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<mods:title id="1B37803645031CEA3A11CD7FEEA14AC6">Reconciliation between neontology and paleontology in the Gryllidea (Orthoptera, Ensifera): reinterpreting the venation of the stridulatory apparatus in crickets</mods:title>
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As currently defined, the †
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and †
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show a similar and relatively stable venation patterns (
<bibRefCitation id="174E4B5AFFD4C73E76F8FCD55D262306" author="PEREZ DE LA FUENTE R. &amp; HEADS S. W. &amp; HINOJOSA-DIAZ I. A. &amp; ENGEL M. S." pageId="11" pageNumber="778" pagination="53 - 58" refId="ref19126" refString="PEREZ DE LA FUENTE R., HEADS S. W., HINOJOSA-DIAZ I. A. &amp; ENGEL M. S. 2012. - The first record of Protogryllinae from the Jurassic of India (Orthoptera: Protogryllidae). Journal of the Kansas Entomological Society 85: 53 - 58. https: // doi. org / 10.2317 / JKES 111103.1" type="journal article" year="2012">
Pérez de la Fuente
<emphasis id="41ABEAB9FFD4C73E74A3FCF65D642306" box="[235,286,821,847]" italics="true" pageId="11" pageNumber="778">et al.</emphasis>
2012
</bibRefCitation>
;
<bibRefCitation id="174E4B5AFFD4C73E7523FCF65E582306" author="WANG H. &amp; FANG Y. N. &amp; FANG Y. &amp; JARZEMBOWSKI E. A. &amp; WANG B. &amp; ZHANG H." box="[363,546,821,848]" pageId="11" pageNumber="778" pagination="1440 - 1444" refId="ref20144" refString="WANG H., FANG Y. N., FANG Y., JARZEMBOWSKI E. A., WANG B. &amp; ZHANG H. 2019. - The earliest fossil record of true crickets belonging to the Baissogryllidae (Insecta, Orthoptera, Grylloidea). Geological Magazine 156: 1440 - 1444. https: // doi. org / 10.1017 / S 0016756818000754" type="journal article" year="2019">
Wang
<emphasis id="41ABEAB9FFD4C73E75F8FCF65D992306" box="[432,483,821,847]" italics="true" pageId="11" pageNumber="778">et al.</emphasis>
2019
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). The general organisation of their forewings is almost similar to that of modern crickets, with a lateral field and a dorsal field, a median fold, and a fan between R and CuA (possibly corresponding to a flexible zone, as in mole crickets).
</paragraph>
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Lateral field. In these fossils, the location of the putative C is often not preserved. The most anterior visible vein is Sc, which has the same pectinate branching as in modern
<taxonomicName id="B4DF4D28FFD4C73E76DCFBD45F7F2467" box="[660,773,1044,1070]" class="Aves" family="Trochilidae" genus="Grylloidea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Apodiformes" pageId="11" pageNumber="776" phylum="Chordata" rank="genus">Grylloidea</taxonomicName>
(
<figureCitation id="EBE42A2EFFD4C73E74C6FBF45CB12407" box="[142,203,1076,1102]" captionStart="FIG" captionStartId="13.[131,142,1941,1959]" captionTargetBox="[186,1379,218,1892]" captionTargetPageId="13" captionText="FIG. 6. — Hypothesis of primary venation homology of male forewing of †Protogryllidae (A, B) and †Baissogryllidae (C-E): A, †Angarogryllus angaricus (Sharov 1968), PIN 1873-16; B, †Falsipseculum karatavicum (Sharov 1968), PIN 3791/1345; C, †Neosharategia paradoxa Gorochov, 1992, PIN 4270-210a; D, †Baissogryllidae sp., CCNH-293;E, †Anglogryllus lyristes Gorochov et al., 2006,MNEMG 2003.46. Abbreviations:“ha”, distally opened harp; “mi”, distally opened mirror, others and colour code, see text. Grey dash lines represent folds. Scale bars: 1 mm." figureDoi="http://doi.org/10.5281/zenodo.10376762" httpUri="https://zenodo.org/record/10376762/files/figure.png" pageId="11" pageNumber="778">Fig. 6</figureCitation>
). Posteriorly, R is present and strongly convex. Then, the M is fused basally with CuA. A lanceolate cell (
<figureCitation id="EBE42A2EFFD4C73E7615FB945EE22427" box="[605,664,1108,1134]" captionStart="FIG" captionStartId="13.[131,142,1941,1959]" captionTargetBox="[186,1379,218,1892]" captionTargetPageId="13" captionText="FIG. 6. — Hypothesis of primary venation homology of male forewing of †Protogryllidae (A, B) and †Baissogryllidae (C-E): A, †Angarogryllus angaricus (Sharov 1968), PIN 1873-16; B, †Falsipseculum karatavicum (Sharov 1968), PIN 3791/1345; C, †Neosharategia paradoxa Gorochov, 1992, PIN 4270-210a; D, †Baissogryllidae sp., CCNH-293;E, †Anglogryllus lyristes Gorochov et al., 2006,MNEMG 2003.46. Abbreviations:“ha”, distally opened harp; “mi”, distally opened mirror, others and colour code, see text. Grey dash lines represent folds. Scale bars: 1 mm." figureDoi="http://doi.org/10.5281/zenodo.10376762" httpUri="https://zenodo.org/record/10376762/files/figure.png" pageId="11" pageNumber="778">Fig. 6</figureCitation>
) is present between R and M: it is basally closed by a short transverse vein (r-m) located at the level of the divergence of M with CuA, and distally closed by a curved RP that partially merges with MA (as in modern
<taxonomicName id="B4DF4D28FFD4C73E74BEFB145D1D24A7" box="[246,359,1236,1262]" class="Aves" family="Trochilidae" genus="Grylloidea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Apodiformes" pageId="11" pageNumber="776" phylum="Chordata" rank="genus">Grylloidea</taxonomicName>
, see above). Contrary to the latter, some †
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have several secondary transverse veins in the lanceolate cell, as for example †
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<emphasis id="41ABEAB9FFD4C73E758CFAD35ED72564" box="[452,685,1299,1325]" italics="true" pageId="11" pageNumber="778">Angarogryllus angaricus</emphasis>
(
<bibRefCitation id="174E4B5AFFD4C73E76F1FAD35CC62504" author="SHAROV A. G." pageId="11" pageNumber="778" pagination="1 - 216" refId="ref19687" refString="SHAROV A. G. 1968. - Filogeniya ortopteroidnykh nasekomykh. Trudy Paleontologicheskogo Instituta, Akademiya Nauk S. S. S. R., 118, 1 - 216, Moskva. [in Russian, translated in english in 1971: Phylogeny of the Orthopteroidea. Israel program for scientific translations, Keter Press, Jerusalem: 1 - 251.]" type="journal article" year="1968">Sharov, 1968</bibRefCitation>
)
</taxonomicName>
(
<figureCitation id="EBE42A2EFFD4C73E749CFAF35D502504" box="[212,298,1331,1357]" captionStart="FIG" captionStartId="13.[131,142,1941,1959]" captionTargetBox="[186,1379,218,1892]" captionTargetPageId="13" captionText="FIG. 6. — Hypothesis of primary venation homology of male forewing of †Protogryllidae (A, B) and †Baissogryllidae (C-E): A, †Angarogryllus angaricus (Sharov 1968), PIN 1873-16; B, †Falsipseculum karatavicum (Sharov 1968), PIN 3791/1345; C, †Neosharategia paradoxa Gorochov, 1992, PIN 4270-210a; D, †Baissogryllidae sp., CCNH-293;E, †Anglogryllus lyristes Gorochov et al., 2006,MNEMG 2003.46. Abbreviations:“ha”, distally opened harp; “mi”, distally opened mirror, others and colour code, see text. Grey dash lines represent folds. Scale bars: 1 mm." figureDoi="http://doi.org/10.5281/zenodo.10376762" httpUri="https://zenodo.org/record/10376762/files/figure.png" pageId="11" pageNumber="778">Fig. 6A</figureCitation>
). In others, like †
<taxonomicName id="B4DF4D28FFD4C73E75A5FAF35D652524" authority="(Sharov, 1968)" baseAuthorityName="Sharov" baseAuthorityYear="1968" class="Insecta" family="Protogryllidae" genus="Falsispeculum" kingdom="Animalia" order="Orthoptera" pageId="11" pageNumber="778" phylum="Arthropoda" rank="species" species="karatavicum">
<emphasis id="41ABEAB9FFD4C73E75A5FAF35F7D2504" box="[493,775,1331,1357]" italics="true" pageId="11" pageNumber="778">Falsispeculum karatavicum</emphasis>
(
<bibRefCitation id="174E4B5AFFD4C73E74C3FA935D6D2524" author="SHAROV A. G." box="[139,279,1363,1389]" pageId="11" pageNumber="778" pagination="1 - 216" refId="ref19687" refString="SHAROV A. G. 1968. - Filogeniya ortopteroidnykh nasekomykh. Trudy Paleontologicheskogo Instituta, Akademiya Nauk S. S. S. R., 118, 1 - 216, Moskva. [in Russian, translated in english in 1971: Phylogeny of the Orthopteroidea. Israel program for scientific translations, Keter Press, Jerusalem: 1 - 251.]" type="journal article" year="1968">Sharov, 1968</bibRefCitation>
)
</taxonomicName>
(
<figureCitation id="EBE42A2EFFD4C73E7565FA935D072524" box="[301,381,1363,1389]" captionStart="FIG" captionStartId="13.[131,142,1941,1959]" captionTargetBox="[186,1379,218,1892]" captionTargetPageId="13" captionText="FIG. 6. — Hypothesis of primary venation homology of male forewing of †Protogryllidae (A, B) and †Baissogryllidae (C-E): A, †Angarogryllus angaricus (Sharov 1968), PIN 1873-16; B, †Falsipseculum karatavicum (Sharov 1968), PIN 3791/1345; C, †Neosharategia paradoxa Gorochov, 1992, PIN 4270-210a; D, †Baissogryllidae sp., CCNH-293;E, †Anglogryllus lyristes Gorochov et al., 2006,MNEMG 2003.46. Abbreviations:“ha”, distally opened harp; “mi”, distally opened mirror, others and colour code, see text. Grey dash lines represent folds. Scale bars: 1 mm." figureDoi="http://doi.org/10.5281/zenodo.10376762" httpUri="https://zenodo.org/record/10376762/files/figure.png" pageId="11" pageNumber="778">Fig. 6B</figureCitation>
), veins between R and M+CuA at the base of the lanceolate cell are very similar and none is stronger than the others, complicating the identification of r-m among the crossveins. This crossvein r-m can either have an oblique direction towards wing base (obliquely inverted) between R and M (
<figureCitation id="EBE42A2EFFD4C73E74E2FA335D412644" box="[170,315,1523,1549]" captionStart="FIG" captionStartId="13.[131,142,1941,1959]" captionTargetBox="[186,1379,218,1892]" captionTargetPageId="13" captionText="FIG. 6. — Hypothesis of primary venation homology of male forewing of †Protogryllidae (A, B) and †Baissogryllidae (C-E): A, †Angarogryllus angaricus (Sharov 1968), PIN 1873-16; B, †Falsipseculum karatavicum (Sharov 1968), PIN 3791/1345; C, †Neosharategia paradoxa Gorochov, 1992, PIN 4270-210a; D, †Baissogryllidae sp., CCNH-293;E, †Anglogryllus lyristes Gorochov et al., 2006,MNEMG 2003.46. Abbreviations:“ha”, distally opened harp; “mi”, distally opened mirror, others and colour code, see text. Grey dash lines represent folds. Scale bars: 1 mm." figureDoi="http://doi.org/10.5281/zenodo.10376762" httpUri="https://zenodo.org/record/10376762/files/figure.png" pageId="11" pageNumber="778">Fig. 6A, B, D</figureCitation>
), or be rather transverse (
<figureCitation id="EBE42A2EFFD4C73E760EFA335ECC2644" box="[582,694,1523,1549]" captionStart="FIG" captionStartId="13.[131,142,1941,1959]" captionTargetBox="[186,1379,218,1892]" captionTargetPageId="13" captionText="FIG. 6. — Hypothesis of primary venation homology of male forewing of †Protogryllidae (A, B) and †Baissogryllidae (C-E): A, †Angarogryllus angaricus (Sharov 1968), PIN 1873-16; B, †Falsipseculum karatavicum (Sharov 1968), PIN 3791/1345; C, †Neosharategia paradoxa Gorochov, 1992, PIN 4270-210a; D, †Baissogryllidae sp., CCNH-293;E, †Anglogryllus lyristes Gorochov et al., 2006,MNEMG 2003.46. Abbreviations:“ha”, distally opened harp; “mi”, distally opened mirror, others and colour code, see text. Grey dash lines represent folds. Scale bars: 1 mm." figureDoi="http://doi.org/10.5281/zenodo.10376762" httpUri="https://zenodo.org/record/10376762/files/figure.png" pageId="11" pageNumber="778">Fig. 6E, C</figureCitation>
). Some †
<taxonomicName id="B4DF4D28FFD4C73E74C6F9D25D512665" authorityName="Gorochov" authorityYear="1985" box="[142,299,1554,1580]" class="Insecta" family="Baissogryllidae" kingdom="Animalia" order="Orthoptera" pageId="11" pageNumber="778" phylum="Arthropoda" rank="family">Baissogryllidae</taxonomicName>
(e.g., †
<taxonomicName id="B4DF4D28FFD4C73E7532F9D25DF82605" authority="Gorochov, Jarzembowski &amp; Coram, 2006" authorityName="Gorochov, Jarzembowski &amp; Coram" authorityYear="2006" class="Insecta" family="Baissogryllidae" genus="Anglogryllus" kingdom="Animalia" order="Orthoptera" pageId="11" pageNumber="778" phylum="Arthropoda" rank="species" species="lyristes">
<emphasis id="41ABEAB9FFD4C73E7532F9D25E412665" box="[378,571,1554,1580]" italics="true" pageId="11" pageNumber="778">Anglogryllus lyristes</emphasis>
<bibRefCitation id="174E4B5AFFD4C73E7609F9D25DF82605" author="GOROCHOV A. V. &amp; JARZEMBOWSKI E. A. &amp; CORAM R. A." pageId="11" pageNumber="778" pagination="641 - 662" refId="ref18426" refString="GOROCHOV A. V., JARZEMBOWSKI E. A. &amp; CORAM R. A. 2006. - Grasshoppers and crickets (Insecta: Orthoptera) from the Lower Cretaceous of Southern England. Cretaceous Research 27: 641 - 662. https: // doi. org / 10.1016 / j. cretres. 2006.03.007" type="journal article" year="2006">Gorochov, Jarzembowski &amp; Coram, 2006</bibRefCitation>
</taxonomicName>
,
<figureCitation id="EBE42A2EFFD4C73E75D9F9F25DA52604" box="[401,479,1586,1613]" captionStart="FIG" captionStartId="13.[131,142,1941,1959]" captionTargetBox="[186,1379,218,1892]" captionTargetPageId="13" captionText="FIG. 6. — Hypothesis of primary venation homology of male forewing of †Protogryllidae (A, B) and †Baissogryllidae (C-E): A, †Angarogryllus angaricus (Sharov 1968), PIN 1873-16; B, †Falsipseculum karatavicum (Sharov 1968), PIN 3791/1345; C, †Neosharategia paradoxa Gorochov, 1992, PIN 4270-210a; D, †Baissogryllidae sp., CCNH-293;E, †Anglogryllus lyristes Gorochov et al., 2006,MNEMG 2003.46. Abbreviations:“ha”, distally opened harp; “mi”, distally opened mirror, others and colour code, see text. Grey dash lines represent folds. Scale bars: 1 mm." figureDoi="http://doi.org/10.5281/zenodo.10376762" httpUri="https://zenodo.org/record/10376762/files/figure.png" pageId="11" pageNumber="778">Fig. 6E</figureCitation>
) have a short r-m making a constriction of the base of the lanceolate cell (
<figureCitation id="EBE42A2EFFD4C73E762DF9925ECF2625" box="[613,693,1618,1644]" captionStart="FIG" captionStartId="13.[131,142,1941,1959]" captionTargetBox="[186,1379,218,1892]" captionTargetPageId="13" captionText="FIG. 6. — Hypothesis of primary venation homology of male forewing of †Protogryllidae (A, B) and †Baissogryllidae (C-E): A, †Angarogryllus angaricus (Sharov 1968), PIN 1873-16; B, †Falsipseculum karatavicum (Sharov 1968), PIN 3791/1345; C, †Neosharategia paradoxa Gorochov, 1992, PIN 4270-210a; D, †Baissogryllidae sp., CCNH-293;E, †Anglogryllus lyristes Gorochov et al., 2006,MNEMG 2003.46. Abbreviations:“ha”, distally opened harp; “mi”, distally opened mirror, others and colour code, see text. Grey dash lines represent folds. Scale bars: 1 mm." figureDoi="http://doi.org/10.5281/zenodo.10376762" httpUri="https://zenodo.org/record/10376762/files/figure.png" pageId="11" pageNumber="778">Fig. 6E</figureCitation>
), a situation similar to that of the modern
<taxonomicName id="B4DF4D28FFD4C73E75BEF9B25E1E26C5" box="[502,612,1650,1676]" class="Aves" family="Trochilidae" genus="Grylloidea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Apodiformes" pageId="11" pageNumber="776" phylum="Chordata" rank="genus">Grylloidea</taxonomicName>
(
<figureCitation id="EBE42A2EFFD4C73E7639F9B25EC826C5" box="[625,690,1650,1676]" captionStart="FIG" captionStartId="8.[132,143,1987,2005]" captionTargetBox="[151,1436,215,1942]" captionTargetId="graphics-64@8.[151,1395,215,1926]" captionTargetPageId="8" captionText="FIG. 3. — Hypothesis of primary homology of venation of male Grylloidea:A, Natula longipennis (Serville, 1838) (MNHN-EO-ENSIF9933, Trigonidiidae);B, Lerneca fuscipennis (Saussure, 1874) (MNHN-EO-ENSIF9780, Phalangopsidae). Abbreviations and colour code: see text. Scale bars:1 mm." figureDoi="http://doi.org/10.5281/zenodo.10376755" httpUri="https://zenodo.org/record/10376755/files/figure.png" pageId="11" pageNumber="778">Figs 3</figureCitation>
;
<figureCitation id="EBE42A2EFFD4C73E76F6F9B25EB426C5" box="[702,718,1650,1676]" captionStart="FIG" captionStartId="10.[133,144,1583,1601]" captionTargetBox="[132,1456,203,1553]" captionTargetId="graphics-253@10.[132,1444,240,1511]" captionTargetPageId="10" captionText="FIG. 4. — Hypothesis of primary homology of venation of male Grylloidea: A, Brachytrupes membranaceus (Drury, 1773) (MNHN-EO-ENSIF9769, Gryllidae); B, Phyllogryllus sp. (MNHN-EO-ENSIF9768, Oecanthidae). Abbreviations and colour code: see text. Grey dash lines represent folds. Scale bars: 5 mm." figureDoi="http://doi.org/10.5281/zenodo.10376758" httpUri="https://zenodo.org/record/10376758/files/figure.png" pageId="11" pageNumber="778">4</figureCitation>
;
<figureCitation id="EBE42A2EFFD4C73E7691F9B25E8626C4" box="[729,764,1650,1677]" captionStart="FIG" captionStartId="12.[132,143,1940,1958]" captionTargetBox="[132,1456,169,1827]" captionTargetId="graphics-22@12.[132,1411,884,1765]" captionTargetPageId="12" captionText="FIG. 5. — Hypothesis of primary venation homology in male Grylloidea with particular forewing venation:A, B, species with shortened wings; B, C, species with reduced stridulatory apparatus.A, Landreva sp. (MNHN-EO-ENSIF9775, Gryllidae); B, Nemobius sylvestris (Bosc, 1792) (MNHN-EO-ENSIF9786, Trigonidiidae); C, Tafalisca lineatipes Bruner,1916 (MNHN-EO-ENSIF9760, Oecanthidae);D, Aphonomorphus sp.(MNHN-EO-ENSIF9764, Oecanthidae).Abbreviations:ha, distally opened harp; mi distally opened mirror; others and colour code: see text. Grey dash lines represent folds. Scale bars: 1 mm." figureDoi="http://doi.org/10.5281/zenodo.10376760" httpUri="https://zenodo.org/record/10376760/files/figure.png" pageId="11" pageNumber="778">5A</figureCitation>
).
</paragraph>
<paragraph id="736036ABFFD4C73E74D3F9535E9327A2" blockId="11.[130,777,694,2027]" pageId="11" pageNumber="778">
R divides distally into a convex anterior branch RA and a rather concave posterior branch RP, well visible in †
<taxonomicName id="B4DF4D28FFD4C73E76E1F9725D5B26A5" baseAuthorityName="Sharov" baseAuthorityYear="1968" class="Insecta" family="Protogryllidae" genus="Angarogryllus" kingdom="Animalia" order="Orthoptera" pageId="11" pageNumber="778" phylum="Arthropoda" rank="species" species="angaricus">
<emphasis id="41ABEAB9FFD4C73E76E1F9725D5B26A5" italics="true" pageId="11" pageNumber="778">Angarogryllus angaricus</emphasis>
</taxonomicName>
(
<figureCitation id="EBE42A2EFFD4C73E7566F9125DD726A5" box="[302,429,1746,1772]" captionStart="APPENDIX" captionStartId="26.[378,390,219,237]" captionText="APPENDIX 3. — Photos of studied material called as supplementary figures in the present paper." pageId="11" pageNumber="778">Appendix 3</figureCitation>
:
<figureCitation id="EBE42A2EFFD4C73E75F0F9125E6D26A4" box="[440,535,1746,1773]" captionStart="FIG" captionStartId="12.[132,143,1940,1958]" captionTargetBox="[132,1456,169,1827]" captionTargetId="graphics-22@12.[132,1411,884,1765]" captionTargetPageId="12" captionText="FIG. 5. — Hypothesis of primary venation homology in male Grylloidea with particular forewing venation:A, B, species with shortened wings; B, C, species with reduced stridulatory apparatus.A, Landreva sp. (MNHN-EO-ENSIF9775, Gryllidae); B, Nemobius sylvestris (Bosc, 1792) (MNHN-EO-ENSIF9786, Trigonidiidae); C, Tafalisca lineatipes Bruner,1916 (MNHN-EO-ENSIF9760, Oecanthidae);D, Aphonomorphus sp.(MNHN-EO-ENSIF9764, Oecanthidae).Abbreviations:ha, distally opened harp; mi distally opened mirror; others and colour code: see text. Grey dash lines represent folds. Scale bars: 1 mm." figureDoi="http://doi.org/10.5281/zenodo.10376760" httpUri="https://zenodo.org/record/10376760/files/figure.png" pageId="11" pageNumber="778">Fig. S5A</figureCitation>
). RP is strongly curved basally and partially fuses with MA. The latter, also well visible in †
<taxonomicName id="B4DF4D28FFD4C73E74E3F8D25DDA2762" baseAuthorityName="Sharov" baseAuthorityYear="1968" box="[171,416,1810,1836]" class="Insecta" family="Protogryllidae" genus="Angarogryllus" kingdom="Animalia" order="Orthoptera" pageId="11" pageNumber="778" phylum="Arthropoda" rank="species" species="angaricus">
<emphasis id="41ABEAB9FFD4C73E74E3F8D25DDA2762" box="[171,416,1810,1836]" italics="true" pageId="11" pageNumber="778">Angarogryllus angaricus</emphasis>
</taxonomicName>
, is clearly convex. The fan is often very poorly preserved in fossils, especially for the MP branch. However, applying a conservative approach, we consider that M divides into two branches MA and MP (notably slightly visible in the unidentified †
<taxonomicName id="B4DF4D28FFD4C73E75E0F8515E3027E2" authorityName="Gorochov" authorityYear="1985" box="[424,586,1937,1963]" class="Insecta" family="Baissogryllidae" kingdom="Animalia" order="Orthoptera" pageId="11" pageNumber="778" phylum="Arthropoda" rank="family">Baissogryllidae</taxonomicName>
no. CCNH-293,
<figureCitation id="EBE42A2EFFD4C73E74CCF8715CA12782" box="[132,219,1969,1995]" captionStart="FIG" captionStartId="13.[131,142,1941,1959]" captionTargetBox="[186,1379,218,1892]" captionTargetPageId="13" captionText="FIG. 6. — Hypothesis of primary venation homology of male forewing of †Protogryllidae (A, B) and †Baissogryllidae (C-E): A, †Angarogryllus angaricus (Sharov 1968), PIN 1873-16; B, †Falsipseculum karatavicum (Sharov 1968), PIN 3791/1345; C, †Neosharategia paradoxa Gorochov, 1992, PIN 4270-210a; D, †Baissogryllidae sp., CCNH-293;E, †Anglogryllus lyristes Gorochov et al., 2006,MNEMG 2003.46. Abbreviations:“ha”, distally opened harp; “mi”, distally opened mirror, others and colour code, see text. Grey dash lines represent folds. Scale bars: 1 mm." figureDoi="http://doi.org/10.5281/zenodo.10376762" httpUri="https://zenodo.org/record/10376762/files/figure.png" pageId="11" pageNumber="778">Fig. 6D</figureCitation>
). These two branches are relatively thin and present in the fan (a situation similar in the modern
<taxonomicName id="B4DF4D28FFD4C73E762EF8115EA727A2" box="[614,733,2001,2027]" class="Aves" family="Trochilidae" genus="Grylloidea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Apodiformes" pageId="11" pageNumber="776" phylum="Chordata" rank="genus">Grylloidea</taxonomicName>
).
</paragraph>
<paragraph id="736036ABFFD4C73E770BFF1758A724A7" blockId="11.[811,1457,215,1262]" pageId="11" pageNumber="778">
Dorsal field. The clearly convex CuA diverges from M and joins CuPa crossing the median fold (well visible in †
<taxonomicName id="B4DF4D28FFD4C73E7125FF375F882178" baseAuthorityName="Sharov" baseAuthorityYear="1968" class="Insecta" family="Protogryllidae" genus="Angarogryllus" kingdom="Animalia" order="Orthoptera" pageId="11" pageNumber="778" phylum="Arthropoda" rank="species" species="angaricus">
<emphasis id="41ABEAB9FFD4C73E7125FF375F882178" italics="true" pageId="11" pageNumber="778">Angarogryllus angaricus</emphasis>
</taxonomicName>
,
<figureCitation id="EBE42A2EFFD4C73E77B7FED758F92178" box="[1023,1155,279,305]" captionStart="APPENDIX" captionStartId="26.[378,390,219,237]" captionText="APPENDIX 3. — Photos of studied material called as supplementary figures in the present paper." pageId="11" pageNumber="778">Appendix 3</figureCitation>
:
<figureCitation id="EBE42A2EFFD4C73E70C6FED75888217B" box="[1166,1266,279,306]" captionStart="FIG" captionStartId="12.[132,143,1940,1958]" captionTargetBox="[132,1456,169,1827]" captionTargetId="graphics-22@12.[132,1411,884,1765]" captionTargetPageId="12" captionText="FIG. 5. — Hypothesis of primary venation homology in male Grylloidea with particular forewing venation:A, B, species with shortened wings; B, C, species with reduced stridulatory apparatus.A, Landreva sp. (MNHN-EO-ENSIF9775, Gryllidae); B, Nemobius sylvestris (Bosc, 1792) (MNHN-EO-ENSIF9786, Trigonidiidae); C, Tafalisca lineatipes Bruner,1916 (MNHN-EO-ENSIF9760, Oecanthidae);D, Aphonomorphus sp.(MNHN-EO-ENSIF9764, Oecanthidae).Abbreviations:ha, distally opened harp; mi distally opened mirror; others and colour code: see text. Grey dash lines represent folds. Scale bars: 1 mm." figureDoi="http://doi.org/10.5281/zenodo.10376760" httpUri="https://zenodo.org/record/10376760/files/figure.png" pageId="11" pageNumber="778">Fig. S5A</figureCitation>
). In the †
<taxonomicName id="B4DF4D28FFD4C73E7111FED75FD52118" authority="and" authorityName="AND" class="Insecta" family="Protogryllidae" kingdom="Animalia" order="Orthoptera" pageId="11" pageNumber="778" phylum="Arthropoda" rank="family">Protogryllidae and</taxonomicName>
some †
<taxonomicName id="B4DF4D28FFD4C73E704BFEF758D72118" authorityName="Gorochov" authorityYear="1985" box="[1027,1197,311,337]" class="Insecta" family="Baissogryllidae" kingdom="Animalia" order="Orthoptera" pageId="11" pageNumber="778" phylum="Arthropoda" rank="family">Baissogryllidae</taxonomicName>
, the CuA is longer and oblique (
<figureCitation id="EBE42A2EFFD4C73E77C4FE9758662138" box="[908,1052,343,369]" captionStart="FIG" captionStartId="13.[131,142,1941,1959]" captionTargetBox="[186,1379,218,1892]" captionTargetPageId="13" captionText="FIG. 6. — Hypothesis of primary venation homology of male forewing of †Protogryllidae (A, B) and †Baissogryllidae (C-E): A, †Angarogryllus angaricus (Sharov 1968), PIN 1873-16; B, †Falsipseculum karatavicum (Sharov 1968), PIN 3791/1345; C, †Neosharategia paradoxa Gorochov, 1992, PIN 4270-210a; D, †Baissogryllidae sp., CCNH-293;E, †Anglogryllus lyristes Gorochov et al., 2006,MNEMG 2003.46. Abbreviations:“ha”, distally opened harp; “mi”, distally opened mirror, others and colour code, see text. Grey dash lines represent folds. Scale bars: 1 mm." figureDoi="http://doi.org/10.5281/zenodo.10376762" httpUri="https://zenodo.org/record/10376762/files/figure.png" pageId="11" pageNumber="778">Fig. 6A, B, C</figureCitation>
), whereas in other †
<taxonomicName id="B4DF4D28FFD4C73E70A7FE9759EC2138" authorityName="Gorochov" authorityYear="1985" box="[1263,1430,343,369]" class="Insecta" family="Baissogryllidae" kingdom="Animalia" order="Orthoptera" pageId="11" pageNumber="778" phylum="Arthropoda" rank="family">Baissogryllidae</taxonomicName>
, it is transverse between the two fields (
<figureCitation id="EBE42A2EFFD4C73E70E0FEB7596D21D8" box="[1192,1303,375,401]" captionStart="FIG" captionStartId="13.[131,142,1941,1959]" captionTargetBox="[186,1379,218,1892]" captionTargetPageId="13" captionText="FIG. 6. — Hypothesis of primary venation homology of male forewing of †Protogryllidae (A, B) and †Baissogryllidae (C-E): A, †Angarogryllus angaricus (Sharov 1968), PIN 1873-16; B, †Falsipseculum karatavicum (Sharov 1968), PIN 3791/1345; C, †Neosharategia paradoxa Gorochov, 1992, PIN 4270-210a; D, †Baissogryllidae sp., CCNH-293;E, †Anglogryllus lyristes Gorochov et al., 2006,MNEMG 2003.46. Abbreviations:“ha”, distally opened harp; “mi”, distally opened mirror, others and colour code, see text. Grey dash lines represent folds. Scale bars: 1 mm." figureDoi="http://doi.org/10.5281/zenodo.10376762" httpUri="https://zenodo.org/record/10376762/files/figure.png" pageId="11" pageNumber="778">Fig. 6D,E</figureCitation>
). CuA merges with CuPaα, after its bifurcation with CuPaβ and before the bifurcation between CuPaα1 and CuPaα2. Between CuPaα and CuPaβ, there is a strong crossvein, which corresponds to d1 (
<figureCitation id="EBE42A2EFFD4C73E7731FE365FC42259" box="[889,958,502,528]" captionStart="FIG" captionStartId="13.[131,142,1941,1959]" captionTargetBox="[186,1379,218,1892]" captionTargetPageId="13" captionText="FIG. 6. — Hypothesis of primary venation homology of male forewing of †Protogryllidae (A, B) and †Baissogryllidae (C-E): A, †Angarogryllus angaricus (Sharov 1968), PIN 1873-16; B, †Falsipseculum karatavicum (Sharov 1968), PIN 3791/1345; C, †Neosharategia paradoxa Gorochov, 1992, PIN 4270-210a; D, †Baissogryllidae sp., CCNH-293;E, †Anglogryllus lyristes Gorochov et al., 2006,MNEMG 2003.46. Abbreviations:“ha”, distally opened harp; “mi”, distally opened mirror, others and colour code, see text. Grey dash lines represent folds. Scale bars: 1 mm." figureDoi="http://doi.org/10.5281/zenodo.10376762" httpUri="https://zenodo.org/record/10376762/files/figure.png" pageId="11" pageNumber="778">Fig. 6</figureCitation>
). Another strong crossvein d2, in alignment with d1, is present between CuPaβ and the anal node at the level of the plectrum. The harp is thus located between CuPa, CuPaβ, d2 and PCuA. In observed †
<taxonomicName id="B4DF4D28FFD4C73E7158FD965F2C22D9" authority="and" authorityName="AND" class="Insecta" family="Protogryllidae" kingdom="Animalia" order="Orthoptera" pageId="11" pageNumber="778" phylum="Arthropoda" rank="family">Protogryllidae and</taxonomicName>
<taxonomicName id="B4DF4D28FFD4C73E772FFDB6587422D9" authorityName="Gorochov" authorityYear="1985" box="[871,1038,630,656]" class="Insecta" family="Baissogryllidae" kingdom="Animalia" order="Orthoptera" pageId="11" pageNumber="778" phylum="Arthropoda" rank="family">Baissogryllidae</taxonomicName>
(see
<figureCitation id="EBE42A2EFFD4C73E700DFDB658B222D9" box="[1093,1224,630,656]" captionStart="APPENDIX" captionStartId="26.[378,390,219,237]" captionText="APPENDIX 3. — Photos of studied material called as supplementary figures in the present paper." pageId="11" pageNumber="778">Appendix 3</figureCitation>
:
<figureCitation id="EBE42A2EFFD4C73E709BFDB6595F22D8" box="[1235,1317,630,657]" captionStart="FIG" captionStartId="12.[132,143,1940,1958]" captionTargetBox="[132,1456,169,1827]" captionTargetId="graphics-22@12.[132,1411,884,1765]" captionTargetPageId="12" captionText="FIG. 5. — Hypothesis of primary venation homology in male Grylloidea with particular forewing venation:A, B, species with shortened wings; B, C, species with reduced stridulatory apparatus.A, Landreva sp. (MNHN-EO-ENSIF9775, Gryllidae); B, Nemobius sylvestris (Bosc, 1792) (MNHN-EO-ENSIF9786, Trigonidiidae); C, Tafalisca lineatipes Bruner,1916 (MNHN-EO-ENSIF9760, Oecanthidae);D, Aphonomorphus sp.(MNHN-EO-ENSIF9764, Oecanthidae).Abbreviations:ha, distally opened harp; mi distally opened mirror; others and colour code: see text. Grey dash lines represent folds. Scale bars: 1 mm." figureDoi="http://doi.org/10.5281/zenodo.10376760" httpUri="https://zenodo.org/record/10376760/files/figure.png" pageId="11" pageNumber="778">Figs S5</figureCitation>
; S
<figureCitation id="EBE42A2EFFD4C73E7108FDB6593522D9" box="[1344,1359,630,656]" captionStart="FIG" captionStartId="13.[131,142,1941,1959]" captionTargetBox="[186,1379,218,1892]" captionTargetPageId="13" captionText="FIG. 6. — Hypothesis of primary venation homology of male forewing of †Protogryllidae (A, B) and †Baissogryllidae (C-E): A, †Angarogryllus angaricus (Sharov 1968), PIN 1873-16; B, †Falsipseculum karatavicum (Sharov 1968), PIN 3791/1345; C, †Neosharategia paradoxa Gorochov, 1992, PIN 4270-210a; D, †Baissogryllidae sp., CCNH-293;E, †Anglogryllus lyristes Gorochov et al., 2006,MNEMG 2003.46. Abbreviations:“ha”, distally opened harp; “mi”, distally opened mirror, others and colour code, see text. Grey dash lines represent folds. Scale bars: 1 mm." figureDoi="http://doi.org/10.5281/zenodo.10376762" httpUri="https://zenodo.org/record/10376762/files/figure.png" pageId="11" pageNumber="778">6</figureCitation>
), we noticed a reduced vein in the harp that could correspond to the CuPb, because of its base connected to that of CuPa. These fossils have two branches of PCu, PCuA and PCuP, with their characteristic strong and curved base (
<figureCitation id="EBE42A2EFFD4C73E7114FD3559D82346" box="[1372,1442,757,783]" captionStart="FIG" captionStartId="13.[131,142,1941,1959]" captionTargetBox="[186,1379,218,1892]" captionTargetPageId="13" captionText="FIG. 6. — Hypothesis of primary venation homology of male forewing of †Protogryllidae (A, B) and †Baissogryllidae (C-E): A, †Angarogryllus angaricus (Sharov 1968), PIN 1873-16; B, †Falsipseculum karatavicum (Sharov 1968), PIN 3791/1345; C, †Neosharategia paradoxa Gorochov, 1992, PIN 4270-210a; D, †Baissogryllidae sp., CCNH-293;E, †Anglogryllus lyristes Gorochov et al., 2006,MNEMG 2003.46. Abbreviations:“ha”, distally opened harp; “mi”, distally opened mirror, others and colour code, see text. Grey dash lines represent folds. Scale bars: 1 mm." figureDoi="http://doi.org/10.5281/zenodo.10376762" httpUri="https://zenodo.org/record/10376762/files/figure.png" pageId="11" pageNumber="778">Fig. 6</figureCitation>
). While it has been suggested that the †
<taxonomicName id="B4DF4D28FFD4C73E7090FCD559052366" authorityName="Gorochov" authorityYear="1985" box="[1240,1407,789,815]" class="Insecta" family="Baissogryllidae" kingdom="Animalia" order="Orthoptera" pageId="11" pageNumber="778" phylum="Arthropoda" rank="family">Baissogryllidae</taxonomicName>
and †
<taxonomicName id="B4DF4D28FFD4C73E777FFCF55FAC2306" box="[823,982,821,847]" class="Insecta" family="Protogryllidae" kingdom="Animalia" order="Orthoptera" pageId="11" pageNumber="778" phylum="Arthropoda" rank="family">Protogryllidae</taxonomicName>
may have a file as in modern crickets (i.e., on the highly curved PCu), no stridulatory teeth are visible on photographs or illustrations of fossil specimens that we have studied. Some teeth are however visible on the PCuA in some undescribed baissogryllids from the lower Jurassic of
<collectingCountry id="0BC8763BFFD4C73E770FFC145FA923A7" box="[839,979,980,1006]" name="Luxembourg" pageId="11" pageNumber="778">Luxembourg</collectingCountry>
(H. J. and A. N., pers. obs.), supporting this hypothesis, but it cannot be generalized to all baissogryllid and protogryllid fossils. Two anal veins, AA and AP, are located posteriorly. In the †
<taxonomicName id="B4DF4D28FFD4C73E700EFBF458972407" authorityName="Gorochov" authorityYear="1985" box="[1094,1261,1076,1102]" class="Insecta" family="Baissogryllidae" kingdom="Animalia" order="Orthoptera" pageId="11" pageNumber="778" phylum="Arthropoda" rank="family">Baissogryllidae</taxonomicName>
that we observed (
<figureCitation id="EBE42A2EFFD4C73E777DFB945FB12427" box="[821,971,1108,1135]" captionStart="FIG" captionStartId="13.[131,142,1941,1959]" captionTargetBox="[186,1379,218,1892]" captionTargetPageId="13" captionText="FIG. 6. — Hypothesis of primary venation homology of male forewing of †Protogryllidae (A, B) and †Baissogryllidae (C-E): A, †Angarogryllus angaricus (Sharov 1968), PIN 1873-16; B, †Falsipseculum karatavicum (Sharov 1968), PIN 3791/1345; C, †Neosharategia paradoxa Gorochov, 1992, PIN 4270-210a; D, †Baissogryllidae sp., CCNH-293;E, †Anglogryllus lyristes Gorochov et al., 2006,MNEMG 2003.46. Abbreviations:“ha”, distally opened harp; “mi”, distally opened mirror, others and colour code, see text. Grey dash lines represent folds. Scale bars: 1 mm." figureDoi="http://doi.org/10.5281/zenodo.10376762" httpUri="https://zenodo.org/record/10376762/files/figure.png" pageId="11" pageNumber="778">Fig. 6C, D, E</figureCitation>
) but not in the †
<taxonomicName id="B4DF4D28FFD4C73E70C9FB94595A2427" box="[1153,1312,1108,1134]" class="Insecta" family="Protogryllidae" kingdom="Animalia" order="Orthoptera" pageId="11" pageNumber="778" phylum="Arthropoda" rank="family">Protogryllidae</taxonomicName>
(
<figureCitation id="EBE42A2EFFD4C73E7167FB9459DB2427" box="[1327,1441,1108,1134]" captionStart="FIG" captionStartId="13.[131,142,1941,1959]" captionTargetBox="[186,1379,218,1892]" captionTargetPageId="13" captionText="FIG. 6. — Hypothesis of primary venation homology of male forewing of †Protogryllidae (A, B) and †Baissogryllidae (C-E): A, †Angarogryllus angaricus (Sharov 1968), PIN 1873-16; B, †Falsipseculum karatavicum (Sharov 1968), PIN 3791/1345; C, †Neosharategia paradoxa Gorochov, 1992, PIN 4270-210a; D, †Baissogryllidae sp., CCNH-293;E, †Anglogryllus lyristes Gorochov et al., 2006,MNEMG 2003.46. Abbreviations:“ha”, distally opened harp; “mi”, distally opened mirror, others and colour code, see text. Grey dash lines represent folds. Scale bars: 1 mm." figureDoi="http://doi.org/10.5281/zenodo.10376762" httpUri="https://zenodo.org/record/10376762/files/figure.png" pageId="11" pageNumber="778">Fig. 6A, B</figureCitation>
), the area between the CuPaα2 and CuPaβ is widened: by its position, this area could correspond to the mirror cell
<emphasis id="41ABEAB9FFD4C73E7133FB5559CA24E7" box="[1403,1456,1173,1198]" italics="true" pageId="11" pageNumber="778">s. str.</emphasis>
(mi) of modern crickets, but it is distally opened and not closed by s1 as it is in modern crickets.
</paragraph>
</subSubSection>
</treatment>
</document>