treatments-xml/data/02/5F/45/025F4552FF999517FF31FDC7FC37A1B9.xml

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<document ID-DOI="http://dx.doi.org/10.11646/zootaxa.3716.2.6" ID-GBIF-Dataset="bf4ff1aa-467d-41e2-a66a-141f5c442865" ID-ISSN="1175-5326" ID-Zenodo-Dep="248083" ID-ZooBank="1D536BCD-4D3B-42B0-98B9-48D8BC31ECE5" checkinTime="1460681022629" checkinUser="plazi" docAuthor="Wedmann, Sonja, Makarkin, Vladimir N., Weiterschan, Thomas &amp; Hörnschemeyer, Thomas" docDate="2013" docId="025F4552FF999517FF31FDC7FC37A1B9" docLanguage="en" docName="zt03716p258.pdf" docOrigin="Zootaxa 3716 (2)" docStyle="DocumentStylede.uka.ipd.idaho.easyIO.settings.Settings@412b93d0" docStyleName="zootaxa.2013.journal_article" docTitle="Berothidae" docType="treatment" docVersion="8" lastPageNumber="250" masterDocId="FE663D2AFF909519FFA6FF8FFFBFA356" masterDocTitle="First fossil larvae of Berothidae (Neuroptera) from Baltic amber, with notes on the biology and termitophily of the family" masterLastPageNumber="258" masterPageNumber="236" pageNumber="245" updateTime="1636695248482" updateUser="ExternalLinkService">
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<mods:title>First fossil larvae of Berothidae (Neuroptera) from Baltic amber, with notes on the biology and termitophily of the family</mods:title>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Wedmann, Sonja</mods:namePart>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Makarkin, Vladimir N.</mods:namePart>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Weiterschan, Thomas</mods:namePart>
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<mods:namePart>Hörnschemeyer, Thomas</mods:namePart>
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<mods:title>Zootaxa</mods:title>
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<mods:date>2013</mods:date>
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<mods:number>3716</mods:number>
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<treatment ID-DOI="http://doi.org/10.5281/zenodo.5677416" ID-GBIF-Taxon="119579572" ID-Zenodo-Dep="5677416" LSID="urn:lsid:plazi:treatment:025F4552FF999517FF31FDC7FC37A1B9" httpUri="http://treatment.plazi.org/id/025F4552FF999517FF31FDC7FC37A1B9" lastPageId="14" lastPageNumber="250" pageId="9" pageNumber="245">
<subSubSection pageId="9" pageNumber="245" type="nomenclature">
<paragraph blockId="9.[151,475,584,643]" box="[151,475,584,610]" pageId="9" pageNumber="245">
<heading bold="true" box="[151,475,584,610]" fontSize="11" level="1" pageId="9" pageNumber="245" reason="1">
<emphasis bold="true" box="[151,475,584,610]" pageId="9" pageNumber="245">
<taxonomicName box="[151,289,584,610]" class="Insecta" family="Berothidae" kingdom="Animalia" order="Neuroptera" pageId="9" pageNumber="245" phylum="Arthropoda" rank="family">Berothidae</taxonomicName>
indet., larva D
</emphasis>
</heading>
</paragraph>
<paragraph blockId="9.[151,475,584,643]" box="[151,270,618,643]" pageId="9" pageNumber="245">
<figureCitation box="[151,270,618,643]" captionStart-0="FIGURE 8" captionStart-1="FIGURE 9" captionStart-2="FIGURE 10" captionStartId-0="10.[151,249,1540,1562]" captionStartId-1="11.[151,249,1278,1300]" captionStartId-2="12.[151,249,1893,1915]" captionTargetBox-0="[151,1436,215,1501]" captionTargetBox-1="[165,1411,207,1228]" captionTargetBox-2="[180,1415,193,1873]" captionTargetId-0="figure@10.[151,1436,193,1518]" captionTargetId-1="figure@11.[151,1427,185,1271]" captionTargetId-2="figure@12.[180,1416,193,1875]" captionTargetPageId-0="10" captionTargetPageId-1="11" captionTargetPageId-2="12" captionText-0="FIGURE 8. Berothidae indet., larva D, photographs of the larva as preserved. A, right side. B, left side. Scale bar is 1 mm." captionText-1="FIGURE 9. Berothidae indet., larva D, drawings of the larva. A, whole larva as preserved, dorsolateral view. B, head and thorax, ventrolateral view. C, head fragment showing stemmata and right antenna, dorsolateral view. an, antenna; cd, cardo; cs, coronal suture; fm, femur; fs, frontal suture; ds, prothoracic dorsal sclerites; lp 1, first segment of labial palpus; md, mandible; mx, maxillary blade; prm, prementum; S 4, S 9, head capsule setae; st, stipes; stm, stemmata; tb, tibia; tr, trochanter. Scale bars are 1 mm (A), 0.5 mm (B), 0.1 mm (C)." captionText-2="FIGURE 10. Berothidae indet., larva D, images reconstructed from synchrotron radiation computer tomography data (false color). A, B, head and prothorax. C, whole larva, partially cut open, showing esophageal tissue in the head and an artificial cavity in the body. ac, artificial cavity; et, esophageal tissue; lcs, lateral clypeal seta; S 1, S 4, S 9, S 11, S 12, head setae. Scale bars are 0.2 mm (A, B), 1.0 mm (C)." httpUri-0="https://zenodo.org/record/248091/files/figure.png" httpUri-1="https://zenodo.org/record/248092/files/figure.png" httpUri-2="https://zenodo.org/record/248093/files/figure.png" pageId="9" pageNumber="245">
Figs.
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</figureCitation>
</paragraph>
</subSubSection>
<subSubSection pageId="9" pageNumber="245" type="materials_examined">
<paragraph blockId="9.[151,1437,690,2011]" pageId="9" pageNumber="245">
<emphasis bold="true" box="[151,387,690,715]" pageId="9" pageNumber="245">Material examined.</emphasis>
Specimen Be
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, deposited in the amber collection of SF; a third instar larva in a small piece of Baltic amber (precise collecting locality is unknown). The inclusion was bought from the collection of Marius Veta,
<collectingCountry box="[301,412,762,787]" name="Lithuania" pageId="9" pageNumber="245">Lithuania</collectingCountry>
.
</paragraph>
</subSubSection>
<subSubSection pageId="9" pageNumber="245" type="description">
<paragraph blockId="9.[151,1437,690,2011]" pageId="9" pageNumber="245">
<emphasis bold="true" box="[199,345,798,823]" pageId="9" pageNumber="245">Description.</emphasis>
Larva as preserved
<quantity box="[576,663,799,823]" metricMagnitude="-3" metricUnit="m" metricValue="3.1" pageId="9" pageNumber="245" unit="mm" value="3.1">3.1 mm</quantity>
long (from tip of cervix to abdomen); actual larval length probably about
<quantity box="[220,285,834,859]" metricMagnitude="-3" metricUnit="m" metricValue="6.0" pageId="9" pageNumber="245" unit="mm" value="6.0">6 mm</quantity>
as abdomen is distorted and posterior part with anal sucker is flapped over.
</paragraph>
<paragraph blockId="9.[151,1437,690,2011]" box="[199,1071,870,895]" pageId="9" pageNumber="245">
Head capsule elongated,
<quantity box="[479,565,870,895]" metricMagnitude="-4" metricUnit="m" metricValue="5.0" pageId="9" pageNumber="245" unit="mm" value="0.5">0.5 mm</quantity>
long,
<quantity box="[635,721,871,895]" metricMagnitude="-4" metricUnit="m" metricValue="3.0" pageId="9" pageNumber="245" unit="mm" value="0.3">0.3 mm</quantity>
wide (length/width ratio
<number box="[1005,1058,870,895]" pageId="9" pageNumber="245" value="1.67">1.67</number>
).
</paragraph>
<paragraph blockId="9.[151,1437,690,2011]" pageId="9" pageNumber="245">
Six stemmata on each side clearly visible; long setae (about
<quantity box="[918,1010,907,931]" metricMagnitude="-4" metricUnit="m" metricValue="2.0" pageId="9" pageNumber="245" unit="mm" value="0.2">0.2 mm</quantity>
) ventral to stemmata (probably S
<number box="[1407,1423,907,931]" pageId="9" pageNumber="245" value="9.0">9</number>
). Dorsally, long seta (about
<quantity box="[446,533,943,967]" metricMagnitude="-4" metricUnit="m" metricValue="2.0" pageId="9" pageNumber="245" unit="mm" value="0.2">0.2 mm</quantity>
) near stemmata (S
<number box="[743,759,943,967]" pageId="9" pageNumber="245" value="4.0">4</number>
); sockets of lateral clypeal (trichobothrian) setae on anterior part of capsule; S
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and S
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between arms of frontal suture, probably S
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just posterior to frontal suture, and some other (unidentified) setae in postero-lateral portion of head capsule clearly visible (
<figureCitation box="[1113,1194,1014,1039]" captionStart="FIGURE 9" captionStartId="11.[151,249,1278,1300]" captionTargetBox="[165,1411,207,1228]" captionTargetId="figure@11.[151,1427,185,1271]" captionTargetPageId="11" captionText="FIGURE 9. Berothidae indet., larva D, drawings of the larva. A, whole larva as preserved, dorsolateral view. B, head and thorax, ventrolateral view. C, head fragment showing stemmata and right antenna, dorsolateral view. an, antenna; cd, cardo; cs, coronal suture; fm, femur; fs, frontal suture; ds, prothoracic dorsal sclerites; lp 1, first segment of labial palpus; md, mandible; mx, maxillary blade; prm, prementum; S 4, S 9, head capsule setae; st, stipes; stm, stemmata; tb, tibia; tr, trochanter. Scale bars are 1 mm (A), 0.5 mm (B), 0.1 mm (C)." httpUri="https://zenodo.org/record/248092/files/figure.png" pageId="9" pageNumber="245">
Figs.
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C,
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A, B). Ecdysial cleavage lines very distinct; consisting of frontal and coronal sutures. Two arms of frontal suture extending for approximately half of head capsule length, convergent from inner margin of antennal tubercles until they touch. Lateral sutures absent. Coronal suture long. Ventrally, coxopodites of maxillae (stipes and cardo) rather clearly visible; stipes and cardo relatively narrow; boundary between them not distinct. Labium not clearly visible; prementum appears short (poorly preserved). Ventral setae not visible.
</paragraph>
<paragraph blockId="9.[151,1437,690,2011]" pageId="9" pageNumber="245">
Mandibles
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long, relatively broad. Maxillary blades very broad basally, gradually narrowed apically; closely associated with mandible to form mandibulomaxillary stilets, not strongly pointed apically.
</paragraph>
<paragraph blockId="9.[151,1437,690,2011]" pageId="9" pageNumber="245">
Antennal sockets rather flat, without distinct antennal tubercle. Right antenna
<quantity box="[1081,1168,1303,1327]" metricMagnitude="-4" metricUnit="m" metricValue="4.0" pageId="9" pageNumber="245" unit="mm" value="0.4">0.4 mm</quantity>
long,
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-segmented; left antenna probably
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-segmented; no apical seta visible. Segment lengths of right antenna (from base to top):
<quantity metricMagnitude="-5" metricUnit="m" metricValue="6.0" pageId="9" pageNumber="245" unit="mm" value="0.06">0.06 mm</quantity>
,
<quantity box="[210,314,1375,1399]" metricMagnitude="-5" metricUnit="m" metricValue="8.0" pageId="9" pageNumber="245" unit="mm" value="0.08">0.08 mm</quantity>
,
<quantity box="[328,432,1375,1399]" metricMagnitude="-5" metricUnit="m" metricValue="4.0" pageId="9" pageNumber="245" unit="mm" value="0.04">0.04 mm</quantity>
,
<quantity box="[445,549,1375,1399]" metricMagnitude="-5" metricUnit="m" metricValue="3.0" pageId="9" pageNumber="245" unit="mm" value="0.03">0.03 mm</quantity>
,
<quantity box="[563,667,1375,1399]" metricMagnitude="-5" metricUnit="m" metricValue="4.0" pageId="9" pageNumber="245" unit="mm" value="0.04">0.04 mm</quantity>
,
<quantity box="[681,785,1375,1399]" metricMagnitude="-5" metricUnit="m" metricValue="8.0" pageId="9" pageNumber="245" unit="mm" value="0.08">0.08 mm</quantity>
,
<quantity box="[798,903,1374,1399]" metricMagnitude="-5" metricUnit="m" metricValue="6.0" pageId="9" pageNumber="245" unit="mm" value="0.06">0.06 mm</quantity>
. All segments
<quantity box="[1074,1176,1375,1399]" metricMagnitude="-5" metricUnit="m" metricValue="2.0" pageId="9" pageNumber="245" unit="mm" value="0.02">0.02 mm</quantity>
wide, except for basal segment which is
<quantity box="[352,438,1411,1435]" metricMagnitude="-4" metricUnit="m" metricValue="3.0" pageId="9" pageNumber="245" unit="mm" value="0.3">0.3 mm</quantity>
wide. Left antenna broken twice, but probably complete.
</paragraph>
<paragraph blockId="9.[151,1437,690,2011]" pageId="9" pageNumber="245">
Labial palps poorly visible, about
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long; probably
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-segmented; second segment longest and terminal segment very short.
</paragraph>
<paragraph blockId="9.[151,1437,690,2011]" pageId="9" pageNumber="245">
Synchroton radiation computer tomography revealed remains of tissue inside of head (
<figureCitation box="[1175,1259,1518,1543]" captionStart="FIGURE 10" captionStartId="12.[151,249,1893,1915]" captionTargetBox="[180,1415,193,1873]" captionTargetId="figure@12.[180,1416,193,1875]" captionTargetPageId="12" captionText="FIGURE 10. Berothidae indet., larva D, images reconstructed from synchrotron radiation computer tomography data (false color). A, B, head and prothorax. C, whole larva, partially cut open, showing esophageal tissue in the head and an artificial cavity in the body. ac, artificial cavity; et, esophageal tissue; lcs, lateral clypeal seta; S 1, S 4, S 9, S 11, S 12, head setae. Scale bars are 0.2 mm (A, B), 1.0 mm (C)." httpUri="https://zenodo.org/record/248093/files/figure.png" pageId="9" pageNumber="245">
Fig.
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C). Oesophagus clearly visible, bifurcated towards mandibulomaxillary stilets.
</paragraph>
<paragraph blockId="9.[151,1437,690,2011]" box="[199,1310,1590,1615]" pageId="9" pageNumber="245">
Cervix probably mainly retracted into prothorax; visible length
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dorsally,
<quantity box="[1120,1206,1591,1615]" metricMagnitude="-4" metricUnit="m" metricValue="1.0" pageId="9" pageNumber="245" unit="mm" value="0.1">0.1 mm</quantity>
laterally.
</paragraph>
<paragraph blockId="9.[151,1437,690,2011]" pageId="9" pageNumber="245">
Prothorax about
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long (as preserved); with two large dorsal sclerites that touch each other at midline. Mesothorax and metathorax crumpled, not measurable; at least mesothorax with indistinctly visible paired oval dorsal sclerites.
</paragraph>
<paragraph blockId="9.[151,1437,690,2011]" pageId="9" pageNumber="245">
Legs rather short, robust. Coxa short; trochanter rather large; femur of right foreleg about
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long, stout; tibia shorter than femur; tarsus short; pretarsus with two short claws and long, basally strongly narrowed trumpetshaped empodium. Left midleg apparently missing; left hindleg stout (visible with Synchroton radiation computer tomography).
</paragraph>
<paragraph blockId="9.[151,1437,690,2011]" pageId="9" pageNumber="245">
Synchroton radiation computer tomography revealed enigmatic, rounded structure on ventro-lateral side of thorax (dorsal of left hind coxa), ca.
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wide, ca.
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long; its surface appears very smooth.
</paragraph>
<paragraph blockId="9.[151,1437,690,2011]" pageId="9" pageNumber="245">Abdomen looks strongly wrinkled; its segmentation vaguely visible. Distal portion of abdomen curved forward; probably with anal sucker.</paragraph>
</subSubSection>
<caption box="[151,1397,1540,1562]" httpUri="https://zenodo.org/record/248091/files/figure.png" pageId="10" pageNumber="246" targetBox="[151,1436,215,1501]" targetPageId="10">
<paragraph blockId="10.[151,1397,1540,1562]" box="[151,1397,1540,1562]" pageId="10" pageNumber="246">
<emphasis bold="true" box="[151,274,1540,1562]" pageId="10" pageNumber="246">FIGURE 8.</emphasis>
<taxonomicName box="[280,391,1540,1562]" class="Insecta" family="Berothidae" kingdom="Animalia" order="Neuroptera" pageId="10" pageNumber="246" phylum="Arthropoda" rank="family">Berothidae</taxonomicName>
indet., larva D, photographs of the larva as preserved. A, right side. B, left side. Scale bar is 1 mm.
</paragraph>
</caption>
<subSubSection lastPageId="14" lastPageNumber="250" pageId="10" pageNumber="246" type="discussion">
<paragraph blockId="10.[151,1436,1601,1986]" pageId="10" pageNumber="246">
<emphasis bold="true" box="[199,315,1601,1626]" pageId="10" pageNumber="246">Remarks.</emphasis>
The strongly deformed, wrinkled abdomen, the absence of one leg, and the broken antenna indicate that the larva was already dead and dry when it became embedded in the resin.
</paragraph>
<paragraph blockId="10.[151,1436,1601,1986]" pageId="10" pageNumber="246">
Fine web threads are visible around the head connecting mouthparts to the body. This specimen was probably ensnared by a spider as prey in order to catch or store it. This is corroborated by one silky thread connected to the abdomen and leading away rectangularily from it. This thread is equipped with more than
<number box="[1241,1269,1746,1770]" pageId="10" pageNumber="246" value="20.0">20</number>
tiny adhesive droplets. These are characteristic of some web spiders. Most probably there are also spider threads wrapped all around the abdomen, though these are not clearly visible. But this does not sufficiently explain its deformation.
</paragraph>
<paragraph blockId="10.[151,1436,1601,1986]" pageId="10" pageNumber="246">Apart from spider predation the larva could have been the victim of another predator or some disease before it became spider prey. Its appearance could be the result of decay.</paragraph>
<paragraph blockId="10.[151,1436,1601,1986]" pageId="10" pageNumber="246">The rounded, oval structure on the ventro-lateral side of the thorax is either a deformed part of the body, or it might be an egg, deposited by a parasite. Further identification of this structure seems not possible.</paragraph>
<caption httpUri="https://zenodo.org/record/248092/files/figure.png" pageId="11" pageNumber="247" targetBox="[165,1411,207,1228]" targetPageId="11">
<paragraph blockId="11.[151,1436,1278,1423]" pageId="11" pageNumber="247">
<emphasis bold="true" box="[151,277,1278,1300]" pageId="11" pageNumber="247">FIGURE 9.</emphasis>
<taxonomicName box="[286,397,1278,1300]" class="Insecta" family="Berothidae" kingdom="Animalia" order="Neuroptera" pageId="11" pageNumber="247" phylum="Arthropoda" rank="family">Berothidae</taxonomicName>
indet., larva D, drawings of the larva. A, whole larva as preserved, dorsolateral view. B, head and thorax, ventrolateral view. C, head fragment showing stemmata and right antenna, dorsolateral view. an, antenna; cd, cardo; cs, coronal suture; fm, femur; fs, frontal suture; ds, prothoracic dorsal sclerites; lp1, first segment of labial palpus; md, mandible; mx, maxillary blade; prm, prementum; S4, S9, head capsule setae; st, stipes; stm, stemmata; tb, tibia; tr, trochanter. Scale bars are 1 mm (A), 0.5 mm (B), 0.1 mm (C).
</paragraph>
</caption>
<paragraph blockId="11.[151,631,1498,1523]" box="[151,631,1498,1523]" pageId="11" pageNumber="247">
<emphasis bold="true" box="[151,631,1498,1523]" pageId="11" pageNumber="247">Review of knowledge on berothid larvae</emphasis>
</paragraph>
<paragraph blockId="11.[151,1436,1570,1955]" pageId="11" pageNumber="247">
Ten subfamilies are currently recognized in
<taxonomicName box="[681,809,1570,1595]" class="Insecta" family="Berothidae" kingdom="Animalia" order="Neuroptera" pageId="11" pageNumber="247" phylum="Arthropoda" rank="family">Berothidae</taxonomicName>
: the Mesozoic Mesithoninae and Paraberothinae, the Rhachiberothinae and Berothinae with fossil and extant species, and the Cyrenoberothinae, Berothimerobiinae, Nyrminae, Protobiellinae, Trichomatinae and Nosybinae which currently include only extant species (Aspöck &amp; Nemeschkal
<number box="[296,355,1679,1703]" pageId="11" pageNumber="247" value="1998.0">1998</number>
;
<collectingCountry box="[366,481,1679,1702]" name="Montserrat" pageId="11" pageNumber="247">Monserrat</collectingCountry>
<number box="[488,547,1678,1703]" pageId="11" pageNumber="247" value="2006.0">2006</number>
; Makarkin &amp; Kupryjanowicz
<number box="[881,940,1679,1703]" pageId="11" pageNumber="247" value="2010.0">2010</number>
; Makarkin
<emphasis box="[1066,1114,1678,1702]" italics="true" pageId="11" pageNumber="247">et al</emphasis>
.
<number box="[1128,1189,1679,1703]" pageId="11" pageNumber="247" value="2012.0">2012</number>
). Aspöck
<emphasis box="[1299,1347,1678,1702]" italics="true" pageId="11" pageNumber="247">et al</emphasis>
. (
<number box="[1369,1428,1679,1703]" pageId="11" pageNumber="247" value="2012.0">2012</number>
) consider the
<collectingCountry box="[295,424,1714,1739]" name="Turkey" pageId="11" pageNumber="247">Asia Minor</collectingCountry>
genus
<taxonomicName authority="Navas, 1933" authorityName="Navas" authorityYear="1933" box="[504,729,1715,1739]" class="Insecta" family="Berothidae" genus="Nyrma" kingdom="Animalia" order="Neuroptera" pageId="11" pageNumber="247" phylum="Arthropoda" rank="genus">
<emphasis box="[504,580,1715,1738]" italics="true" pageId="11" pageNumber="247">Nyrma</emphasis>
Navás,
<number box="[673,729,1715,1739]" pageId="11" pageNumber="247" value="1933.0">1933</number>
</taxonomicName>
and the Chilean
<taxonomicName authority="Blanchard, 1851" authorityName="Blanchard" authorityYear="1851" box="[921,1271,1714,1739]" class="Insecta" family="Berothidae" genus="Ormiscocerus" kingdom="Animalia" order="Neuroptera" pageId="11" pageNumber="247" phylum="Arthropoda" rank="genus">
<emphasis box="[921,1078,1715,1738]" italics="true" pageId="11" pageNumber="247">Ormiscocerus</emphasis>
Blanchard,
<number box="[1215,1271,1714,1739]" pageId="11" pageNumber="247" value="1851.0">1851</number>
</taxonomicName>
as comprising the subfamily Nyrminae. However, the body, legs and wings of
<taxonomicName box="[877,1034,1751,1774]" class="Insecta" family="Berothidae" genus="Ormiscocerus" kingdom="Animalia" order="Neuroptera" pageId="11" pageNumber="247" phylum="Arthropoda" rank="genus">
<emphasis box="[877,1034,1751,1774]" italics="true" pageId="11" pageNumber="247">Ormiscocerus</emphasis>
</taxonomicName>
are much more similar to the other Chilean genus
<taxonomicName authority="Monserrat et Deretsky, 1999 (Berothimerobiinae)" authorityName="Monserrat et Deretsky, 1999 (Berothimerobiinae" box="[316,1062,1786,1811]" class="Insecta" family="Berothidae" genus="Berothimerobius" kingdom="Animalia" order="Neuroptera" pageId="11" pageNumber="247" phylum="Arthropoda" rank="genus">
<emphasis box="[316,502,1786,1810]" italics="true" pageId="11" pageNumber="247">Berothimerobius</emphasis>
<collectingCountry box="[509,624,1787,1810]" name="Montserrat" pageId="11" pageNumber="247">Monserrat</collectingCountry>
<emphasis box="[631,651,1787,1810]" italics="true" pageId="11" pageNumber="247">et</emphasis>
Deretsky,
<number box="[771,827,1787,1811]" pageId="11" pageNumber="247" value="1999.0">1999</number>
(Berothimerobiinae)
</taxonomicName>
than to
<taxonomicName box="[1152,1228,1787,1810]" class="Insecta" family="Berothidae" genus="Nyrma" kingdom="Animalia" order="Neuroptera" pageId="11" pageNumber="247" phylum="Arthropoda" rank="genus">
<emphasis box="[1152,1228,1787,1810]" italics="true" pageId="11" pageNumber="247">Nyrma</emphasis>
</taxonomicName>
(cf. Aspöck
<number box="[1372,1431,1787,1811]" pageId="11" pageNumber="247" value="1989.0">1989</number>
;
<collectingCountry box="[151,266,1823,1846]" name="Montserrat" pageId="11" pageNumber="247">Monserrat</collectingCountry>
&amp; Deretsky
<number box="[430,489,1823,1847]" pageId="11" pageNumber="247" value="1999.0">1999</number>
; Penny &amp; Winterton
<number box="[753,814,1823,1847]" pageId="11" pageNumber="247" value="2007.0">2007</number>
).
<taxonomicName box="[839,996,1823,1846]" class="Insecta" family="Berothidae" genus="Ormiscocerus" kingdom="Animalia" order="Neuroptera" pageId="11" pageNumber="247" phylum="Arthropoda" rank="genus">
<emphasis box="[839,996,1823,1846]" italics="true" pageId="11" pageNumber="247">Ormiscocerus</emphasis>
</taxonomicName>
and
<taxonomicName box="[1064,1250,1822,1846]" class="Insecta" family="Berothidae" genus="Berothimerobius" kingdom="Animalia" order="Neuroptera" pageId="11" pageNumber="247" phylum="Arthropoda" rank="genus">
<emphasis box="[1064,1250,1822,1846]" italics="true" pageId="11" pageNumber="247">Berothimerobius</emphasis>
</taxonomicName>
might be even synonyms. Eggs of
<taxonomicName box="[381,458,1859,1882]" class="Insecta" family="Berothidae" genus="Nyrma" kingdom="Animalia" order="Neuroptera" pageId="11" pageNumber="247" phylum="Arthropoda" rank="genus">
<emphasis box="[381,458,1859,1882]" italics="true" pageId="11" pageNumber="247">Nyrma</emphasis>
</taxonomicName>
are stemmed as in most other berothids (R. Dobosz, pers. comm.), whereas eggs of
<taxonomicName box="[151,337,1894,1918]" class="Insecta" family="Berothidae" genus="Berothimerobius" kingdom="Animalia" order="Neuroptera" pageId="11" pageNumber="247" phylum="Arthropoda" rank="genus">
<emphasis box="[151,337,1894,1918]" italics="true" pageId="11" pageNumber="247">Berothimerobius</emphasis>
</taxonomicName>
are sessile (
<collectingCountry box="[488,605,1895,1919]" name="Montserrat" pageId="11" pageNumber="247">Monserrat</collectingCountry>
<number box="[618,679,1894,1919]" pageId="11" pageNumber="247" value="2006.0">2006</number>
). Therefore, both subfamilies, Nyrminae and Berothimerobiinae, appear to be valid.
</paragraph>
<caption httpUri="https://zenodo.org/record/248093/files/figure.png" pageId="12" pageNumber="248" targetBox="[180,1415,193,1873]" targetPageId="12">
<paragraph blockId="12.[151,1436,1893,2008]" pageId="12" pageNumber="248">
<emphasis bold="true" box="[151,290,1893,1915]" pageId="12" pageNumber="248">FIGURE 10.</emphasis>
<taxonomicName box="[298,409,1893,1915]" class="Insecta" family="Berothidae" kingdom="Animalia" order="Neuroptera" pageId="12" pageNumber="248" phylum="Arthropoda" rank="family">Berothidae</taxonomicName>
indet., larva D, images reconstructed from synchrotron radiation computer tomography data (false color). A, B, head and prothorax. C, whole larva, partially cut open, showing esophageal tissue in the head and an artificial cavity in the body. ac, artificial cavity; et, esophageal tissue; lcs, lateral clypeal seta; S1,S4, S9, S11, S12, head setae. Scale bars are 0.2 mm (A, B), 1.0 mm (C).
</paragraph>
</caption>
<caption httpUri="https://zenodo.org/record/248094/files/figure.png" pageId="13" pageNumber="249" targetBox="[151,1436,195,990]" targetPageId="13">
<paragraph blockId="13.[151,1436,1011,1064]" pageId="13" pageNumber="249">
<emphasis bold="true" box="[151,286,1011,1033]" pageId="13" pageNumber="249">FIGURE 11.</emphasis>
<taxonomicName box="[292,403,1011,1033]" class="Insecta" family="Berothidae" kingdom="Animalia" order="Neuroptera" pageId="13" pageNumber="249" phylum="Arthropoda" rank="family">Berothidae</taxonomicName>
indet., larva E. A, whole larva as preserved. B, anterior part of the larva (dorsal view). Scale bars are 1 mm (A, B).
</paragraph>
</caption>
<paragraph blockId="13.[151,1437,1108,2033]" pageId="13" pageNumber="249">
The larvae of few species of the following subfamilies of extant berothids are described or illustrated: Rhachiberothinae, Berothimerobiinae, Nyrminae and Berothinae (
<tableCitation box="[950,1045,1144,1169]" captionStart="TABLE 1" captionStartId="1.[151,236,152,175]" captionTargetBox="[159,1425,202,451]" captionTargetPageId="1" captionText="TABLE 1. List of extant species of Berothidae whose larvae are known." httpUri="http://table.plazi.org/id/DE89A4CCFF919518FF31FF17FC24A3F9" pageId="13" pageNumber="249" tableUuid="DE89A4CCFF919518FF31FF17FC24A3F9">
Table
<number box="[1032,1045,1145,1169]" pageId="13" pageNumber="249" value="1.0">1</number>
</tableCitation>
). The larvae of the presumed cyrenoberothine
<taxonomicName authority="Aspock et Aspock, 1988" authorityName="Aspock et Aspock" authorityYear="1988" box="[339,1028,1181,1206]" class="Insecta" family="Berothidae" genus="Manselliberotha" kingdom="Animalia" order="Neuroptera" pageId="13" pageNumber="249" phylum="Arthropoda" rank="species" species="neuropterologorum">
<emphasis box="[339,747,1181,1205]" italics="true" pageId="13" pageNumber="249">Manselliberotha neuropterologorum</emphasis>
Aspöck et Aspöck,
<number box="[972,1028,1181,1205]" pageId="13" pageNumber="249" value="1988.0">1988</number>
</taxonomicName>
are known but neither described nor illustrated (Aspöck &amp; Aspöck
<number box="[508,567,1217,1241]" pageId="13" pageNumber="249" value="1988.0">1988</number>
, p.
<number box="[613,629,1217,1241]" pageId="13" pageNumber="249" value="9.0">9</number>
). A larva from
<collectingCountry box="[817,920,1216,1241]" name="Australia" pageId="13" pageNumber="249">Australia</collectingCountry>
not attributed to a taxon is figured by Riek (
<number box="[159,220,1253,1277]" pageId="13" pageNumber="249" value="1970.0">1970</number>
). Judged from its shape and from the relatively small head, it is probably fully-grown and does not belong to the Berothinae because of the position of lateral sutures, which are located distally to the frontal suture (see Riek
<number box="[151,211,1325,1349]" pageId="13" pageNumber="249" value="1970.0">1970</number>
: Fig.
<number box="[275,338,1325,1349]" pageId="13" pageNumber="249" value="29.1">29.10</number>
G, H; New
<number box="[466,526,1325,1349]" pageId="13" pageNumber="249" value="1991.0">1991</number>
: Fig.
<number box="[589,654,1325,1349]" pageId="13" pageNumber="249" value="34.1">34.10</number>
A, B). Such a position of the lateral sutures is similar to that found in Rhachiberothinae, and it is probably plesiomorphic compared with the condition found in Berothinae, in which the lateral sutures originate from the arms of the frontal suture. Three berothid subfamilies occur in
<collectingCountry box="[1325,1432,1397,1422]" name="Australia" pageId="13" pageNumber="249">Australia</collectingCountry>
: Protobiellinae, Trichomatinae and Berothinae. The figured larva might belong to Protobiellinae, a sister group to the higher
<taxonomicName box="[289,412,1469,1494]" class="Insecta" family="Berothidae" kingdom="Animalia" order="Neuroptera" pageId="13" pageNumber="249" phylum="Arthropoda" rank="family">Berothidae</taxonomicName>
(i.e., Trichomatinae + Nosybinae + Berothinae) (Aspöck &amp; Nemeschkal
<number box="[1227,1284,1469,1493]" pageId="13" pageNumber="249" value="1998.0">1998</number>
) judged from its more primitive character states (i.e., four-segmented antennae and palpi, position of the lateral sutures).
</paragraph>
<paragraph blockId="13.[151,1437,1108,2033]" pageId="13" pageNumber="249">
The most comprehensive and detailed studies on berothid larvae can be found in the unpublished thesis of Möller (
<number box="[246,307,1577,1601]" pageId="13" pageNumber="249" value="2003.0">2003</number>
), which is provided with numerous SEM photomicrographs and drawings of the African species
<taxonomicName box="[151,366,1613,1637]" class="Insecta" family="Berothidae" genus="Podallea" kingdom="Animalia" order="Neuroptera" pageId="13" pageNumber="249" phylum="Arthropoda" rank="species" species="vasseana">
<emphasis box="[151,366,1613,1637]" italics="true" pageId="13" pageNumber="249">Podallea vasseana</emphasis>
</taxonomicName>
, and the dissertation of MacLeod (
<number box="[780,840,1613,1638]" pageId="13" pageNumber="249" value="1964.0">1964</number>
), who examined the head capsule of the American species of
<taxonomicName authority="Banks, 1905" authorityName="Banks" authorityYear="1905" box="[272,544,1648,1673]" class="Insecta" family="Berothidae" genus="Lomamyia" kingdom="Animalia" order="Neuroptera" pageId="13" pageNumber="249" phylum="Arthropoda" rank="genus">
<emphasis box="[272,390,1649,1672]" italics="true" pageId="13" pageNumber="249">Lomamyia</emphasis>
Banks,
<number box="[485,544,1648,1673]" pageId="13" pageNumber="249" value="1905.0">1905</number>
</taxonomicName>
. The majority of other descriptions are based on the first instar larvae hatched from eggs in the cage (Tillyard
<number box="[512,571,1685,1710]" pageId="13" pageNumber="249" value="1916.0">1916</number>
; Toschi
<number box="[667,726,1685,1710]" pageId="13" pageNumber="249" value="1964.0">1964</number>
; Minter
<number box="[823,882,1685,1709]" pageId="13" pageNumber="249" value="1990.0">1990</number>
;
<collectingCountry box="[895,1010,1686,1709]" name="Montserrat" pageId="13" pageNumber="249">Monserrat</collectingCountry>
<number box="[1019,1080,1685,1710]" pageId="13" pageNumber="249" value="2006.0">2006</number>
). Third instar larvae are known only for two genera of Berothinae (
<taxonomicName box="[552,653,1721,1745]" class="Insecta" family="Berothidae" genus="Podallea" kingdom="Animalia" order="Neuroptera" pageId="13" pageNumber="249" phylum="Arthropoda" rank="genus">
<emphasis box="[552,653,1721,1745]" italics="true" pageId="13" pageNumber="249">Podallea</emphasis>
</taxonomicName>
and
<taxonomicName box="[709,827,1721,1744]" class="Insecta" family="Berothidae" genus="Lomamyia" kingdom="Animalia" order="Neuroptera" pageId="13" pageNumber="249" phylum="Arthropoda" rank="genus">
<emphasis box="[709,827,1721,1744]" italics="true" pageId="13" pageNumber="249">Lomamyia</emphasis>
</taxonomicName>
) (Gurney
<number box="[945,1004,1721,1745]" pageId="13" pageNumber="249" value="1947.0">1947</number>
; Tauber &amp; Tauber
<number box="[1217,1276,1720,1745]" pageId="13" pageNumber="249" value="1968.0">1968</number>
; Möller
<number box="[1372,1431,1721,1745]" pageId="13" pageNumber="249" value="2003.0">2003</number>
; Möller
<emphasis box="[234,283,1757,1781]" italics="true" pageId="13" pageNumber="249">et al</emphasis>
.
<number box="[297,358,1757,1782]" pageId="13" pageNumber="249" value="2006.0">2006</number>
).
</paragraph>
<paragraph blockId="13.[151,1437,1108,2033]" pageId="13" pageNumber="249">
Hitherto, three fossil larvae of
<taxonomicName box="[543,666,1792,1817]" class="Insecta" family="Berothidae" kingdom="Animalia" order="Neuroptera" pageId="13" pageNumber="249" phylum="Arthropoda" rank="family">Berothidae</taxonomicName>
have been described, all from the Cretaceous: from the Valanginian/ Hauterivian of Jezzine (Jouar Es-Souss), Lebanese amber; from the earliest Cenomanian of Burmese amber (the age is after Shi
<emphasis box="[325,373,1865,1889]" italics="true" pageId="13" pageNumber="249">et al</emphasis>
.
<number box="[388,449,1865,1889]" pageId="13" pageNumber="249" value="2012.0">2012</number>
); and from the Campanian of Grassy Lake, Canadian amber.
</paragraph>
<paragraph blockId="13.[151,1437,1108,2033]" lastBlockId="14.[151,1436,151,752]" lastPageId="14" lastPageNumber="250" pageId="13" pageNumber="249">
The fossil larva described from Lebanese amber is the oldest known berothid larva. Its family affinity as belonging to
<taxonomicName box="[305,428,1936,1961]" class="Insecta" family="Berothidae" kingdom="Animalia" order="Neuroptera" pageId="13" pageNumber="249" phylum="Arthropoda" rank="family">Berothidae</taxonomicName>
is undoubted, judged from the structure of its mouthparts, ecdysial cleavage lines and antennae (see Whalley
<number box="[413,473,1973,1997]" pageId="13" pageNumber="249" value="1980.0">1980</number>
:
<number box="[484,528,1973,1998]" pageId="13" pageNumber="249" value="161.0">161</number>
,
<figureCitation box="[541,664,1973,1998]" captionStart-0="FIGURE 9" captionStart-1="FIGURE 10" captionStartId-0="11.[151,249,1278,1300]" captionStartId-1="12.[151,249,1893,1915]" captionTargetBox-0="[165,1411,207,1228]" captionTargetBox-1="[180,1415,193,1873]" captionTargetId-0="figure@11.[151,1427,185,1271]" captionTargetId-1="figure@12.[180,1416,193,1875]" captionTargetPageId-0="11" captionTargetPageId-1="12" captionText-0="FIGURE 9. Berothidae indet., larva D, drawings of the larva. A, whole larva as preserved, dorsolateral view. B, head and thorax, ventrolateral view. C, head fragment showing stemmata and right antenna, dorsolateral view. an, antenna; cd, cardo; cs, coronal suture; fm, femur; fs, frontal suture; ds, prothoracic dorsal sclerites; lp 1, first segment of labial palpus; md, mandible; mx, maxillary blade; prm, prementum; S 4, S 9, head capsule setae; st, stipes; stm, stemmata; tb, tibia; tr, trochanter. Scale bars are 1 mm (A), 0.5 mm (B), 0.1 mm (C)." captionText-1="FIGURE 10. Berothidae indet., larva D, images reconstructed from synchrotron radiation computer tomography data (false color). A, B, head and prothorax. C, whole larva, partially cut open, showing esophageal tissue in the head and an artificial cavity in the body. ac, artificial cavity; et, esophageal tissue; lcs, lateral clypeal seta; S 1, S 4, S 9, S 11, S 12, head setae. Scale bars are 0.2 mm (A, B), 1.0 mm (C)." httpUri-0="https://zenodo.org/record/248092/files/figure.png" httpUri-1="https://zenodo.org/record/248093/files/figure.png" pageId="13" pageNumber="249">
Figs.
<number box="[604,664,1973,1997]" pageId="13" pageNumber="249" value="9.5" valueMax="10.0" valueMin="9.0">910</number>
</figureCitation>
). The larva is
<quantity box="[825,919,1973,1998]" metricMagnitude="-3" metricUnit="m" metricValue="7.0" metricValueMax="8.0" metricValueMin="6.0" pageId="13" pageNumber="249" unit="mm" value="7.0" valueMax="8.0" valueMin="6.0">68 mm</quantity>
long (i.e., probably a third instar), very hairy, and possesses a trumpet-shaped empodium; rounded head; five pairs of stemmata; four- or five-segmented labial palps; antennae with an apical seta (but the number of segments is not clear); and “a prominent epicranial suture” [=ecdysial cleavage lines] (Whalley
<number box="[560,620,188,212]" pageId="14" pageNumber="250" value="1980.0">1980</number>
: p.
<number box="[658,703,188,213]" pageId="14" pageNumber="250" value="161.0">161</number>
). The structure of the latter character is very important; the photo (Whalley
<number box="[262,322,224,248]" pageId="14" pageNumber="250" value="1980.0">1980</number>
:
<figureCitation box="[333,414,223,248]" captionStart="FIGURE 10" captionStartId="12.[151,249,1893,1915]" captionTargetBox="[180,1415,193,1873]" captionTargetId="figure@12.[180,1416,193,1875]" captionTargetPageId="12" captionText="FIGURE 10. Berothidae indet., larva D, images reconstructed from synchrotron radiation computer tomography data (false color). A, B, head and prothorax. C, whole larva, partially cut open, showing esophageal tissue in the head and an artificial cavity in the body. ac, artificial cavity; et, esophageal tissue; lcs, lateral clypeal seta; S 1, S 4, S 9, S 11, S 12, head setae. Scale bars are 0.2 mm (A, B), 1.0 mm (C)." httpUri="https://zenodo.org/record/248093/files/figure.png" pageId="14" pageNumber="250">
Fig.
<number box="[385,414,224,248]" pageId="14" pageNumber="250" value="10.0">10</number>
</figureCitation>
) shows that all three sutures of the ecdysial cleavage lines (frontal, coronal and lateral) are probably present. These structures of this larva are most similar to those of the larva of the extant Chilean genus
<taxonomicName box="[151,337,296,320]" class="Insecta" family="Berothidae" genus="Berothimerobius" kingdom="Animalia" order="Neuroptera" pageId="14" pageNumber="250" phylum="Arthropoda" rank="genus">
<emphasis box="[151,337,296,320]" italics="true" pageId="14" pageNumber="250">Berothimerobius</emphasis>
</taxonomicName>
(although only a first instar larva of the latter species is known and the structure of the ecdysial cleavage lines is unknown, see
<collectingCountry box="[502,617,333,356]" name="Montserrat" pageId="14" pageNumber="250">Monserrat</collectingCountry>
<number box="[624,685,332,357]" pageId="14" pageNumber="250" value="2006.0">2006</number>
).
</paragraph>
<paragraph blockId="14.[151,1436,151,752]" pageId="14" pageNumber="250">
The larva from the Burmese amber (about
<quantity box="[717,787,368,392]" metricMagnitude="-3" metricUnit="m" metricValue="4.0" pageId="14" pageNumber="250" unit="mm" value="4.0">4 mm</quantity>
long without mouthparts) is of rather typical berothid appearance (Engel &amp; Grimaldi
<number box="[502,562,404,428]" pageId="14" pageNumber="250" value="2008.0">2008</number>
: Figs. 42, 43). It possesses the following principal character states: apparently six small stemmata; palpi with eight palpomeres, with “apical most palpomere slightly broadened, with rounded apex” (however, Fig.
<number box="[398,426,476,500]" pageId="14" pageNumber="250" value="43.0">43</number>
shows four palpomeres with short, acute terminal segment); antennae with
<number box="[1293,1321,476,501]" pageId="14" pageNumber="250" value="15.0">15</number>
“articles” (but Fig.
<number box="[259,287,512,536]" pageId="14" pageNumber="250" value="43.0">43</number>
shows the number of segments of antennae to be much less,
<number box="[996,1039,511,536]" pageId="14" pageNumber="250" value="6.0" valueMax="7.0" valueMin="5.0">57</number>
segments, with two terminal setae preserved in one antenna). The structure of the ecdysial cleavage lines is unknown.
</paragraph>
<paragraph blockId="14.[151,1436,151,752]" pageId="14" pageNumber="250">
The newly-hatched larva reported from Canadian amber as belonging to the family
<taxonomicName box="[1177,1318,583,608]" class="Insecta" family="Chrysopidae" kingdom="Animalia" order="Neuroptera" pageId="14" pageNumber="250" phylum="Arthropoda" rank="family">Chrysopidae</taxonomicName>
(Engel &amp; Grimaldi
<number box="[274,334,620,644]" pageId="14" pageNumber="250" value="2008.0">2008</number>
:
<number box="[360,391,620,644]" pageId="14" pageNumber="250" value="30.0">30</number>
, Figs.
<number box="[495,566,620,644]" pageId="14" pageNumber="250" value="13.0" valueMax="14.0" valueMin="12.0">1214</number>
) surely belongs to
<taxonomicName box="[841,967,620,645]" class="Insecta" family="Berothidae" kingdom="Animalia" order="Neuroptera" pageId="14" pageNumber="250" phylum="Arthropoda" rank="family">Berothidae</taxonomicName>
. Its size (about
<quantity box="[1203,1304,620,645]" metricMagnitude="-4" metricUnit="m" metricValue="6.0" pageId="14" pageNumber="250" unit="mm" value="0.6">0.6 mm</quantity>
excluding mandibulomaxillary stylets, too small for chrysopid larvae) and the head morphology are typical of
<taxonomicName box="[1264,1392,655,680]" class="Insecta" family="Berothidae" kingdom="Animalia" order="Neuroptera" pageId="14" pageNumber="250" phylum="Arthropoda" rank="family">Berothidae</taxonomicName>
: the mouthparts are straight, with maxillae which are broad basally and narrowed apically, not curved as in chrysopids; the antennae are
<number box="[339,351,727,752]" pageId="14" pageNumber="250" value="3.0">3</number>
-segmented, and bear a long strong terminal seta.
</paragraph>
</subSubSection>
</treatment>
</document>