="etymology">
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<paragraph pageId="5" pageNumber="22">Etymology.</paragraph>
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<paragraph pageId="5" pageNumber="22">This species is named for Unni E. H. Fyhn, a postdoctoral student in my laboratory in the 1970s and who continued to work on the genetic control of fish hemoglobins until her untimely death from cancer.</paragraph>
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</subSubSection>
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<subSubSection pageId="5" pageNumber="22" type="diagnosis">
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<paragraph pageId="5" pageNumber="22">Diagnosis.</paragraph>
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<paragraph pageId="5" pageNumber="22">
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Males are usually smaller but otherwise are indistinguishable from those of
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<taxonomicName class="Insecta" family="Geometridae" genus="Hagnagora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hagnagora elianne" order="Lepidoptera" pageId="5" pageNumber="22" phylum="Arthropoda" rank="species" species="elianne">Hagnagora elianne</taxonomicName>
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. They may be distinguished from
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<taxonomicName class="Insecta" family="Geometridae" genus="Hagnagora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hagnagora anicata" order="Lepidoptera" pageId="5" pageNumber="22" phylum="Arthropoda" rank="species" species="anicata">Hagnagora anicata</taxonomicName>
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by the swollen distal half of the uncus (as opposed to gently tapered) and the absence of a moderately large, upcurved spine at the end of the costa. Females may be distinguished from those of
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<taxonomicName class="Insecta" family="Geometridae" genus="Hagnagora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hagnagora elianne" order="Lepidoptera" pageId="5" pageNumber="22" phylum="Arthropoda" rank="species" species="elianne">Hagnagora elianne</taxonomicName>
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by their shorter, less complex signum. The female of
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<taxonomicName class="Insecta" family="Geometridae" genus="Hagnagora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hagnagora anicata" order="Lepidoptera" pageId="5" pageNumber="22" phylum="Arthropoda" rank="species" species="anicata">Hagnagora anicata</taxonomicName>
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is undescribed.
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</paragraph>
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<paragraph pageId="5" pageNumber="22">
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Specimens from higher altitudes are larger (see
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<bibRefCitation author="Sullivan, JB" journalOrPublisher="Journal Lepidopterists' Society" pageId="8" pageNumber="25" pagination="72 - 77" title="Intraspecific body size variation in macrolepidoptera as related to altitude of capture site and seasonal generation." volume="61" year="2007">Sullivan and Miller 2007</bibRefCitation>
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) as are most
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<taxonomicName class="Insecta" family="Geometridae" genus="Hagnagora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hagnagora elianne" order="Lepidoptera" pageId="5" pageNumber="22" phylum="Arthropoda" rank="species" species="elianne">Hagnagora elianne</taxonomicName>
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. Specimens from lower altitudes are smaller as are most
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<taxonomicName class="Insecta" family="Geometridae" genus="Hagnagora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hagnagora unnia" order="Lepidoptera" pageId="5" pageNumber="22" phylum="Arthropoda" rank="species" species="unnia">Hagnagora unnia</taxonomicName>
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. Although size alone cannot always be used to distinguish the species, it is often an excellent indicator, particularly where both species occur together. The expanse of the male valves (open and flattened) is often much smaller in
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<taxonomicName class="Insecta" family="Geometridae" genus="Hagnagora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hagnagora unnia" order="Lepidoptera" pageId="5" pageNumber="22" phylum="Arthropoda" rank="species" species="unnia">Hagnagora unnia</taxonomicName>
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; however, this character is not always definitive. In females, the lengths of the ladder-like signa do not overlap for the two species and the sample size was larger than for males (only a single barcoded male of
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<taxonomicName class="Insecta" family="Geometridae" genus="Hagnagora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hagnagora unnia" order="Lepidoptera" pageId="5" pageNumber="22" phylum="Arthropoda" rank="species" species="unnia">Hagnagora unnia</taxonomicName>
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was available). Additionally, the structure of the signum in
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<taxonomicName class="Insecta" family="Geometridae" genus="Hagnagora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hagnagora elianne" order="Lepidoptera" pageId="5" pageNumber="22" phylum="Arthropoda" rank="species" species="elianne">Hagnagora elianne</taxonomicName>
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is broader and more complex.
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</paragraph>
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</subSubSection>
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<subSubSection lastPageId="6" lastPageNumber="23" pageId="5" pageNumber="22" type="description">
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<paragraph pageId="5" pageNumber="22">Description.</paragraph>
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<paragraph lastPageId="6" lastPageNumber="23" pageId="5" pageNumber="22">
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Male. Head - palps porrect, cream with black scaling on lateral and dorsal sides of segments 2 and 3. Middle segment 2-3 x as long as other segments. Eyes round. Frons scaling slightly raised near base of tongue, brown, square, bordered with cream scales. Head brown with intermingled cream scales, particularly on edges. Collar mostly brown with scattered cream scales. Scape brown laterally and dorsally, ventral cream. Antenna simple, no visible setae, orange tan ventrally with dorsal scaling dark brown, often 70 segments. Thorax and abdomen - thorax brown dorsally; tegulae brown with thin cream scaling at outer edges. Abdomen brown with distal scaling on each segment forming a thin cream ring. Ventral abdomen similar, but with extensive cream scaling on basal segments. Legs with mixture of brown and cream scales, becoming pure cream on proximal segments. Spurs on 2nd and 3rd tibiae small. Epiphysis reaching distal end of tibia. Wings - forewing (16 mm) with cream-white band from middle of costa traversing diagonally almost to anal angle. Distal color almost black. Wing color proximal to diagonal band as dark as outer third, then becoming brown to base. Wing margins with thin cream crescents. Hindwing uniformly brown with terminal cream crescents at very tips. Leading edge of costa with two yellow spots evenly spaced between base and white band. Cream band on underside of forewing similar to that upperside. Outer third of wing rust colored becoming blackish in tornal half near wing band. Wing surface basal to white band black, crossed by thin white lines. Retinaculum thumb-like, distinct, rusty. Hindwing rusty, crossed by thin cream-white lines. Medial area of wing (corresponding to white band on forewing) lighter, distinct. Veins distinct, orange scaled. Anal edge of forewing with white scaling. Female. No differences from male except antennal segments thinner, retinaculum absent, and
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<pageBreakToken pageId="6" pageNumber="23" start="start">white</pageBreakToken>
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scaling at ventral base of scape extends onto antennal shaft. Forewing length 19.33 mm (16-21mm, N=12). Male genitalia - terminal half of moderately long uncus distinctly swollen; tip slightly pointed but not hooked. Tegumen broad, sclerotized on lateral and proximal edges that fuse below base of uncus, often forming a distinct X-shaped structure. Vinculum moderately broad, rather uneven along its edges. Vinculum fused to tegument via its proximal sclerotized edge; distal edge ends in a circular flap. Anellar arms project finger-like to medial area but do not fuse. Juxta a large circular structure. Saccus with broad, almost diamond-shaped structure, but it often is folded behind valves during slide preparation. Valva moderately broad and curves upward toward apex. Costa sclerotized but edge is not well defined; ends in a small slightly upcurved spine. Valva with pronounced, swollen, medial ridge; sacular area lightly sclerotized. Medial area of valva largely unsclerotized, but with large hair brush anchored at base. Additional hairs are on medial ridge and saccular region. Aedeagus sclerotized to manica with out third unsclerotized, but with striations. Ductus emerges subbasally. Aedeagus without sclerotized ridges or projections. Vesica expanded basally into two dorsal striated
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<normalizedToken originalValue="“wings.”">"wings."</normalizedToken>
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Ventral vesica slightly striated. Vesica narrowed toward terminus with dorsal surface crinkled and striated, but separated from striations of dorsal
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<normalizedToken originalValue="“wings.”">"wings."</normalizedToken>
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Female genitalia - anal papillae slightly pointed and striated toward distal tips. Posterior apophyses
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<normalizedToken originalValue="1½">11/2</normalizedToken>
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<normalizedToken originalValue="×">x</normalizedToken>
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length of anterior apophyses; apophyses end proximally in a terminally rounded, paddle-like structure. Segment surfaces unsclerotized, but with tightly adhering scales. Ostial area poorly defined and unsclerotized. Ostial opening expanded laterally, tapered to ductus, which is moderately long and thin with a proximal collar at base. Bursa an expanded sac, stippled dorsally and with two, ventral ladder-like signa. Signa look like corrugated pipes, tapering anteriorly. Length of longest sclerotized signum is best character for distinguishing
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<taxonomicName class="Insecta" family="Geometridae" genus="Hagnagora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hagnagora elianne" order="Lepidoptera" pageId="6" pageNumber="23" phylum="Arthropoda" rank="species" species="elianne">Hagnagora elianne</taxonomicName>
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from
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<taxonomicName class="Insecta" family="Geometridae" genus="Hagnagora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hagnagora unnia" order="Lepidoptera" pageId="6" pageNumber="23" phylum="Arthropoda" rank="species" species="unnia">Hagnagora unnia</taxonomicName>
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. No distinct structures on pelt. Tympanum with ansa swollen distally for half its length then narrowed before expanding at proximal end to form a T-like structure with a swollen base.
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</paragraph>
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<caption pageId="6" pageNumber="23">
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<paragraph pageId="6" pageNumber="23">
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Figure 3.
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<taxonomicName class="Insecta" family="Geometridae" genus="Hagnagora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hagnagora unnia" order="Lepidoptera" pageId="6" pageNumber="23" phylum="Arthropoda" rank="species" species="unnia">Hagnagora unnia</taxonomicName>
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Sullivan a adult male holotype before dissection b genital capsule c aedeagus d adult female e female genitalia.
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</paragraph>
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</caption>
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<caption pageId="6" pageNumber="23">
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<paragraph pageId="6" pageNumber="23">
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Figure 4. Comparison of female genitalia of
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<taxonomicName class="Insecta" family="Geometridae" genus="Hagnagora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hagnagora elianne" order="Lepidoptera" pageId="6" pageNumber="23" phylum="Arthropoda" rank="species" species="elianne">Hagnagora elianne</taxonomicName>
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(left) and
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<taxonomicName class="Insecta" family="Geometridae" genus="Hagnagora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hagnagora unnia" order="Lepidoptera" pageId="6" pageNumber="23" phylum="Arthropoda" rank="species" species="unnia">Hagnagora unnia</taxonomicName>
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(right) illustrating differences in length and complexity of the signa.
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</paragraph>
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</caption>
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</subSubSection>
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<subSubSection lastPageId="7" lastPageNumber="24" pageId="6" pageNumber="23" type="discussion">
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<paragraph pageId="6" pageNumber="23">Discussion.</paragraph>
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<paragraph lastPageId="7" lastPageNumber="24" pageId="6" pageNumber="23">
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The
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<taxonomicName class="Insecta" family="Geometridae" genus="Hagnagora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hagnagora anicata" order="Lepidoptera" pageId="6" pageNumber="23" phylum="Arthropoda" rank="species" species="anicata">Hagnagora anicata</taxonomicName>
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complex in Costa Rica illustrates a frequently occurring example in barcode work with neotropical
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<taxonomicName class="Magnoliopsida" family="Malvaceae" genus="Lepidoptera" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="Lepidoptera" order="Malvales" pageId="6" pageNumber="23" phylum="Tracheophyta" rank="genus">Lepidoptera</taxonomicName>
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. Species with large geographical distributions are frequently a complex of species, often seemingly identical in maculation. When genitalic examination is applied to distinct barcode clusters, they usually resolve these genetic clusters, reinforcing the concept of distinct species. Additionally, other characteristics such as elevational or geographical distributions, food plant usage, and behavior, often also support the distinctness of these clusters. In the case of the
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<taxonomicName class="Insecta" family="Geometridae" genus="Hagnagora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hagnagora anicata" order="Lepidoptera" pageId="6" pageNumber="23" phylum="Arthropoda" rank="species" species="anicata">Hagnagora anicata</taxonomicName>
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complex in Costa Rica, initial barcoding revealed one species above 3000 m and another below 2200 m, the species differing in more than 5% of their sequences and with little intraspecific sequence variability. When additional sampling was done at intermediate altitudes the results still supported two species. Although additional sampling needs to be done,
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<taxonomicName class="Insecta" family="Geometridae" genus="Hagnagora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hagnagora elianne" order="Lepidoptera" pageId="6" pageNumber="23" phylum="Arthropoda" rank="species" species="elianne">Hagnagora elianne</taxonomicName>
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continues to occur at higher altitudes and
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<taxonomicName class="Insecta" family="Geometridae" genus="Hagnagora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hagnagora unnia" order="Lepidoptera" pageId="6" pageNumber="23" phylum="Arthropoda" rank="species" species="unnia">Hagnagora unnia</taxonomicName>
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occurs at lower altitudes with the two species overlapping between 1400 and 3000 m. Based on genital characters, both species are distinct from each other and from
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<taxonomicName class="Insecta" family="Geometridae" genus="Hagnagora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hagnagora anicata" order="Lepidoptera" pageId="6" pageNumber="23" phylum="Arthropoda" rank="species" species="anicata">Hagnagora anicata</taxonomicName>
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, which may not extend north of Colombia. Material
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<pageBreakToken pageId="7" pageNumber="24" start="start">from</pageBreakToken>
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western Colombia (Valle province) differs in genital characters from
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<taxonomicName class="Insecta" family="Geometridae" genus="Hagnagora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hagnagora anicata" order="Lepidoptera" pageId="7" pageNumber="24" phylum="Arthropoda" rank="species" species="anicata">Hagnagora anicata</taxonomicName>
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(Sullivan, unpubl. data) and barcoded samples from Ecuador (Brehm, unpubl. data) reveal at least two species there, both genetically distinct from the Costa Rican species. Barcoding of hundreds of species of the montane geometrid fauna from Ecuador and Costa Rica demonstrates that very few species are actually common to both regions (Brehm and Sullivan, unpubl. data).
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</paragraph>
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<paragraph pageId="7" pageNumber="24">
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We know little about the biology of either new species, although, based on observations at 3300 m, presumed
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<taxonomicName class="Insecta" family="Geometridae" genus="Hagnagora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hagnagora elianne" order="Lepidoptera" pageId="7" pageNumber="24" phylum="Arthropoda" rank="species" species="elianne">Hagnagora elianne</taxonomicName>
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do fly diurnally, which may be true of all
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<taxonomicName class="Insecta" family="Geometridae" genus="Hagnagora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hagnagora" order="Lepidoptera" pageId="7" pageNumber="24" phylum="Arthropoda" rank="genus">Hagnagora</taxonomicName>
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species (
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<bibRefCitation author="Brehm, G" journalOrPublisher="Zeitschrift" pageId="7" pageNumber="24" pagination="256 - 260" title="Unusual flight activity of a new species of Hagnagora Druce, 1885 (Lepidoptera: Geometridae) from Costa Rica. Ent." volume="115" year="2005">Brehm and Sullivan 2005</bibRefCitation>
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). Known food plants used by
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<taxonomicName class="Insecta" family="Geometridae" genus="Hagnagora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hagnagora" order="Lepidoptera" pageId="7" pageNumber="24" phylum="Arthropoda" rank="genus">Hagnagora</taxonomicName>
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are in the
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<taxonomicName genus="Boraginaceae" lsidName="Boraginaceae" pageId="7" pageNumber="24" rank="genus">Boraginaceae</taxonomicName>
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(e.g.,
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<taxonomicName class="Insecta" family="Aphrophoridae" genus="Cordia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cordia" order="Hemiptera" pageId="7" pageNumber="24" phylum="Arthropoda" rank="genus">Cordia</taxonomicName>
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L.) and the
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<taxonomicName genus="Clethraceae" lsidName="Clethraceae" pageId="7" pageNumber="24" rank="genus">Clethraceae</taxonomicName>
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(e.g.,
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<taxonomicName class="Magnoliopsida" family="Clethraceae" genus="Clethra" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Clethra" order="Ericales" pageId="7" pageNumber="24" phylum="Tracheophyta" rank="genus">Clethra</taxonomicName>
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L.) (
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<bibRefCitation pageId="7" pageNumber="24">Brehm 2002</bibRefCitation>
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) and the
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<taxonomicName class="Insecta" family="Geometridae" genus="Hagnagora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hagnagora anicata" order="Lepidoptera" pageId="7" pageNumber="24" phylum="Arthropoda" rank="species" species="anicata">Hagnagora anicata</taxonomicName>
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complex likely uses the same groups of plants. Recently,
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<bibRefCitation pageId="7" pageNumber="24">King and Parra (2011)</bibRefCitation>
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reported captive larvae of
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<taxonomicName class="Insecta" family="Geometridae" genus="Hagnagora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hagnagora vittata" order="Lepidoptera" pageId="7" pageNumber="24" phylum="Arthropoda" rank="species" species="vittata">Hagnagora vittata</taxonomicName>
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(Philippi) that were reared on
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<taxonomicName genus="Fuschia" lsidName="Fuschia magellanica" pageId="7" pageNumber="24" rank="species" species="magellanica">Fuschia magellanica</taxonomicName>
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Lamarck (
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<taxonomicName genus="Onagraceae" lsidName="Onagraceae" pageId="7" pageNumber="24" rank="genus">Onagraceae</taxonomicName>
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).
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</paragraph>
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<paragraph pageId="7" pageNumber="24">
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The
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<taxonomicName class="Insecta" family="Geometridae" genus="Hagnagora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hagnagora anicata" order="Lepidoptera" pageId="7" pageNumber="24" phylum="Arthropoda" rank="species" species="anicata">Hagnagora anicata</taxonomicName>
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complex extends from Mexico to Bolivia and east into western Venezuela paralleling the Andes. A population occurs on Jamaica. While traditionally considered a single species, barcode and genitalic analyses indicate that five or more species are probably involved in the complex. It is not unreasonable to assume that because such complexes are common in the neotropics, the estimated number of geometrid species described from there will likely continue to increase, perhaps dramatically.
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</paragraph>
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</subSubSection>
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</treatment>
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</document> |