257 lines
36 KiB
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257 lines
36 KiB
XML
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<mods:title id="6EA2B3CBDE823ECB8B003C982098FE5A">Lepidoptera phylogeny and systematics: the state of inventorying moth and butterfly diversity</mods:title>
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<treatment id="039C87C3FFC9DC2DFF3EE4EAFEE5FE42" ID-DOI="http://doi.org/10.5281/zenodo.5613517" ID-GBIF-Taxon="119382764" ID-Zenodo-Dep="5613517" LSID="urn:lsid:plazi:treatment:039C87C3FFC9DC2DFF3EE4EAFEE5FE42" httpUri="http://treatment.plazi.org/id/039C87C3FFC9DC2DFF3EE4EAFEE5FE42" lastPageId="9" lastPageNumber="706" pageId="4" pageNumber="703">
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<subSubSection id="C32F655EFFC9DC20FF3EE4EAFE70F85A" pageId="4" pageNumber="703" type="nomenclature">
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<paragraph id="8B8A36D5FFC9DC20FF3EE4EAFC3EFA74" blockId="4.[151,924,1450,1476]" box="[151,924,1450,1476]" pageId="4" pageNumber="703">
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<heading id="D0C281B9FFC9DC20FF3EE4EAFC3EFA74" bold="true" box="[151,924,1450,1476]" fontSize="11" level="1" pageId="4" pageNumber="703" reason="1">
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<emphasis id="B941EAC7FFC9DC20FF3EE4EAFC3EFA74" bold="true" box="[151,924,1450,1476]" pageId="4" pageNumber="703">
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<taxonomicName id="4C354D56FFC9DC20FF3EE4EAFE90FA74" ID-CoL="LP" box="[151,306,1450,1476]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="4" pageNumber="703" phylum="Arthropoda" rank="order">Lepidoptera</taxonomicName>
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phylogeny, and the inventory of
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<taxonomicName id="4C354D56FFC9DC20FD62E4EAFCBAFA74" box="[715,792,1450,1476]" class="Insecta" family="Agathiphagidae" genus="Agathiphaga" kingdom="Animalia" order="Lepidoptera" pageId="4" pageNumber="703" phylum="Arthropoda" rank="species" species="major">major</taxonomicName>
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groupings
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</emphasis>
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</heading>
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</paragraph>
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<paragraph id="8B8A36D5FFC9DC20FF3EE4ADFE70F85A" blockId="4.[151,1436,1517,2026]" pageId="4" pageNumber="703">
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Some key points in the history of lepidopteran classification were given by
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<bibRefCitation id="EFA44B24FFC9DC20FBB3E4ADFB61F9B7" author="Scoble" box="[1050,1219,1517,1543]" pageId="4" pageNumber="740" refString="Scoble, M. J. (1992) The Lepidoptera: Form, Function and Diversity. 404 pp., Oxford University Press. Scoble, M. J. (2003) Changing roles and perceptions in European natural history collections: from idiosynchracy to infrastructucture. In: Scoble, M. J. (ed.): ENHSIN: The European Natural History Specimen Information Network. The Natural History Museum, London, pp. 11 - 20." type="book chapter" year="1992">Scoble (1992)</bibRefCitation>
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, and what follows immediately below is based on that account. The most prominent early landmark in lepidopteran classification is the 10th edition of Linnaeus’
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<emphasis id="B941EAC7FFC9DC20FDA8E700FD6AF9E9" box="[513,712,1600,1625]" italics="true" pageId="4" pageNumber="703">Systema Naturae</emphasis>
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(
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<bibRefCitation id="EFA44B24FFC9DC20FD7EE700FC2BF9EA" author="Linnaeus" box="[727,905,1600,1626]" pageId="4" pageNumber="738" refString="Linnaeus, C. (1758) Systema Naturae per regna tria naturae, secundum classes, ordines genera, species, cum characteribus, differentiis, synonymis, locis. 10 th ed. Holmiae. 534 pp." type="book" year="1758">Linnaeus 1758</bibRefCitation>
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), which, other than for spiders, represents the beginning of zoological nomenclature as accepted by the
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<emphasis id="B941EAC7FFC9DC20FCFCE728FA37F931" box="[853,1429,1640,1665]" italics="true" pageId="4" pageNumber="703">International Code of Zoological Nomenclature</emphasis>
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. Although published work on lepidopteran natural history dates back to Aristotle (384–322 BC), and while later pre-Linnaean authors, notably Albertus Magnus, Marcel Malpighius, Jan Swammerdam, René Reaumur, and Charles De Geer, made significant contributions, it was Linnaeus who provided a classification, comprehensive for the time, in which we see the germ of subsequent systems.
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<bibRefCitation id="EFA44B24FFC9DC20FC7BE648FB31F892" author="Linnaeus" box="[978,1171,1800,1826]" pageId="4" pageNumber="738" refString="Linnaeus, C. (1758) Systema Naturae per regna tria naturae, secundum classes, ordines genera, species, cum characteribus, differentiis, synonymis, locis. 10 th ed. Holmiae. 534 pp." type="book" year="1758">Linnaeus (1758)</bibRefCitation>
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recognised three main divisions of the
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<taxonomicName id="4C354D56FFC9DC20FEF2E670FE4BF8FA" box="[347,489,1840,1866]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="4" pageNumber="703" phylum="Arthropoda" rank="order">Lepidoptera</taxonomicName>
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-
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<taxonomicName id="4C354D56FFC9DC20FDAFE670FDFEF8F9" box="[518,604,1840,1865]" class="Insecta" family="Papilionidae" genus="Papilio" kingdom="Animalia" order="Lepidoptera" pageId="4" pageNumber="703" phylum="Arthropoda" rank="genus">
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<emphasis id="B941EAC7FFC9DC20FDAFE670FDFEF8F9" box="[518,604,1840,1865]" italics="true" pageId="4" pageNumber="703">Papilio</emphasis>
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</taxonomicName>
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,
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<taxonomicName id="4C354D56FFC9DC20FDC4E670FD1FF8F9" box="[621,701,1840,1865]" class="Insecta" family="Sphingidae" genus="Sphinx" kingdom="Animalia" order="Lepidoptera" pageId="4" pageNumber="703" phylum="Arthropoda" rank="genus">
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<emphasis id="B941EAC7FFC9DC20FDC4E670FD1FF8F9" box="[621,701,1840,1865]" italics="true" pageId="4" pageNumber="703">Sphinx</emphasis>
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</taxonomicName>
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, and
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<taxonomicName id="4C354D56FFC9DC20FCAAE670FCD1F8F9" box="[771,883,1840,1865]" class="Insecta" family="Noctuidae" genus="Phalaena" higherTaxonomySource="CoL" kingdom="Animalia" order="Lepidoptera" pageId="4" pageNumber="703" phylum="Arthropoda" rank="genus">
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<emphasis id="B941EAC7FFC9DC20FCAAE670FCD1F8F9" box="[771,883,1840,1865]" italics="true" pageId="4" pageNumber="703">Phalaena</emphasis>
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</taxonomicName>
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.
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<taxonomicName id="4C354D56FFC9DC20FC2DE670FC56F8F9" box="[900,1012,1840,1865]" class="Insecta" family="Aleyrodidae" genus="Phaleana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hemiptera" pageId="4" pageNumber="703" phylum="Arthropoda" rank="genus">
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<emphasis id="B941EAC7FFC9DC20FC2DE670FC56F8F9" box="[900,1012,1840,1865]" italics="true" pageId="4" pageNumber="703">Phaleana</emphasis>
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</taxonomicName>
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was divided into seven subgroups. Today, all but
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<taxonomicName id="4C354D56FFC9DC20FEEDE618FE16F8C1" box="[324,436,1880,1905]" class="Insecta" family="Noctuidae" genus="Phalaena" higherTaxonomySource="CoL" kingdom="Animalia" order="Lepidoptera" pageId="4" pageNumber="703" phylum="Arthropoda" rank="genus">
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<emphasis id="B941EAC7FFC9DC20FEEDE618FE16F8C1" box="[324,436,1880,1905]" italics="true" pageId="4" pageNumber="703">Phalaena</emphasis>
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</taxonomicName>
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are still used for nine superfamilies. Following the Linnaean sequence these superfamilies are: Papilionoidea, Sphingoidea, Bombycoidea, Noctuoidea, Geometroidea, Tortricoidea, Pyraloidea, Tineoidea, and Alucitoidea. Linnaeus’ coverage was remarkable and his system, evidently, is embedded in our higher classification today.
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</paragraph>
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||
</subSubSection>
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<subSubSection id="C32F655EFFC8DC21FF6FE1D8FC3BF85D" pageId="5" pageNumber="704" type="description">
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<paragraph id="8B8A36D5FFC8DC21FF6FE1D8FF75FEE2" blockId="5.[151,1437,152,578]" pageId="5" pageNumber="704">Linnaeus based his classification on features of the antennae and the wings, particularly the position of the wings at rest, and on whether the insects flew by day or by night. He also took account of the mouthparts. He noted that the antennae of butterflies were clubbed, and that some members of what he called ‘Bombyces’ lacked a tongue. A considerable variety of features of larvae and adults is to be found in his species descriptions.</paragraph>
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<paragraph id="8B8A36D5FFC8DC21FF6FE020FC42FDF2" blockId="5.[151,1437,152,578]" pageId="5" pageNumber="704">
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Lepidopteran classification developed from the Linnaean foundation in several ways. Numerous additional species were described, as discussed elsewhere in this paper and Linnaeus’ divisions and subdivisions were further divided and formal categorical ranks were established for them. Moreover, many supraspecific taxa were added to those introduced by Linnaeus. Primary
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<typeStatus id="548E8877FFC8DC21FCEDE098FC20FE42" box="[836,898,472,498]" pageId="5" pageNumber="704">types</typeStatus>
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of many Linnean
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<taxonomicName id="4C354D56FFC8DC21FBFFE098FB46FE42" box="[1110,1252,472,498]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="5" pageNumber="704" phylum="Arthropoda" rank="order">Lepidoptera</taxonomicName>
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species survive to this day, and thus remain available for study. Linnaeus’ own collection is housed in the Linnean Society of London (see http://www.linnean.org/); examples are shown in
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<figureCitation id="130E2A50FFC8DC21FCDAE368FC7EFDF2" box="[883,988,552,578]" captionStart-0="FIGURE 2" captionStart-1="FIGURE 3" captionStartId-0="5.[151,255,1544,1568]" captionStartId-1="6.[151,255,1934,1958]" captionTargetBox-0="[150,1436,621,1504]" captionTargetBox-1="[153,1436,185,1926]" captionTargetId-0="figure@5.[149,1438,619,1506]" captionTargetPageId-0="5" captionTargetPageId-1="6" captionText-0="FIGURE 2. Lectotype of Papilio machaon Linnaeus, 1758, type species of the genus Papiio Linnaeus, 1758. “ … if this is not a butterfly in a nomenclatural sense, then nothing is ” (Vane-Wright 2007: 59). Courtesy of The Linnean Society of London." captionText-1="FIGURE 3. Two unit trays of moths from Linnaeus's collection. While these trays indeed contain ' Microlepidoptera' (typewritten labels in upper left corners) in the sense that the moths were described in Pyralis and Tinea, several of the species are now placed in the macrolepidopteran family Noctuidae. Courtesy of The Linnean Society of London." httpUri-0="https://zenodo.org/record/274046/files/figure.png" httpUri-1="https://zenodo.org/record/274047/files/figure.png" pageId="5" pageNumber="704">Figs 2–3</figureCitation>
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.
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</paragraph>
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<caption id="DF4A665DFFC8DC21FF3EE748FF53F9D5" httpUri="https://zenodo.org/record/274046/files/figure.png" pageId="5" pageNumber="704" targetBox="[150,1436,621,1504]" targetPageId="5">
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<paragraph id="8B8A36D5FFC8DC21FF3EE748FF53F9D5" blockId="5.[151,1436,1544,1637]" pageId="5" pageNumber="704">
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<emphasis id="B941EAC7FFC8DC21FF3EE748FEBBF990" bold="true" box="[151,281,1544,1568]" pageId="5" pageNumber="704">FIGURE 2.</emphasis>
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Lectotype of
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<taxonomicName id="4C354D56FFC8DC21FE06E749FCB6F9AF" authority="Linnaeus, 1758" authorityName="Linnaeus" authorityYear="1758" box="[431,788,1544,1568]" class="Insecta" family="Papilionidae" genus="Papilio" kingdom="Animalia" order="Lepidoptera" pageId="5" pageNumber="704" phylum="Arthropoda" rank="species" species="machaon">
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<emphasis id="B941EAC7FFC8DC21FE06E749FDC4F990" box="[431,614,1545,1568]" italics="true" pageId="5" pageNumber="704">Papilio machaon</emphasis>
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Linnaeus, 1758
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</taxonomicName>
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, type species of the genus
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<emphasis id="B941EAC7FFC8DC21FB9BE749FBDBF990" box="[1074,1145,1545,1568]" italics="true" pageId="5" pageNumber="704">Papiio</emphasis>
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Linnaeus, 1758. “…
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<emphasis id="B941EAC7FFC8DC21FAC9E749FD5CF9F2" italics="true" pageId="5" pageNumber="704">if this is not a butterfly in a nomenclatural sense, then nothing is</emphasis>
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” (Vane-Wright 2007: 59). Courtesy of The Linnean Society of London.
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</paragraph>
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</caption>
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<paragraph id="8B8A36D5FFC8DC21FF6FE7D3FC3BF85D" blockId="5.[151,1437,1683,2029]" pageId="5" pageNumber="704">
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Prior to Linnaeus’ death, and just under twenty years after the publication of
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<emphasis id="B941EAC7FFC8DC21FBF6E7D3FA8DF91C" box="[1119,1327,1683,1708]" italics="true" pageId="5" pageNumber="704">Systema Naturae,</emphasis>
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<bibRefCitation id="EFA44B24FFC8DC21FA91E7D3FE38F965" author="Denis" pageId="5" pageNumber="735" refString="Denis, M. & Schiffermuller, I. (1775) Ankundung eines systematischen Werkes von der Schmetterlingen der Wienergegend. 323 pp., 3. Pls. Wien." type="book" year="1775">Denis & Schiffermüller (1775)</bibRefCitation>
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produced a work on the
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<taxonomicName id="4C354D56FFC8DC21FD67E7FBFCFEF965" box="[718,860,1723,1749]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="5" pageNumber="704" phylum="Arthropoda" rank="order">Lepidoptera</taxonomicName>
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of the Vienna area. Their system was based on Linnaeus’ classification, but they added further observations on structure and examined many additional species. They subdivided the Linnaean groups and based the names of many of the subdivisions on both larva (e.g., Larvae Punctatae) and adult (e.g., Ph[alaenae] Geometrae Unicolores). Building on these foundations, the Danish entomologist Fabricius, who was a student of Linnaeus, described many new species and named some of the Linnaean subgroups (
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<bibRefCitation id="EFA44B24FFC8DC21FD8BE6C5FC6FF82C" author="Fabricius" box="[546,973,1925,1948]" pageId="5" pageNumber="736" refString="Fabricius, J. C. (1775). Systema entomologiae, sistens insectorum classes, ordines genera, species, adiectis synonymis, locis descriptionibus, observationibus. xxx + 832 pp. Flensburgi et Lipsiae." type="book" year="1775">Fabricius 1775 - and several later works</bibRefCitation>
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). Fabricius’ impact on
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<taxonomicName id="4C354D56FFC8DC21FB73E6C3FACAF82D" box="[1242,1384,1923,1949]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="5" pageNumber="704" phylum="Arthropoda" rank="order">Lepidoptera</taxonomicName>
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taxonomy (indeed insect taxonomy broadly) was notable. Slightly later,
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<bibRefCitation id="EFA44B24FFC8DC21FC6FE6EBFBDCF875" author="Latreille" box="[966,1150,1963,1989]" pageId="5" pageNumber="738" refString="Latreille, P. A. (1796) Precis des Caracteres generiques des insects, dispose dans un ordre naturel par le Citoyen Latreille. 14 + 210 + 5 pp. Brive, Bordeaux." type="book" year="1796">Latreille (1796)</bibRefCitation>
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added further supraspecific groups, most of which represent genera in the modern sense.
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</paragraph>
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</subSubSection>
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<caption id="DF4A665DFFCBDC22FF3EE6CEFAF3F85B" httpUri="https://zenodo.org/record/274047/files/figure.png" pageId="6" pageNumber="705" targetBox="[153,1436,185,1926]" targetPageId="6">
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<paragraph id="8B8A36D5FFCBDC22FF3EE6CEFAF3F85B" blockId="6.[151,1436,1934,2027]" pageId="6" pageNumber="705">
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<emphasis id="B941EAC7FFCBDC22FF3EE6CEFEB9F816" bold="true" box="[151,283,1934,1958]" pageId="6" pageNumber="705">FIGURE 3.</emphasis>
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Two unit trays of moths from Linnaeus's collection. While these trays indeed contain 'Microlepidoptera' (typewritten labels in upper left corners) in the sense that the moths were described in
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<taxonomicName id="4C354D56FFCBDC22FB97E6F1FB29F878" box="[1086,1163,1969,1992]" class="Insecta" family="Pyralidae" genus="Pyralis" kingdom="Animalia" order="Lepidoptera" pageId="6" pageNumber="705" phylum="Arthropoda" rank="genus">
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<emphasis id="B941EAC7FFCBDC22FB97E6F1FB29F878" box="[1086,1163,1969,1992]" italics="true" pageId="6" pageNumber="705">Pyralis</emphasis>
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</taxonomicName>
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and
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<taxonomicName id="4C354D56FFCBDC22FB68E6F1FB59F878" box="[1217,1275,1969,1992]" class="Insecta" family="Tineidae" genus="Tinea" kingdom="Animalia" order="Lepidoptera" pageId="6" pageNumber="705" phylum="Arthropoda" rank="genus">
|
||
<emphasis id="B941EAC7FFCBDC22FB68E6F1FB59F878" box="[1217,1275,1969,1992]" italics="true" pageId="6" pageNumber="705">Tinea</emphasis>
|
||
</taxonomicName>
|
||
, several of the species are now placed in the macrolepidopteran family
|
||
<taxonomicName id="4C354D56FFCBDC22FD59E694FCC3F85B" box="[752,865,2004,2027]" class="Insecta" family="Noctuidae" kingdom="Animalia" order="Lepidoptera" pageId="6" pageNumber="705" phylum="Arthropoda" rank="family">Noctuidae</taxonomicName>
|
||
. Courtesy of The Linnean Society of London.
|
||
</paragraph>
|
||
</caption>
|
||
<subSubSection id="C32F655EFFCADC23FF6FE1D8FCF3FDDA" pageId="7" pageNumber="706" type="reference_group">
|
||
<paragraph id="8B8A36D5FFCADC23FF6FE1D8FB76FECA" blockId="7.[151,1437,152,2018]" pageId="7" pageNumber="706">
|
||
<taxonomicName id="4C354D56FFCADC23FF6FE1D8FE8BFF02" box="[198,297,152,178]" class="Insecta" family="Agathiphagidae" genus="Agathiphaga" kingdom="Animalia" order="Lepidoptera" pageId="7" pageNumber="706" phylum="Arthropoda" rank="species" species="major">A major</taxonomicName>
|
||
step in the evolution of lepidopteran taxonomy was noted by
|
||
<bibRefCitation id="EFA44B24FFCADC23FBAFE1D8FB13FF02" author="Emmet" box="[1030,1201,152,178]" pageId="7" pageNumber="736" refString="Emmet, A. M. (1992) The scientific names of British Lepidoptera: Their history and meaning. Harley Books, Colchester, 221 pp." type="book" year="1992">Emmet (1992)</bibRefCitation>
|
||
who suggested that our modern concept of the genus in
|
||
<taxonomicName id="4C354D56FFCADC23FDE2E180FD7AFF6A" ID-CoL="LP" box="[587,728,192,218]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="7" pageNumber="706" phylum="Arthropoda" rank="order">Lepidoptera</taxonomicName>
|
||
was provided by Schrank (
|
||
<date id="FF8B1015FFCADC23FB8BE180FB2CFF6A" box="[1058,1166,192,218]" pageId="7" pageNumber="706" value="1801" valueMax="1802">1801–02</date>
|
||
), who added this level between the Linnaeus’ ‘genus’ and species. Many modern genera were established by Jacob Hübner, a designer and artist, who described and illustrated many of the genera (called
|
||
<emphasis id="B941EAC7FFCADC23FBBCE050FBC7FE99" box="[1045,1125,272,297]" italics="true" pageId="7" pageNumber="706">Stirpes</emphasis>
|
||
by him) that we recognize today (
|
||
<bibRefCitation id="EFA44B24FFCADC23FF41E078FE0AFEE2" author="Hemming" box="[232,424,312,338]" pageId="7" pageNumber="736" refString="Hemming, F. (1937) Hubner. A bibliographical and systematic account of the entomological works of Jacob Hubner and of the supplements by Carl Geyer, Gotfried Franz von Fr ¨ lich and Gottlieb August Wilhelm Herrich-Schaffer." type="book" year="1937">Hemming 1937</bibRefCitation>
|
||
). The lepidopteran genus was firmly established by
|
||
<bibRefCitation id="EFA44B24FFCADC23FBBFE078FAAFFEE2" author="Ochsenheimer" box="[1046,1293,312,338]" pageId="7" pageNumber="739" refString="Ochsenheimer, F. (1807 - 1835). Die Schmetterlinge von Europa 1 - 10. (vol. 5 - 10, 1825 - 1836 by F. Treitschke). Er. Fleischer, Leipzig." type="book chapter" year="1807">
|
||
Ochsenheimer (
|
||
<date id="FF8B1015FFCADC23FB79E078FAAFFEE2" box="[1232,1293,312,338]" pageId="7" pageNumber="706" value="1807" valueMax="1835">1807</date>
|
||
</bibRefCitation>
|
||
–1835), in a series of works completed by Treitsche, in his synthesis of the lepidopteran fauna of Europe.
|
||
</paragraph>
|
||
<paragraph id="8B8A36D5FFCADC23FF6FE0C8FCF3FDDA" blockId="7.[151,1437,152,2018]" pageId="7" pageNumber="706">
|
||
Many of the higher taxa of
|
||
<taxonomicName id="4C354D56FFCADC23FDB4E0C8FD09FE12" ID-CoL="LP" box="[541,683,392,418]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="7" pageNumber="706" phylum="Arthropoda" rank="order">Lepidoptera</taxonomicName>
|
||
that we still accept in a recognizable form emerged from the impressive and very influential six volume study by G.A.W. Herrich-Schäffer. The work was published at regular intervals between
|
||
<date id="FF8B1015FFCADC23FE60E098FDD8FE42" box="[457,634,472,498]" pageId="7" pageNumber="706" value="1843" valueMax="1856">1843 and 1856</date>
|
||
(
|
||
<bibRefCitation id="EFA44B24FFCADC23FD23E098FC3EFE42" author="Herrich-Schaffer" box="[650,924,472,498]" pageId="7" pageNumber="737" refString="Herrich-Schaffer, G. A. W. (1843 - 56) Systematische Bearbeitung der Schmetterlinge von Europa, als Text, Revision und Supplement zu J. Huebners Sammlung europaischer Schmetterlinge. Vol. 1 - 6. Manz, Regensburg." type="book chapter" year="1843">
|
||
Herrich-Schäffer,
|
||
<date id="FF8B1015FFCADC23FCC9E098FC3EFE42" box="[864,924,472,498]" pageId="7" pageNumber="706" value="1843" valueMax="1856">1843</date>
|
||
</bibRefCitation>
|
||
–1856). Although, as for earlier works, this study was based on the European fauna, the implications are applicable far more widely. To a significant extent, Herrich-Schaeffer established the taxa on wing venation. He provided numerous morphological illustrations, which are remarkable in their detail and execution.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="C32F655EFFCADC2DFF6FE338FEE5FE42" lastPageId="9" lastPageNumber="708" pageId="7" pageNumber="706" type="description">
|
||
<paragraph id="8B8A36D5FFCADC23FF6FE338FCBFFC1A" blockId="7.[151,1437,152,2018]" pageId="7" pageNumber="706">
|
||
With collections being made from beyond Europe, a further dimension to the classification was added. Apart from the fact that they described numerous new genera and species, Edward Meyrick (particularly for microlepidopterans) and Sir George Hampson (particularly for macrolepidopterans) had a marked impact on the higher classification of the order.
|
||
<bibRefCitation id="EFA44B24FFCADC23FDE0E3B0FCA0FCBA" author="Meyrick" box="[585,770,752,778]" pageId="7" pageNumber="739" refString="Meyrick, E. (1895) A handbook of British Lepidoptera. [viii] + 844 pp. London." type="book" year="1895">Meyrick (1895)</bibRefCitation>
|
||
proposed that the
|
||
<taxonomicName id="4C354D56FFCADC23FC70E3B0FBC4FCBA" box="[985,1126,752,778]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="7" pageNumber="706" phylum="Arthropoda" rank="order">Lepidoptera</taxonomicName>
|
||
should be divided into ten ‘phyla’. Meyrick, like Herrich-Schäffer, relied to a significant extent on wing venation because he considered the pattern to be of low adaptive (‘physiological’) value and likely therefore to be little altered by external factors. Hampson’s contribution to
|
||
<taxonomicName id="4C354D56FFCADC23FDBEE228FD06FC32" box="[535,676,872,898]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="7" pageNumber="706" phylum="Arthropoda" rank="order">Lepidoptera</taxonomicName>
|
||
higher classification focused mainly on Noctuoidea (see
|
||
<bibRefCitation id="EFA44B24FFCADC23FAF8E228FF5CFC1A" author="Kitching" pageId="7" pageNumber="737" refString="Kitching, I. J. (1984) A historical review of the higher classification of the Noctuidae (Lepidoptera). Bulletin of the British Museum (Natural Hististory) Entomology, 49, 153 - 234." type="journal article" year="1984">Kitching 1984</bibRefCitation>
|
||
for an analysis), Pyraloidea, and Thyridoidea.
|
||
</paragraph>
|
||
<paragraph id="8B8A36D5FFCADC23FF6FE2F8FEEAFA3A" blockId="7.[151,1437,152,2018]" pageId="7" pageNumber="706">
|
||
Since the mid nineteenth century many other systems have caused the earlier classifications to be modified and expanded. Some writers emphasized other character sets e.g., from larvae (
|
||
<bibRefCitation id="EFA44B24FFCADC23FB2BE2A0FA80FC4A" author="Fracker" box="[1154,1314,992,1018]" pageId="7" pageNumber="736" refString="Fracker, S. B. (1915) The classification of lepidopterous larvae. Illinois Biological Monographsi, 2, 1 - 169." type="journal article" year="1915">Fracker 1915</bibRefCitation>
|
||
) or pupae (
|
||
<bibRefCitation id="EFA44B24FFCADC23FF36E548FEE3FB92" author="Mosher" box="[159,321,1032,1058]" pageId="7" pageNumber="739" refString="Mosher, E. (1916) A classification of the Lepidoptera based on characters of the pupa. Bulletin of the Illinois State Laboratory of natural History, 12, 1 - 158." type="journal article" year="1916">Mosher 1916</bibRefCitation>
|
||
). Probably the
|
||
<taxonomicName id="4C354D56FFCADC23FE58E548FD94FB92" box="[497,566,1032,1058]" class="Insecta" family="Agathiphagidae" genus="Agathiphaga" kingdom="Animalia" order="Lepidoptera" pageId="7" pageNumber="706" phylum="Arthropoda" rank="species" species="major">major</taxonomicName>
|
||
contribution to the higher classification of the order concerns the division above the superfamily. Landmark works were written by
|
||
<bibRefCitation id="EFA44B24FFCADC23FCECE570FC45FBFA" author="Borner" box="[837,999,1072,1098]" pageId="7" pageNumber="734" refString="Borner, C. (1925) Lepidoptera, Schmetterlinge. In: Brohmer, P.: Fauna von Deutschland (3. edn), 358 - 387." type="journal article" year="1925">Börner (1925</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="EFA44B24FFCADC23FC5CE570FB93FBFA" author="Borner" box="[1013,1073,1072,1098]" pageId="7" pageNumber="734" refString="Borner, C. (1939) Die Grundlagen meines Lepidopterensystems. Verhandlungen des VIII. Internationalen Kongresses fur Entomologie Berlin 1938, 2, 1372 - 1424." type="journal article" year="1939">1939</bibRefCitation>
|
||
) who proposed a fundamental division of the
|
||
<taxonomicName id="4C354D56FFCADC23FEE1E518FE77FBC2" box="[328,469,1112,1138]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="7" pageNumber="706" phylum="Arthropoda" rank="order">Lepidoptera</taxonomicName>
|
||
into Monotrysia and Ditrysia on the basis of the structure of female genitalia. The Monotrysia (
|
||
<emphasis id="B941EAC7FFCADC23FE9AE5C0FED6FB29" box="[307,372,1152,1177]" italics="true" pageId="7" pageNumber="706">sensu</emphasis>
|
||
Börner), which included only 5 percent of the
|
||
<taxonomicName id="4C354D56FFCADC23FC14E5C0FBECFB2A" box="[957,1102,1152,1178]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="7" pageNumber="706" phylum="Arthropoda" rank="order">Lepidoptera</taxonomicName>
|
||
, are not monophyletic. But Börner’s recognition of the Ditrysia as a natural group was an important step in understanding the phylogenetic structure of the order. So also was his appreciation of the systematic value of many morphological characters in the classification and diagnosis of lepidopteran families and superfamilies. Building on the works of Börner as well as of Kiriakoff (1948) and in particular
|
||
<bibRefCitation id="EFA44B24FFCADC23FC99E460FC7EFA8A" author="Hinton" box="[816,988,1312,1338]" pageId="7" pageNumber="737" refString="Hinton, H. E. (1946) On the homology and nomenclature of the setae of lepidopterous larvae, with some notes on the phylogeny of the Lepidoptera. Transactions of the Royal Entomological Society of London, 97, 1 - 37" type="journal article" year="1946">Hinton (1946)</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="EFA44B24FFCADC23FC43E460FB3BFA8A" author="Hennig" box="[1002,1177,1312,1338]" pageId="7" pageNumber="736" refString="Hennig, W. (1953) Kritische Bemerkungen zum phylogenetischen System der Insekten. Beitrage zur Entomologie, 3 (Sonderheft), 1 - 85." type="journal article" year="1953">Hennig (1953)</bibRefCitation>
|
||
outlined the relationships of the most basal lineages within the order in a benchmark article, which foreshadowed subsequent work on this subject.
|
||
</paragraph>
|
||
<paragraph id="8B8A36D5FFCADC2DFF6FE4D8FEE5FE42" blockId="7.[151,1437,152,2018]" lastBlockId="9.[151,1436,152,498]" lastPageId="9" lastPageNumber="708" pageId="7" pageNumber="706">
|
||
One hundred and twenty-four lepidopteran families (52 of them with a subfamily classification) were recognized in the recent treatment of the order in the
|
||
<emphasis id="B941EAC7FFCADC23FD5CE480FC54FA69" box="[757,1014,1472,1497]" italics="true" pageId="7" pageNumber="706">Handbook of Zoology</emphasis>
|
||
(Kristensen ed. 1998, 2003). These families were grouped into 47 diagnosed superfamilies, and a provisional phylogeny at the superfamily level was presented. About the same time an alternative classification was presented by
|
||
<bibRefCitation id="EFA44B24FFCADC23FBCDE750FA83F99A" author="Heppner" box="[1124,1313,1552,1578]" pageId="7" pageNumber="736" refString="Heppner, J. B. (1998) Classification of Lepidoptera. Part 1. Introduction. Holarctic Lepidoptera, 5 (Suppl. 1): 1 - 148." type="journal article" year="1998">Heppner (1998)</bibRefCitation>
|
||
, who later (2005) expanded it to tribal level. The two classifications are broadly similar, but some differences exist, partly because Heppner’s classification expressly does not aim to reflect monophyla in the 'Hennigian' (cladistic) sense of phylogenetic systematics. Moreover, the superfamilies and higher-level entities in Heppner’s work were not diagnosed. The superfamily/family/subfamily classification adopted in the
|
||
<emphasis id="B941EAC7FFCADC23FB72E7F0FAF7F979" box="[1243,1365,1712,1737]" italics="true" pageId="7" pageNumber="706">Handbook</emphasis>
|
||
, with some
|
||
<taxonomicName id="4C354D56FFCADC23FF77E798FE80F942" box="[222,290,1752,1778]" class="Insecta" family="Agathiphagidae" genus="Agathiphaga" kingdom="Animalia" order="Lepidoptera" pageId="7" pageNumber="706" phylum="Arthropoda" rank="species" species="major">major</taxonomicName>
|
||
modifications suggested in subsequent work, is given in Appendix 1. Although the superfamily cladogram printed in the
|
||
<emphasis id="B941EAC7FFCADC23FE68E640FD99F8A9" box="[449,571,1792,1817]" italics="true" pageId="7" pageNumber="706">Handbook</emphasis>
|
||
(and see
|
||
<figureCitation id="130E2A50FFCADC23FD07E640FD51F8AA" box="[686,755,1792,1818]" captionStart="FIGURE 3" captionStartId="6.[151,255,1934,1958]" captionTargetBox="[153,1436,185,1926]" captionTargetPageId="6" captionText="FIGURE 3. Two unit trays of moths from Linnaeus's collection. While these trays indeed contain ' Microlepidoptera' (typewritten labels in upper left corners) in the sense that the moths were described in Pyralis and Tinea, several of the species are now placed in the macrolepidopteran family Noctuidae. Courtesy of The Linnean Society of London." httpUri="https://zenodo.org/record/274047/files/figure.png" pageId="7" pageNumber="706">
|
||
Fig.
|
||
<quantity id="4CCD9B30FFCADC23FD4CE640FD51F8AA" box="[741,755,1792,1818]" metricMagnitude="-2" metricUnit="m" metricValue="7.62" pageId="7" pageNumber="706" unit="in" value="3.0">3</quantity>
|
||
</figureCitation>
|
||
in the present paper) has been referred to as representing the “consensus” position (
|
||
<bibRefCitation id="EFA44B24FFCADC23FE6EE668FD66F8F2" author="Holloway" box="[455,708,1832,1858]" pageId="7" pageNumber="737" refString="Holloway, J. D., Kibby, G. & Peggie, D. (2001) The families of Malesian moths and butterflies. Fauna Malesiana 3, i - xi, 1 - 455. Brill, Leiden, Boston & Koln." type="journal article" year="2001">
|
||
Holloway
|
||
<emphasis id="B941EAC7FFCADC23FDEDE668FDD5F8F1" box="[580,631,1832,1857]" italics="true" pageId="7" pageNumber="706">et al</emphasis>
|
||
. 2001
|
||
</bibRefCitation>
|
||
), its provisional nature should be emphasized: its lack of resolution of
|
||
<taxonomicName id="4C354D56FFCADC23FEAEE610FEEEF8DA" box="[263,332,1872,1898]" class="Insecta" family="Agathiphagidae" genus="Agathiphaga" kingdom="Animalia" order="Lepidoptera" pageId="7" pageNumber="706" phylum="Arthropoda" rank="species" species="major">major</taxonomicName>
|
||
branches is conspicuous. It serves, however, as a starting point for some ongoing inquiries into
|
||
<taxonomicName id="4C354D56FFCADC23FF7DE638FEC3F822" box="[212,353,1912,1938]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="7" pageNumber="706" phylum="Arthropoda" rank="order">Lepidoptera</taxonomicName>
|
||
phylogenetic systematics (see, e.g., the http://www.tolweb.org/
|
||
<taxonomicName id="4C354D56FFCADC23FBDEE638FAA9F822" box="[1143,1291,1912,1938]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="7" pageNumber="706" phylum="Arthropoda" rank="order">Lepidoptera</taxonomicName>
|
||
and http:// www.lepsys.eu/ websites). In
|
||
<figureCitation id="130E2A50FFCADC23FDAAE6E0FDEAF80A" box="[515,584,1952,1978]" captionStart="FIGURE 4" captionStartId="8.[151,255,1915,1939]" captionTargetBox="[237,1350,199,1878]" captionTargetId="figure@8.[224,1370,151,1909]" captionTargetPageId="8" captionText="FIGURE 4. Cladogram of lepidopteran superfamilies / superfamily assemblages. Widths of clade bars pertaining to larger clades (> 1.000 species) reflect estimated numbers of described species. Scale bar = 10,000 species. From Kristensen (ed.) (2003)." httpUri="https://zenodo.org/record/274048/files/figure.png" pageId="7" pageNumber="706">Fig. 4</figureCitation>
|
||
the approximate species numbers for superfamilies/superfamily assemblages with more than 1,000 described species is indicated by the width of the clade lines. It is immediately apparent, that the most species-rich lineages are cladistically quite subordinate. This particular phylogenetic pattern was ascribed considerable general significance in Hennig's writings (e.g., 1953, 1966): indeed, the basic diversification pattern within the
|
||
<taxonomicName id="4C354D56FFC4DC2DFDF7E1A8FD4EFEB2" box="[606,748,232,258]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="9" pageNumber="708" phylum="Arthropoda" rank="order">Lepidoptera</taxonomicName>
|
||
remains an unusually instructive example of what has subsequently come to be known as a 'Hennigian comb', with the first diverged extant lineages exhibiting a step-by-step acquisition of the apomorphies which characterize the most subordinate (and successful) groups. Undoubtedly the proportional representation of individual superfamilies will change markedly as descriptive
|
||
<taxonomicName id="4C354D56FFC4DC2DFF3EE0C8FE87FE12" box="[151,293,392,418]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="9" pageNumber="708" phylum="Arthropoda" rank="order">Lepidoptera</taxonomicName>
|
||
taxonomy approaches completion. Thus only relatively modest growth is foreseeable in the butterfly and bombycoid assemblages, while particularly strong increases are expected for the Gelechioidea and the Pyraloidea.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |