174 lines
21 KiB
XML
174 lines
21 KiB
XML
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<mods:title id="567092D25F1C6B3D4E49F11F7807C0BF">COMPARATIVE ANATOMY OF THE INSECT TRACHEAL SYSTEM PART 1: INTRODUCTION, APTERYGOTES, PALEOPTERA, POLYNEOPTERA</mods:title>
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<mods:namePart id="F7EBE00487AF859F4B2E2F4788A304C7">Herhold, Hollister W</mods:namePart>
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<mods:namePart id="A2ED20361199CD33B66E0E2E53993CAF">Degrey, Samuel P</mods:namePart>
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<treatment id="038D8781FFCA2077FEE8FA64A431FBDB" ID-DOI="http://doi.org/10.5281/zenodo.7733218" ID-GBIF-Taxon="209443381" ID-Zenodo-Dep="7733218" LSID="urn:lsid:plazi:treatment:038D8781FFCA2077FEE8FA64A431FBDB" httpUri="http://treatment.plazi.org/id/038D8781FFCA2077FEE8FA64A431FBDB" lastPageId="51" pageId="46">
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<subSubSection id="C33E651CFFCA206AFEE8FA64A1C1FF22" box="[275,469,1531,1555]" pageId="46" type="nomenclature">
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<paragraph id="8B9B3697FFCA206AFEE8FA64A1C1FF22" blockId="46.[275,469,1531,1555]" box="[275,469,1531,1555]" pageId="46">
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<heading id="D0D381FBFFCA206AFEE8FA64A1C1FF22" box="[275,469,1531,1555]" centered="true" fontSize="10" level="2" pageId="46" reason="4">
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ORDER
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<taxonomicName id="4C244D14FFCA206AFE93FA64A1C1FF22" ID-CoL="3P7" authorityName="Several D Tillyard" authorityYear="1914" box="[360,469,1531,1555]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="46" phylum="Arthropoda" rank="order">ODONATA</taxonomicName>
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</heading>
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</paragraph>
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</subSubSection>
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<subSubSection id="C33E651CFFCA2077FF77F9B3A431FBDB" lastPageId="51" pageId="46" type="description">
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<paragraph id="8B9B3697FFCA206AFF77F9B3A3AEFD64" blockId="46.[108,636,1580,1737]" lastBlockId="46.[684,1212,226,1738]" pageId="46">
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Respiration in
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<taxonomicName id="4C244D14FFCA206AFEC6F9B3A1B4FF75" authorityName="Several D Tillyard" authorityYear="1914" box="[317,416,1580,1604]" class="Magnoliopsida" higherTaxonomySource="GBIF" kingdom="Plantae" order="Odonata" pageId="46" phylum="Tracheophyta" rank="order">Odonata</taxonomicName>
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, especially aquatic naiad immatures, has been the subject of investigation for some time, although much of the morphological work was performed more than a century ago.
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<bibRefCitation id="EFB54B66FFCA206AFEFBF92EA19DFFF9" author="Scott, G. G." box="[256,393,1712,1737]" pageId="46" pagination="341 - 354" refId="ref57639" refString="Scott, G. G. 1905. The distribution of tracheae in the nymph of Plathemis lydia. Biological Bulletin 9 (6): 341 - 354." type="journal article" year="1905">Scott (1905)</bibRefCitation>
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presented one of the earliest detailed studies on dragonfly tracheae using a libellulid immature, highlighting notable characteristics such as the crossover of longitudinal trunks leading from the thorax to the abdomen, a trait retained in the adult.
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<bibRefCitation id="EFB54B66FFCA206AFB9DFEF9A2E7F8AE" author="Tillyard, R. J." pageId="46" refId="ref58551" refString="Tillyard, R. J. 1917. The biology of dragonflies (Odonata or Paraneuroptera). London: Cambridge University Press." type="book" year="1917">Tillyard (1917)</bibRefCitation>
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remains the standard for dragonfly morphology, and he carefully diagrammed the tracheal architecture via dissection, although also with an emphasis on immatures.
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<bibRefCitation id="EFB54B66FFCA206AFBB5FE75A2E1FB12" author="Chapman, R. N." pageId="46" pagination="27 - 51" refId="ref53960" refString="Chapman, R. N. 1918. The basal connections of the tracheae of the wings of insects. In The Wings of insects: 27 - 51. Ithaca, NY: Cornell University Press." type="book chapter" year="1918">Chapman (1918)</bibRefCitation>
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included both
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<taxonomicName id="4C244D14FFCA206AFC54FD93A3FEFB12" box="[943,1002,524,547]" class="Insecta" family="Aeshnidae" genus="Anax" kingdom="Animalia" order="Odonata" pageId="46" phylum="Arthropoda" rank="genus">
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<emphasis id="B950EA85FFCA206AFC54FD93A3FEFB12" box="[943,1002,524,547]" italics="true" pageId="46">Anax</emphasis>
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</taxonomicName>
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and
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<taxonomicName id="4C244D14FFCA206AFBD7FD93A478FB12" box="[1068,1132,524,547]" class="Insecta" family="Lestidae" genus="Lestes" kingdom="Animalia" order="Odonata" pageId="46" phylum="Arthropoda" rank="genus">
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<emphasis id="B950EA85FFCA206AFBD7FD93A478FB12" box="[1068,1132,524,547]" italics="true" pageId="46">Lestes</emphasis>
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</taxonomicName>
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in his comparative study of leg-wing tracheal morphology.
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<bibRefCitation id="EFB54B66FFCA206AFD1AFDD2A380FB54" author="Kennedy, C. H." box="[737,916,589,613]" pageId="46" pagination="84 - 90" refId="ref55753" refString="Kennedy, C. H. 1922. The homologies of the tracheal branches in the respiratory system of insects. Ohio Journal of Science 22 (3): 84 - 90." type="journal article" year="1922">Kennedy (1922)</bibRefCitation>
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built on Chapman’s work, extending it to abdominal tracheae; his extension to multiple orders insects using only odonate specimens is shown here to be insufficient. Recent studies have investigated physiological aspects, including active tracheal compression (likely related to respiration) in
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<taxonomicName id="4C244D14FFCA206AFBA2FC8BA4A8FA1D" authorityName="Several D Tillyard" authorityYear="1914" box="[1113,1212,788,812]" class="Magnoliopsida" higherTaxonomySource="GBIF" kingdom="Plantae" order="Odonata" pageId="46" phylum="Tracheophyta" rank="order">Odonata</taxonomicName>
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observed through synchrotron imaging by
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<bibRefCitation id="EFB54B66FFCA206AFB86FCAAA37DFA5F" author="Westneat, M. W." pageId="46" pagination="558 - 560" refId="ref59240" refString="Westneat, M. W., et al. 2003. Tracheal respiration in insects visualized with synchrotron X-ray imaging. Science 299: 558 - 560." type="journal article" year="2003">Westneat et al. (2003)</bibRefCitation>
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. For this study, three odonates were scanned, an aeshnid dragonfly larva at 30 µm resolution, an adult aeshnid at 21 µm resolution, and a calopterygid damselfly at 19 µm. As noted above, the wings of the adult aeshnid were removed and scanned separately; wing scans are not included here, though wing-base tracheae are retained and labeled.
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</paragraph>
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<paragraph id="8B9B3697FFCA206AFD37FBC1A39BFCF0" blockId="46.[684,1212,226,1738]" pageId="46">
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<bibRefCitation id="EFB54B66FFCA206AFD37FBC1A364FD46" author="Tillyard, R. J." box="[716,880,1118,1143]" pageId="46" refId="ref58551" refString="Tillyard, R. J. 1917. The biology of dragonflies (Odonata or Paraneuroptera). London: Cambridge University Press." type="book" year="1917">Tillyard (1917)</bibRefCitation>
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identified three pairs of longitudinal trunks in his figure 75 (included here as
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<figureCitation id="131F2A12FFCA206AFD57FB3EA2E2FD88" box="[684,758,1185,1209]" captionStart="FIGURE 28" captionStartId="51.[107,195,1581,1602]" captionTargetBox="[147,596,224,1553]" captionTargetId="figure-10@51.[147,597,224,1554]" captionTargetPageId="51" captionText="FIGURE 28. Tillyard’s (1917) figure 75, showing three longitudinal trunks: dorsal trunk DT, visceral trunk VT, and ventral trunk VNT." figureDoi="http://doi.org/10.5281/zenodo.7730225" httpUri="https://zenodo.org/record/7730225/files/figure.png" pageId="46">fig. 28</figureCitation>
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), whereas the aeshnid and calopterygid specimens here possess four, a condition not seen in any other order. A
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<emphasis id="B950EA85FFCA206AFC3DFB7CA3C0FDCB" box="[966,980,1251,1274]" italics="true" pageId="46">n</emphasis>
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-DLT is comprised of elongated and compartmentalized air-saclike tracheae connected segmentally or perhaps intermittently by regular tubular tracheae, and these were likely misidentified by Tillyard as air sacs rather than tracheae, which were discernible as such with micro-CT.
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</paragraph>
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<paragraph id="8B9B3697FFCA2077FD37FA55A39FF80D" blockId="46.[684,1212,226,1738]" lastBlockId="51.[684,1212,226,746]" lastPageId="51" pageId="46">
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<bibRefCitation id="EFB54B66FFCA206AFD37FA55A340FCD3" author="Scott, G. G." box="[716,852,1482,1506]" pageId="46" pagination="341 - 354" refId="ref57639" refString="Scott, G. G. 1905. The distribution of tracheae in the nymph of Plathemis lydia. Biological Bulletin 9 (6): 341 - 354." type="journal article" year="1905">Scott (1905)</bibRefCitation>
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indicated several tracheae present in the mid and hind legs of immatures but absent in adults, also noted by
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<bibRefCitation id="EFB54B66FFCA206AFC06F993A4ACFF15" author="Chapman, R. N." box="[1021,1208,1548,1572]" pageId="46" pagination="27 - 51" refId="ref53960" refString="Chapman, R. N. 1918. The basal connections of the tracheae of the wings of insects. In The Wings of insects: 27 - 51. Ithaca, NY: Cornell University Press." type="book chapter" year="1918">Chapman (1918)</bibRefCitation>
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, although with his own terminology. It is possible that these extra tracheae develop into the air-sac tracheae, which seem to be largely absent overall in immatures (otherwise they would have difficulty swimming underwater). Future studies should investigate this proliferation of tracheae and their possible development into the numerous air-sac tracheae.
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</paragraph>
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<caption id="DF5B661FFFCB206BFF90F94DA3DFFFD9" ID-DOI="http://doi.org/10.5281/zenodo.7730215" ID-Zenodo-Dep="7730215" box="[107,971,1744,1769]" httpUri="https://zenodo.org/record/7730215/files/figure.png" pageId="47" startId="47.[107,193,1746,1767]" targetBox="[308,1009,225,1710]" targetPageId="47">
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<paragraph id="8B9B3697FFCB206BFF90F94DA3DFFFD9" blockId="47.[107,971,1744,1769]" box="[107,971,1744,1769]" pageId="47">
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FIGURE 24.
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<taxonomicName id="4C244D14FFCB206BFF16F94FA1DEFFD9" box="[237,458,1744,1768]" class="Insecta" family="Baetidae" genus="Neocloeon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ephemeroptera" pageId="47" phylum="Arthropoda" rank="species" species="triangulifer">
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<emphasis id="B950EA85FFCB206BFF16F94FA1DEFFD9" box="[237,458,1744,1768]" italics="true" pageId="47">Neocloeon triangulifer</emphasis>
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</taxonomicName>
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(
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<taxonomicName id="4C244D14FFCB206BFE2CF94FA26CFFD9" authority="Hyatt & Arms, 1891" authorityName="Hyatt & Arms" authorityYear="1891" box="[471,632,1744,1768]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ephemeroptera" pageId="47" phylum="Arthropoda" rank="order">Ephemeroptera</taxonomicName>
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:
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<taxonomicName id="4C244D14FFCB206BFD79F94FA2CFFFD9" authorityName="Leach" authorityYear="1815" box="[642,731,1744,1768]" class="Insecta" family="Baetidae" kingdom="Animalia" order="Ephemeroptera" pageId="47" phylum="Arthropoda" rank="family">Baetidae</taxonomicName>
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) subimago lateral view.
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</paragraph>
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</caption>
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<caption id="DF5B661FFFD42074FF97F906A4A9FF9E" ID-DOI="http://doi.org/10.5281/zenodo.7730217" ID-Zenodo-Dep="7730217" box="[108,1213,1687,1711]" httpUri="https://zenodo.org/record/7730217/files/figure.png" pageId="48" startId="48.[108,194,1689,1710]" targetBox="[216,1104,234,1663]" targetPageId="48">
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<paragraph id="8B9B3697FFD42074FF97F906A4A9FF9E" blockId="48.[108,1213,1687,1711]" box="[108,1213,1687,1711]" pageId="48">
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FIGURE 25.
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<taxonomicName id="4C244D14FFD42074FF10F908A1D1FF9E" box="[235,453,1687,1711]" class="Insecta" family="Baetidae" genus="Neocloeon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ephemeroptera" pageId="48" phylum="Arthropoda" rank="species" species="triangulifer">
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<emphasis id="B950EA85FFD42074FF10F908A1D1FF9E" box="[235,453,1687,1711]" italics="true" pageId="48">Neocloeon triangulifer</emphasis>
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</taxonomicName>
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(
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<taxonomicName id="4C244D14FFD42074FE2AF908A265FF9E" authorityName="Several L D Landa" authorityYear="1948" box="[465,625,1687,1711]" class="Magnoliopsida" higherTaxonomySource="GBIF" kingdom="Plantae" order="Ephemeroptera" pageId="48" phylum="Tracheophyta" rank="order">Ephemeroptera</taxonomicName>
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:
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<taxonomicName id="4C244D14FFD42074FD81F908A2C6FF9E" authorityName="Leach" authorityYear="1815" box="[634,722,1687,1711]" class="Insecta" family="Baetidae" kingdom="Animalia" order="Ephemeroptera" pageId="48" phylum="Arthropoda" rank="family">Baetidae</taxonomicName>
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) subimago dorsal (left) and ventral (right) views.
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</paragraph>
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</caption>
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<caption id="DF5B661FFFD52075FF96F936A38BFF8E" ID-DOI="http://doi.org/10.5281/zenodo.7730219" ID-Zenodo-Dep="7730219" box="[109,927,1703,1727]" httpUri="https://zenodo.org/record/7730219/files/figure.png" pageId="49" startId="49.[109,195,1705,1726]" targetBox="[311,1010,226,1677]" targetPageId="49">
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<paragraph id="8B9B3697FFD52075FF96F936A38BFF8E" blockId="49.[109,927,1703,1727]" box="[109,927,1703,1727]" pageId="49">
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FIGURE 26.
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<taxonomicName id="4C244D14FFD52075FF0BF938A1D8FF8E" box="[240,460,1703,1727]" class="Insecta" family="Baetidae" genus="Neocloeon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ephemeroptera" pageId="49" phylum="Arthropoda" rank="species" species="triangulifer">
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<emphasis id="B950EA85FFD52075FF0BF938A1D8FF8E" box="[240,460,1703,1727]" italics="true" pageId="49">Neocloeon triangulifer</emphasis>
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</taxonomicName>
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(
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<taxonomicName id="4C244D14FFD52075FE21F938A26FFF8E" authority="Hyatt & Arms, 1891" authorityName="Hyatt & Arms" authorityYear="1891" box="[474,635,1703,1727]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ephemeroptera" pageId="49" phylum="Arthropoda" rank="order">Ephemeroptera</taxonomicName>
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:
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<taxonomicName id="4C244D14FFD52075FD7EF938A2CAFF8E" authorityName="Leach" authorityYear="1815" box="[645,734,1703,1727]" class="Insecta" family="Baetidae" kingdom="Animalia" order="Ephemeroptera" pageId="49" phylum="Arthropoda" rank="family">Baetidae</taxonomicName>
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) adult lateral view.
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</paragraph>
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</caption>
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<paragraph id="8B9B3697FFD62076FF97F913A4B1FF93" blockId="50.[108,1189,1674,1699]" box="[108,1189,1674,1699]" pageId="50">
|
||
FIGURE 27.
|
||
<taxonomicName id="4C244D14FFD62076FF14F915A1DFFF93" box="[239,459,1674,1698]" class="Insecta" family="Baetidae" genus="Neocloeon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ephemeroptera" pageId="50" phylum="Arthropoda" rank="species" species="triangulifer">
|
||
<emphasis id="B950EA85FFD62076FF14F915A1DFFF93" box="[239,459,1674,1698]" italics="true" pageId="50">Neocloeon triangulifer</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<taxonomicName id="4C244D14FFD62076FE23F915A26DFF93" authority="Hyatt & Arms, 1891" authorityName="Hyatt & Arms" authorityYear="1891" box="[472,633,1674,1698]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ephemeroptera" pageId="50" phylum="Arthropoda" rank="order">Ephemeroptera</taxonomicName>
|
||
:
|
||
<taxonomicName id="4C244D14FFD62076FD78F915A2C8FF93" authorityName="Leach" authorityYear="1815" box="[643,732,1674,1698]" class="Insecta" family="Baetidae" kingdom="Animalia" order="Ephemeroptera" pageId="50" phylum="Arthropoda" rank="family">Baetidae</taxonomicName>
|
||
) adult dorsal (left) and ventral (right) views.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="8B9B3697FFD72077FD37FEDAA431FBDB" blockId="51.[684,1212,226,746]" pageId="51">
|
||
The thorax of both specimens features a large number of air spaces. The flight musculature of odonates is thought to grow substantially during maturation of the adult (
|
||
<bibRefCitation id="EFB54B66FFD72077FBF4FE37A4A3F8F1" author="Marden, J. H." box="[1039,1207,424,448]" pageId="51" pagination="505 - 521" refId="ref56423" refString="Marden, J. H. 1989. Bodybuilding dragonflies: costs and benefits of maximizing flight muscle. Physiological Zoology 62 (2): 505 - 521." type="journal article" year="1989">Marden, 1989</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFB54B66FFD72077FD57FE56A394F8D0" author="Anholt, B. R. & J. H. Marden & D. M. Jenkins" box="[684,896,457,481]" pageId="51" pagination="1156 - 1163" refId="ref53507" refString="Anholt, B. R., J. H. Marden, and D. M. Jenkins. 1991. Patterns of mass gain and sexual dimorphism in adult dragonflies (Insecta: Odonata). Canadian Journal of Zoology 69 (5): 1156 - 1163." type="journal article" year="1991">Anholt et al., 1991</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFB54B66FFD72077FC76FE56A43AF8D0" author="Marden, J. H." box="[909,1070,457,481]" pageId="51" pagination="157 - 178" refId="ref56453" refString="Marden, J. H. 2000. Variability in the size, composition, and function of insect flight muscles. Annual Review of Physiology 62: 157 - 178." type="journal article" year="2000">Marden, 2000</bibRefCitation>
|
||
). It could be that these air spaces allow expansion of flight muscle—as the adult exoskeleton is fixed in size, empty volume must be allocated in advance to allow for an increase in flight muscle mass. Interestingly, this would be the inverse of the “gas bag in the gut” phenomenon (see Discussion), where the alimentary canal is coopted into a large air space in the adult.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |