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<mods:title id="9E0137F5076DD240E4D8D2880DA9AE45">The first cave associated genus of Berothidae (Insecta: Neuroptera), and a new interpretation of the subfamily Cyrenoberothinae</mods:title>
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<mods:namePart id="EB40047BD13A9BF07A5B25AA82ADF302">Machado, Renato Jose Pires</mods:namePart>
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<taxonomicName id="4C02DECA9C6AFFCBFEBC39E4FE7A06A7" authority="Machado &amp; Martins &amp; Aspöck &amp; Tavares &amp; Aspöck, 2022" authorityName="Machado &amp; Martins &amp; Aspöck &amp; Tavares &amp; Aspöck" authorityYear="2022" box="[305,510,402,426]" class="Insecta" family="Berothidae" genus="Speleoberotha" kingdom="Animalia" order="Neuroptera" pageId="4" pageNumber="1426" phylum="Arthropoda" rank="genus" status="gen. nov.">SPELEOBEROTHA</taxonomicName>
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<taxonomicNameLabel id="A245C4209C6AFFCBFD8839E3FDF806A4" box="[517,636,402,426]" pageId="4" pageNumber="1426" rank="genus">GEN. NOV.</taxonomicNameLabel>
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<emphasis id="B976795B9C6AFFCBFF2E39B3FE3F06D5" box="[163,443,453,475]" italics="true" pageId="4" pageNumber="1426">Zoobank registration:</emphasis>
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urn:lsid:zoobank.org:act:
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.
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<paragraph id="8BBDA5499C6AFFCBFF2E3A55FD710578" blockId="4.[163,780,546,630]" pageId="4" pageNumber="1426">
<emphasis id="B976795B9C6AFFCBFF2E3A55FE2D0536" box="[163,425,546,568]" italics="true" pageId="4" pageNumber="1426">E t y m o l o g y: S p e l e o</emphasis>
(f r o m G r e e k σ Π ηλαίων, spelaion = cave) and
<taxonomicName id="4C02DECA9C6AFFCBFE073A37FE600558" authorityName="Walker" authorityYear="1860" box="[394,484,577,598]" class="Insecta" family="Berothidae" genus="Berotha" kingdom="Animalia" order="Neuroptera" pageId="4" pageNumber="1426" phylum="Arthropoda" rank="genus">
<emphasis id="B976795B9C6AFFCBFE073A37FE600558" box="[394,484,577,598]" italics="true" pageId="4" pageNumber="1426">Berotha</emphasis>
</taxonomicName>
,
<typeStatus id="54B91BEB9C6AFFCBFE623A34FDA40559" box="[495,544,578,599]" pageId="4" pageNumber="1426">type</typeStatus>
genus of
<taxonomicName id="4C02DECA9C6AFFCBFD0A3A37FC820559" authorityName="Handlirsch" authorityYear="1908" box="[647,774,577,599]" class="Insecta" family="Berothidae" kingdom="Animalia" order="Neuroptera" pageId="4" pageNumber="1426" phylum="Arthropoda" rank="family">Berothidae</taxonomicName>
, a common suffix for genera in this group of animals.
</paragraph>
<paragraph id="8BBDA5499C6AFFCBFF2E3AE9FD1B05BA" blockId="4.[163,671,670,692]" box="[163,671,670,692]" pageId="4" pageNumber="1426">
<emphasis id="B976795B9C6AFFCBFF2E3AE9FDC105BD" box="[163,581,670,692]" italics="true" pageId="4" pageNumber="1426">
<typeStatus id="54B91BEB9C6AFFCBFF2E3AE9FF5C05BA" box="[163,216,671,692]" pageId="4" pageNumber="1426">Type</typeStatus>
species:
<taxonomicName id="4C02DECA9C6AFFCBFEB23AE8FDC105BD" authority="Machado &amp; Martins &amp; Aspöck &amp; Tavares &amp; Aspöck, 2022" authorityName="Machado &amp; Martins &amp; Aspöck &amp; Tavares &amp; Aspöck" authorityYear="2022" box="[319,581,670,691]" class="Insecta" family="Berothidae" genus="Speleoberotha" kingdom="Animalia" order="Neuroptera" pageId="4" pageNumber="1426" phylum="Arthropoda" rank="species" species="palomae" status="sp. nov.">Speleoberotha palomae</taxonomicName>
</emphasis>
<taxonomicNameLabel id="A245C4209C6AFFCBFDC63AE9FD1B05BA" box="[587,671,671,692]" pageId="4" pageNumber="1426" rank="species">sp. nov.</taxonomicNameLabel>
</paragraph>
<paragraph id="8BBDA5499C6AFFCBFF2E3AABFEC7043E" blockId="4.[163,779,733,816]" pageId="4" pageNumber="1426">
<emphasis id="B976795B9C6AFFCBFF2E3AABFEF105FC" box="[163,373,733,754]" italics="true" pageId="4" pageNumber="1426">Autapomorphies:</emphasis>
Antennae longer than body; Sc and RA not fused apically; male gonocoxites 9 as an unpaired bow.
</paragraph>
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<emphasis id="B976795B9C6AFFCBFF2E3B1FFEAB048E" box="[163,303,873,896]" italics="true" pageId="4" pageNumber="1426">Description</emphasis>
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<paragraph id="8BBDA5499C6AFFCBFF2E3BE7FDA4029C" blockId="4.[163,779,913,1426]" pageId="4" pageNumber="1426">Head with frons elongated; antennae moniliform and longer than body length. Pronotum wider than long, with one transversal furrow. Legs cursorial. Wings with membrane mostly hyaline but with dark markings surrounding the major forks and crossveins; margins beaded; Sc and RA not fused apically; RP with two or three major forks. Forewing with humeral recurrent vein; subcostal veinlets forked. Hindwing with CuP extremely reduced. Abdomen with ninth tergite separated from the ectoproct. Male genital sclerites without bristles; gonocoxites and gonostyli 10 (mediuncus) fused and forming an elongate and acute structure; gonocoxites 11 and 9 (gonarcus and parameres, respectively) are two unpaired bows fused basally. Female genitalia with tiny curved sclerites representing gonapophyses 9; gonocoxites 8 and gonapophyses 8 absent, and spermatheca elongated and coiled.</paragraph>
<paragraph id="8BBDA5499C6AFFCBFF2E3DB3FE9402D3" blockId="4.[163,272,1477,1501]" box="[163,272,1477,1501]" pageId="4" pageNumber="1426">
<emphasis id="B976795B9C6AFFCBFF2E3DB3FE9402D3" box="[163,272,1477,1501]" italics="true" pageId="4" pageNumber="1426">Remarks</emphasis>
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The two species included in the new genus seem to be cave dwelling, because both species were collected in or nearby caves and rock overhangs, probably living around the sheltered cave entrance area, not deep into it. This is the first record of any
<taxonomicName id="4C02DECA9C6AFFCBFDAD3E1EFD250170" authorityName="Handlirsch" authorityYear="1908" box="[544,673,1640,1662]" class="Insecta" family="Berothidae" kingdom="Animalia" order="Neuroptera" pageId="4" pageNumber="1426" phylum="Arthropoda" rank="family">Berothidae</taxonomicName>
living in this
<typeStatus id="54B91BEB9C6AFFCBFF593EF1FE810192" box="[212,261,1671,1692]" pageId="4" pageNumber="1426">type</typeStatus>
of habitat. In some of the dissected specimens of both species presented here, the abdomen is full of pollen (
<figureCitation id="1339B9CC9C6AFFCBFF793EB2FED801D7" box="[244,348,1732,1754]" captionStart="Figure 2" captionStartId="5.[146,225,1290,1312]" captionTargetBox="[146,1426,195,1250]" captionTargetId="figure-279@5.[146,1426,195,1250]" captionTargetPageId="5" captionText="Figure 2. Speleoberotha palomae sp. nov. (Berothidae: Cyrenoberothinae). A, B, female abdomen with pollen in its gut. C, vegetation near the collection locality of Speleoberotha palomae sp. nov. paratype in Pernambuco State, Brazil." figureDoi="http://doi.org/10.5281/zenodo.6994561" httpUri="https://zenodo.org/record/6994561/files/figure.png" pageId="4" pageNumber="1426">Fig. 2A,B</figureCitation>
), indicating an herbivorous diet. Pollen feeding has been reported before for different species of
<taxonomicName id="4C02DECA9C6AFFCBFF4F3F77FECD0019" authorityName="Handlirsch" authorityYear="1908" box="[194,329,1793,1815]" class="Insecta" family="Berothidae" kingdom="Animalia" order="Neuroptera" pageId="4" pageNumber="1426" phylum="Arthropoda" rank="family">Berothidae</taxonomicName>
, such as
<taxonomicName id="4C02DECA9C6AFFCBFE373F77FE68003B" authority="Monserrat &amp; Deretsky, 1999" authorityName="Monserrat &amp; Deretsky" authorityYear="1999" class="Insecta" family="Berothidae" genus="Berothimerobius" kingdom="Animalia" order="Neuroptera" pageId="4" pageNumber="1426" phylum="Arthropoda" rank="species" species="reticulatus">
<emphasis id="B976795B9C6AFFCBFE373F77FC8F0018" box="[442,779,1793,1814]" italics="true" pageId="4" pageNumber="1426">Berothimerobius reticulatus</emphasis>
<bibRefCitation id="EF93D8B89C6AFFCBFF2E3F56FE68003B" author="Monserrat VJ &amp; Deretsky Z" box="[163,492,1824,1846]" pageId="4" pageNumber="1426" pagination="45 - 66" refId="ref12489" refString="Monserrat VJ, Deretsky Z. 1999. New faunistical, taxonomic and systematic data on brown lacewings (Neuroptera: Hemerobiidae). Journal of Neuropterology 2: 45 - 66." type="journal article" year="1999">Monserrat &amp; Deretsky, 1999</bibRefCitation>
</taxonomicName>
,
<taxonomicName id="4C02DECA9C6AFFCBFE743F56FE5E005A" authority="Navas, 1933" authorityName="Navas" authorityYear="1933" class="Insecta" family="Berothidae" genus="Nyrma" kingdom="Animalia" order="Neuroptera" pageId="4" pageNumber="1426" phylum="Arthropoda" rank="species" species="kervillea">
<emphasis id="B976795B9C6AFFCBFE743F56FD31003B" box="[505,693,1824,1845]" italics="true" pageId="4" pageNumber="1426">Nyrma kervillea</emphasis>
Navás, 1933 (
<bibRefCitation id="EF93D8B89C6AFFCBFF7B3F49FE4A005A" author="Monserrat VJ" box="[246,462,1854,1876]" pageId="4" pageNumber="1426" pagination="173 - 207" refId="ref12459" refString="Monserrat VJ. 2006. Nuevos datos sobre algunas especies de la familia Berothidae (Insecta: Neuroptera). Heteropterus Revista de Entomologia 6: 173 - 207." type="journal article" year="2006">Monserrat, 2006</bibRefCitation>
)
</taxonomicName>
, the
<taxonomicName id="4C02DECA9C6AFFCBFDAE3F48FC8D005A" authorityName="MacLeod &amp; Adams" authorityYear="1967" box="[547,777,1854,1876]" class="Insecta" family="Berothidae" kingdom="Animalia" order="Neuroptera" pageId="4" pageNumber="1442" phylum="Arthropoda" rank="subFamily" subFamily="Cyrenoberothinae">Cyrenoberothinae</taxonomicName>
<taxonomicName id="4C02DECA9C6AFFCBFF2E3F2BFF7B007D" authorityName="Macleod &amp; Adams" authorityYear="1967" box="[163,255,1885,1907]" class="Insecta" family="Berothidae" genus="Cyrenoberotha" kingdom="Animalia" order="Neuroptera" pageId="4" pageNumber="1426" phylum="Arthropoda" rank="species" species="penai">
<emphasis id="B976795B9C6AFFCBFF2E3F2BFF7B007D" box="[163,255,1885,1907]" italics="true" pageId="4" pageNumber="1426">C. penai</emphasis>
</taxonomicName>
as reported by
<bibRefCitation id="EF93D8B89C6AFFCBFE233F2BFD57007D" author="MacLeod EG &amp; Adams PA" box="[430,723,1885,1907]" pageId="4" pageNumber="1426" pagination="237 - 265" refId="ref12293" refString="MacLeod EG, Adams PA. 1967. A review of the taxonomy and morphology of the Berothidae, with the description of a new subfamily from Chile (Neuroptera). Psyche 74: 237 - 265." type="journal article" year="1967">MacLeod &amp; Adams (1967)</bibRefCitation>
, and
<taxonomicName id="4C02DECA9C6AFFCBFCB638B3FC7007D4" authorityName="Aspock &amp; Aspock" authorityYear="1988" box="[827,1012,197,218]" class="Insecta" family="Berothidae" genus="Manselliberotha" kingdom="Animalia" order="Neuroptera" pageId="4" pageNumber="1426" phylum="Arthropoda" rank="genus">
<emphasis id="B976795B9C6AFFCBFCB638B3FC7007D4" box="[827,1012,197,218]" italics="true" pageId="4" pageNumber="1426">Manselliberotha</emphasis>
</taxonomicName>
, as verified here by the dissection of a few specimens. This feeding behaviour suggests that the adults of the new genus can fly outside the caves to feed on nearby plants from the Atlantic Forest biome (
<figureCitation id="1339B9CC9C6AFFCBFCCE3936FC16065B" box="[835,914,320,342]" captionStart="Figure 2" captionStartId="5.[146,225,1290,1312]" captionTargetBox="[146,1426,195,1250]" captionTargetId="figure-279@5.[146,1426,195,1250]" captionTargetPageId="5" captionText="Figure 2. Speleoberotha palomae sp. nov. (Berothidae: Cyrenoberothinae). A, B, female abdomen with pollen in its gut. C, vegetation near the collection locality of Speleoberotha palomae sp. nov. paratype in Pernambuco State, Brazil." figureDoi="http://doi.org/10.5281/zenodo.6994561" httpUri="https://zenodo.org/record/6994561/files/figure.png" pageId="4" pageNumber="1426">Fig. 2C</figureCitation>
) and return to the safety of the cave entrances.
</paragraph>
<paragraph id="8BBDA5499C6AFFCBFCDE3928FA1E0586" blockId="4.[827,1444,197,1016]" pageId="4" pageNumber="1426">
The unpaired male gonocoxites 9 seems to be an autapomorphy of
<taxonomicName id="4C02DECA9C6AFFCBFB94390BFB41069C" authorityName="Machado &amp; Martins &amp; Aspöck &amp; Tavares &amp; Aspöck" authorityYear="2022" box="[1049,1221,381,402]" class="Insecta" family="Berothidae" genus="Speleoberotha" kingdom="Animalia" order="Neuroptera" pageId="4" pageNumber="1426" phylum="Arthropoda" rank="genus">
<emphasis id="B976795B9C6AFFCBFB94390BFB41069C" box="[1049,1221,381,402]" italics="true" pageId="4" pageNumber="1426">Speleoberotha</emphasis>
</taxonomicName>
in
<taxonomicName id="4C02DECA9C6AFFCBFB79390BFA1A069D" baseAuthorityName="Leach" baseAuthorityYear="1815" box="[1268,1438,381,403]" pageId="4" pageNumber="1443" rank="superFamily" superFamily="Mantispoidea">Mantispoidea</taxonomicName>
. Among the superfamily, this particular genital piece is generally constituted by a pair of rods that are associated basally with the gonocoxites 11 (gonarcus) (
<bibRefCitation id="EF93D8B89C6AFFCBFAB839AFFC540503" author="Aspock U &amp; Aspock H" pageId="4" pageNumber="1426" pagination="97 - 127" refId="ref11704" refString="Aspock U, Aspock H. 2008. Phylogenetic relevance of the genital sclerites of Neuropterida (Insecta: Holometabola). Systematic Entomology 33: 97 - 127." type="journal article" year="2008">Aspöck &amp; Aspöck, 2008</bibRefCitation>
) and are generally important for species determination, particularly in
<taxonomicName id="4C02DECA9C6AFFCBFB163A60FBC50544" authority="(ArdilaCamacho et al., 2021)" baseAuthorityName="ArdilaCamacho" baseAuthorityYear="2021" class="Insecta" family="Mantispidae" kingdom="Animalia" order="Neuroptera" pageId="4" pageNumber="1426" phylum="Arthropoda" rank="subFamily" subFamily="Symphrasinae">
Symphrasinae (ArdilaCamacho
<emphasis id="B976795B9C6AFFCBFC3C3A40FC690544" box="[945,1005,565,587]" italics="true" pageId="4" pageNumber="1426">et al.</emphasis>
, 2021)
</taxonomicName>
. In the new genus, these two rods are fused, forming a bow similar to gonocoxite 11, representing a unique characteristic of
<taxonomicName id="4C02DECA9C6AFFCBFB7B3A04FA170589" authorityName="Machado &amp; Martins &amp; Aspöck &amp; Tavares &amp; Aspöck" authorityYear="2022" box="[1270,1427,626,647]" class="Insecta" family="Berothidae" genus="Speleoberotha" kingdom="Animalia" order="Neuroptera" pageId="4" pageNumber="1426" phylum="Arthropoda" rank="genus">
<emphasis id="B976795B9C6AFFCBFB7B3A04FA170589" box="[1270,1427,626,647]" italics="true" pageId="4" pageNumber="1426">Speleoberotha</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="8BBDA5499C6AFFCBFCDE3AE7FA9604F6" blockId="4.[827,1444,197,1016]" pageId="4" pageNumber="1426">
The long and acute structure in the male terminalia (formerly mediuncus) is interpreted here as the fusion of the unpaired gonocoxites and gonostyli 10. This interpretation follows
<bibRefCitation id="EF93D8B89C6AFFCBFBC83A9BFA18040C" author="Ardila-Camacho A &amp; Martins CC &amp; Aspock U &amp; Ramos A" box="[1093,1436,749,771]" pageId="4" pageNumber="1426" pagination="1 - 89" refId="ref11599" refString="Ardila-Camacho A, Martins CC, Aspock U, Contreras- Ramos A. 2021. Comparative morphology of extant raptorial Mantispoidea (Neuroptera: Mantispidae, Rhachiberothidae) suggests a non-monophyletic Mantispidae and a single origin of the raptorial condition within the superfamily. Zootaxa 4992: 1 - 89." type="journal article" year="2021">
Ardila-Camacho
<emphasis id="B976795B9C6AFFCBFA833A98FACE040C" box="[1294,1354,749,771]" italics="true" pageId="4" pageNumber="1426">et al.</emphasis>
(2021)
</bibRefCitation>
, who demonstrated that in
<taxonomicName id="4C02DECA9C6AFFCBFBE63B7AFA82042C" baseAuthorityName="Leach" baseAuthorityYear="1815" box="[1131,1286,780,802]" pageId="4" pageNumber="1443" rank="superFamily" superFamily="Mantispoidea">Mantispoidea</taxonomicName>
the structure traditionally called the mediuncus is formed by the gonocoxite and the gonostyli and that these are sometimes clearly distinguishable, as in
<taxonomicName id="4C02DECA9C6AFFCBFB703B1EFA240470" box="[1277,1440,872,894]" class="Insecta" family="Mantispidae" kingdom="Animalia" order="Neuroptera" pageId="4" pageNumber="1426" phylum="Arthropoda" rank="subFamily" subFamily="Symphrasinae">Symphrasinae</taxonomicName>
, or fused, as in
<taxonomicName id="4C02DECA9C6AFFCBFC6E3BF0FB0D0495" authorityName="MacLeod &amp; Adams" authorityYear="1967" box="[995,1161,902,923]" class="Insecta" family="Berothidae" genus="Cyrenoberotha" kingdom="Animalia" order="Neuroptera" pageId="4" pageNumber="1426" phylum="Arthropoda" rank="genus">
<emphasis id="B976795B9C6AFFCBFC6E3BF0FB0D0495" box="[995,1161,902,923]" italics="true" pageId="4" pageNumber="1426">Cyrenoberotha</emphasis>
</taxonomicName>
and
<taxonomicName id="4C02DECA9C6AFFCBFB4E3BF0FAF80495" authorityName="Aspock &amp; Aspock" authorityYear="1988" box="[1219,1404,902,923]" class="Insecta" family="Berothidae" genus="Manselliberotha" kingdom="Animalia" order="Neuroptera" pageId="4" pageNumber="1426" phylum="Arthropoda" rank="genus">
<emphasis id="B976795B9C6AFFCBFB4E3BF0FAF80495" box="[1219,1404,902,923]" italics="true" pageId="4" pageNumber="1426">Manselliberotha</emphasis>
</taxonomicName>
. In
<taxonomicName id="4C02DECA9C6AFFCBFCB63BD3FC5E04B4" authorityName="Machado &amp; Martins &amp; Aspöck &amp; Tavares &amp; Aspöck" authorityYear="2022" box="[827,986,933,954]" class="Insecta" family="Berothidae" genus="Speleoberotha" kingdom="Animalia" order="Neuroptera" pageId="4" pageNumber="1426" phylum="Arthropoda" rank="genus">
<emphasis id="B976795B9C6AFFCBFCB63BD3FC5E04B4" box="[827,986,933,954]" italics="true" pageId="4" pageNumber="1426">Speleoberotha</emphasis>
</taxonomicName>
, this elongated structure shows a small central hollow that is considered here as the fusion point between the gonocoxites and the gonostyli.
</paragraph>
<paragraph id="8BBDA5499C6AFFCBFCB63C57FC600358" blockId="4.[827,1442,1057,1110]" pageId="4" pageNumber="1426">
<emphasis id="B976795B9C6AFFCBFCB63C57FBD00338" box="[827,1108,1057,1079]" italics="true" pageId="4" pageNumber="1426">
Distribution (
<figureCitation id="1339B9CC9C6AFFCBFC5B3C54FBC10338" box="[982,1093,1057,1079]" captionStart="Figure 3" captionStartId="6.[164,242,1509,1531]" captionTargetBox="[167,1447,198,1466]" captionTargetId="figure-149@6.[164,1450,195,1469]" captionTargetPageId="6" captionText="Figure 3. Distribution of Cyrenoberothinae (Neuroptera: Berothidae). A, world map showing the distribution of all genera included here in Cyrenoberothinae. B, map of South America showing the records of Speleoberotha. C, maps showing in detail the three Brazilian states where Speleoberotha were collected: Ceará, Minas Gerais and Pernambuco." figureDoi="http://doi.org/10.5281/zenodo.6994563" httpUri="https://zenodo.org/record/6994563/files/figure.png" pageId="4" pageNumber="1426">Fig. 3B, C</figureCitation>
):
</emphasis>
<collectingCountry id="F315E5D99C6AFFCBFBE53C57FB2A0339" box="[1128,1198,1057,1079]" name="Brazil" pageId="4" pageNumber="1426">Brazil</collectingCountry>
(
<collectingRegion id="49C66BAB9C6AFFCBFB303C57FA800338" box="[1213,1284,1057,1078]" country="Brazil" name="Ceara" pageId="4" pageNumber="1426">Ceará</collectingRegion>
,
<collectingRegion id="49C66BAB9C6AFFCBFA823C57FA1A0339" box="[1295,1438,1057,1079]" country="Brazil" name="Pernambuco" pageId="4" pageNumber="1426">Pernambuco</collectingRegion>
,
<collectingRegion id="49C66BAB9C6AFFCBFCB63C36FC5C0358" box="[827,984,1088,1110]" country="Brazil" name="Minas Gerais" pageId="4" pageNumber="1426">Minas Gerais</collectingRegion>
).
</paragraph>
</subSubSection>
</treatment>
</document>