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<mods:title>Leveraging female genitalic characters for generic and species delimitation in Nilomantis Werner, 1907 and Ilomantis Giglio-Tos, 1915 (Mantodea, Nilomantinae)</mods:title>
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<mods:namePart>Sydney K. Brannoch</mods:namePart>
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<mods:namePart>Gavin J. Svenson</mods:namePart>
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<mods:title>Insect Systematics &amp; Evolution</mods:title>
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<mods:date>2016</mods:date>
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<heading bold="true" box="[98,482,480,507]" fontSize="11" level="1" pageId="29" pageNumber="238" reason="0">
<taxonomicName authority="Roy, 1975" authorityName="Roy" authorityYear="1975" box="[98,482,480,507]" class="Insecta" family="Iridopterygidae" genus="Nilomantis" kingdom="Animalia" order="Mantodea" pageId="29" pageNumber="238" phylum="Arthropoda" rank="species" species="edmundsi">
<emphasis bold="true" box="[98,482,480,507]" pageId="29" pageNumber="238">
<emphasis bold="true" box="[98,354,480,507]" italics="true" pageId="29" pageNumber="238">Nilomantis edmundsi</emphasis>
<bibRefCitation author="Roy, R. &amp; Leston, D." box="[361,482,481,507]" journalOrPublisher="Bulletin de l'Institut Fondamental d'Afrique Noire (IFAN) Serie A" pageId="29" pageNumber="238" pagination="297 - 344" part="37" refString="Roy, R. &amp; Leston, D. (1975) Mantodea of Ghana: new species, further records and habitats. Bulletin de l'Institut Fondamental d'Afrique Noire (IFAN) Serie A 37: 297 - 344." title="Mantodea of Ghana: new species, further records and habitats" type="journal article" year="1975">
Roy, 
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</emphasis>
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</heading>
</paragraph>
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<paragraph blockId="29.[98,1066,533,595]" pageId="29" pageNumber="238">
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<taxonomicName authority="Roy, 1975: 313" authorityName="Roy" authorityPageNumber="313" authorityYear="1975" box="[98,515,533,560]" class="Insecta" family="Iridopterygidae" genus="Nilomantis" kingdom="Animalia" order="Mantodea" pageId="29" pageNumber="238" phylum="Arthropoda" rank="species" species="edmundsi">
<emphasis box="[98,331,533,559]" italics="true" pageId="29" pageNumber="238">Nilomantis edmundsi</emphasis>
<treatmentCitation author="Roy, R. &amp; Leston, D." box="[339,515,533,560]" page="313" pageId="29" pageNumber="238" year="1975">
<bibRefCitation author="Roy, R. &amp; Leston, D." box="[339,459,533,560]" journalOrPublisher="Bulletin de l'Institut Fondamental d'Afrique Noire (IFAN) Serie A" pageId="29" pageNumber="238" pagination="297 - 344" part="37" refString="Roy, R. &amp; Leston, D. (1975) Mantodea of Ghana: new species, further records and habitats. Bulletin de l'Institut Fondamental d'Afrique Noire (IFAN) Serie A 37: 297 - 344." title="Mantodea of Ghana: new species, further records and habitats" type="journal article" year="1975">
Roy, 
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: 
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; 
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<bibRefCitation author="Edmunds, M." box="[588,768,533,559]" journalOrPublisher="Zoological Journal of the Linnean Society" pageId="29" pageNumber="238" pagination="1 - 37" part="58" refString="Edmunds, M. (1976) The defensive behaviour of Ghanaian praying mantids with a discussion of territoriality. Zoological Journal of the Linnean Society 58: 1 - 37." title="The defensive behaviour of Ghanaian praying mantids with a discussion of territoriality" type="journal article" year="1976">
Edmunds 
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: 
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–7, 34; 
<treatmentCitation author="Edmunds, M." page="41" pageId="29" pageNumber="238" year="1986">
<bibRefCitation author="Edmunds, M." box="[883,1062,533,559]" journalOrPublisher="Journal of Tropical Ecology" pageId="29" pageNumber="238" pagination="39 - 50" part="2" refString="Edmunds, M. (1986) The phenology and diversity of praying mantids in Ghana. Journal of Tropical Ecology 2: 39 - 50." title="The phenology and diversity of praying mantids in Ghana" type="journal article" year="1986">
Edmunds 
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: 41
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, 45; 
<treatmentCitation author="Ehrmann, R." box="[186,421,568,594]" page="242" pageId="29" pageNumber="238" year="2002">
<bibRefCitation author="Ehrmann, R." box="[186,363,568,594]" journalOrPublisher="Munster, Natur und Tier Verlag Munster" pageId="29" pageNumber="238" refString="Ehrmann, R. (2002) Mantodea: Gottesanberinnen der Welt. Munster, Natur und Tier Verlag Munster." title="Mantodea: Gottesanberinnen der Welt" type="book" year="2002">
Ehrmann 
<number box="[301,363,568,594]" pageId="29" pageNumber="238" value="2002.0">2002</number>
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: 
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; 
<treatmentCitation author="Otte, D. &amp; Spearman, L. A." box="[433,762,568,595]" page="120" pageId="29" pageNumber="238" year="2005">
<bibRefCitation author="Otte, D. &amp; Spearman, L. A." box="[433,706,568,595]" journalOrPublisher="Catalog of the mantids of the world" pageId="29" pageNumber="238" part="1" refString="Otte, D. &amp; Spearman, L. A. (2005) Mantida species file. Catalog of the mantids of the world, volume 1. Association of Insect Diversity, Philadelphia, PA." title="Mantida species file" type="journal volume" year="2005">
Otte &amp; Spearman 
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: 
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</paragraph>
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<subSubSection pageId="29" pageNumber="238" type="materials_examined">
<paragraph blockId="29.[98,319,637,663]" box="[98,319,637,663]" pageId="29" pageNumber="238">
<heading box="[98,319,637,663]" fontSize="11" level="2" pageId="29" pageNumber="238" reason="1">
<emphasis box="[98,319,637,663]" italics="true" pageId="29" pageNumber="238">Examined specimens</emphasis>
</heading>
</paragraph>
<paragraph blockId="29.[98,1066,689,751]" pageId="29" pageNumber="238">
The list consists of 
<specimenCount box="[339,476,689,715]" pageId="29" pageNumber="238" type="generic">
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specimen
</specimenCount>
. 
<collectingCountry box="[494,573,689,715]" name="Ghana" pageId="29" pageNumber="238">Ghana</collectingCountry>
; 
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♀, 
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001500, MN 
<number box="[890,935,689,715]" pageId="29" pageNumber="238" value="244.0">244</number>
Secondary voucher, Genitalia No. 0 0 0 
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0 
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S
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. K. Brannoch.
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<subSubSection lastPageId="30" lastPageNumber="239" pageId="29" pageNumber="238" type="description">
<paragraph blockId="29.[98,379,793,819]" box="[98,379,793,819]" pageId="29" pageNumber="238">
<heading box="[98,379,793,819]" fontSize="11" level="2" pageId="29" pageNumber="238" reason="1">
<emphasis box="[98,379,793,819]" italics="true" pageId="29" pageNumber="238">Male genitalia description</emphasis>
</heading>
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<paragraph blockId="29.[98,1066,845,1115]" pageId="29" pageNumber="238">
An examination of the plates presented by 
<bibRefCitation author="Roy, R. &amp; Leston, D." box="[602,855,845,872]" journalOrPublisher="Bulletin de l'Institut Fondamental d'Afrique Noire (IFAN) Serie A" pageId="29" pageNumber="238" pagination="297 - 344" part="37" refString="Roy, R. &amp; Leston, D. (1975) Mantodea of Ghana: new species, further records and habitats. Bulletin de l'Institut Fondamental d'Afrique Noire (IFAN) Serie A 37: 297 - 344." title="Mantodea of Ghana: new species, further records and habitats" type="journal article" year="1975">
Roy &amp; Leston (
<number box="[786,846,845,872]" pageId="29" pageNumber="238" value="1975.0">1975</number>
)
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indicates that the anterior process (ap) is narrow. Apical process (paa) broad and recurved. Apophysis phalloide (L 
<number box="[239,249,915,941]" pageId="29" pageNumber="238" value="1.0">1</number>
) does not retain the apophysis; lobo membranoso (loa) lightly sclerotized and crenulated on the distal margin. Ventral sclerotization of the left phallomeric process (L 
<number box="[176,190,984,1010]" pageId="29" pageNumber="238" value="4.0">4</number>
A) terminates in two lobes; processo distale (pda) largely rounded and well separated from a secondary lobe that is rounded and relatively smaller. Processo ventrale sclerificato (pva) and piastra ventrale (pia) sclerotized and more well defined than in 
<taxonomicName box="[129,238,1088,1114]" class="Insecta" family="Iridopterygidae" genus="Nilomantis" kingdom="Animalia" order="Mantodea" pageId="29" pageNumber="238" phylum="Arthropoda" rank="species" species="floweri">
<emphasis box="[129,238,1088,1114]" italics="true" pageId="29" pageNumber="238">N. floweri</emphasis>
</taxonomicName>
; right arm (bm) pointed distally and lightly sclerotized.
</paragraph>
<paragraph blockId="29.[98,584,1157,1183]" box="[98,584,1157,1183]" pageId="29" pageNumber="238">
<heading box="[98,584,1157,1183]" fontSize="11" level="2" pageId="29" pageNumber="238" reason="1">
<emphasis box="[98,584,1157,1183]" italics="true" pageId="29" pageNumber="238">
Female genitalia description (
<figureCitation box="[414,481,1157,1183]" captionStart="Fig. 4" captionStartId="8.[106,139,1540,1562]" captionTargetBox="[155,1024,390,1513]" captionTargetId="figure@8.[155,1024,390,1515]" captionTargetPageId="8" captionText="Fig. 4. Female genitalic complex of Nilomantis shown in the lateral perspective. (A) Nilomantis floweri; (B) Nilomantis edmundsi. Abbreviations: KLP, knob-like projection; RD, round dilation. This figure is published in colour in the online edition of this journal, which can be accessed via http: // booksand journals. brillonline. com / content / journals / 1876312 x." httpUri="https://zenodo.org/record/269843/files/figure.png" pageId="29" pageNumber="238">
Figs 
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</figureCitation>
B, 
<number box="[512,528,1157,1183]" pageId="29" pageNumber="238" value="6.0">6</number>
C,D)
</emphasis>
</heading>
</paragraph>
<paragraph blockId="29.[98,1067,1209,1652]" lastBlockId="30.[106,1074,134,403]" lastPageId="30" lastPageNumber="239" pageId="29" pageNumber="238">Gonapophysis VIII (GA) slender, variably ciliated, slightly arcuate, and visibly curved from both the ventral (VH) and lateral (LH) perspectives; GA surfaced with fossettes. From VH, gonocoxa VIII (VF) internal margin narrow, rounded, lending VF an angular appearance; from LH, VF possesses a distinct knob-like projection that is directed dorsad. From VH, GA gently curves inward. From LH, GA obtusely curved ventrad; GA ventral margin features a rugose, sparsely ciliated medial outgrowth (MO), which projects posterior to the crest of the curve. GA external margin possesses a moderately sclerotized tubercle anterior to MO. From VH, apical lobe (AL) ovoid, pitted, distinctly lacking cilia across surface majority; from LH, AL thin, arched, pitted, and approximately pointed; no visible cilia observed. AL has a pronounced membrane that projects from the internal dorsal margin of the structure, forming a shallow pocket as the membrane merges with the dorsal margin of GA, lending AL the appearance of being bilobed or mitten-shaped. GA dorsal margin invaginated at the region that bends towards the ventral habitus. Gonapophysis IX (GP) relatively short, broadening at the site of the medial tine (MT) before narrowing into a tapered, acuminate apex; GP ventral margin invaginated, covered anteriorly by MT. Gonoplac (GL) shorter than GA, broad, and obtusely arched ventrad; from LH, GL base has a rounded dilation on the anteroventral margin. GL narrows toward the apex; from LH, GL apex slants ventrad; apical cleft (AC) reduced, resulting in a slight apical notch. From LH, GL possesses a pronounced sclerotization that traverses the length of the structure, terminating just before the AC. GL dorsal and ventral margins relatively smooth.</paragraph>
</subSubSection>
<subSubSection lastPageId="31" lastPageNumber="240" pageId="30" pageNumber="239" type="discussion">
<paragraph blockId="30.[106,237,465,491]" box="[106,237,465,491]" pageId="30" pageNumber="239">
<heading bold="true" box="[106,237,465,491]" fontSize="11" level="1" pageId="30" pageNumber="239" reason="0">
<emphasis bold="true" box="[106,237,465,491]" pageId="30" pageNumber="239">Discussion</emphasis>
</heading>
</paragraph>
<paragraph blockId="30.[106,1074,517,1653]" pageId="30" pageNumber="239">
As a means to consider the validity of 
<taxonomicName box="[535,638,517,543]" class="Insecta" family="Iridopterygidae" genus="Ilomantis" kingdom="Animalia" order="Mantodea" pageId="30" pageNumber="239" phylum="Arthropoda" rank="genus">
<emphasis box="[535,638,517,543]" italics="true" pageId="30" pageNumber="239">Ilomantis</emphasis>
</taxonomicName>
, we examined the female genital complexes of the species included in 
<taxonomicName box="[488,611,552,578]" class="Insecta" family="Iridopterygidae" genus="Nilomantis" kingdom="Animalia" order="Mantodea" pageId="30" pageNumber="239" phylum="Arthropoda" rank="genus">
<emphasis box="[488,611,552,578]" italics="true" pageId="30" pageNumber="239">Nilomantis</emphasis>
</taxonomicName>
and 
<taxonomicName box="[672,775,552,578]" class="Insecta" family="Iridopterygidae" genus="Ilomantis" kingdom="Animalia" order="Mantodea" pageId="30" pageNumber="239" phylum="Arthropoda" rank="genus">
<emphasis box="[672,775,552,578]" italics="true" pageId="30" pageNumber="239">Ilomantis</emphasis>
</taxonomicName>
for potential intergeneric diagnostic characters. The shape of the dilation on the anteroventral margin of the gonoplac (GL) varies between the genera, with 
<taxonomicName box="[645,768,621,647]" class="Insecta" family="Iridopterygidae" genus="Nilomantis" kingdom="Animalia" order="Mantodea" pageId="30" pageNumber="239" phylum="Arthropoda" rank="genus">
<emphasis box="[645,768,621,647]" italics="true" pageId="30" pageNumber="239">Nilomantis</emphasis>
</taxonomicName>
possessing a rounded dilation, whereas 
<taxonomicName box="[266,369,656,682]" class="Insecta" family="Iridopterygidae" genus="Ilomantis" kingdom="Animalia" order="Mantodea" pageId="30" pageNumber="239" phylum="Arthropoda" rank="genus">
<emphasis box="[266,369,656,682]" italics="true" pageId="30" pageNumber="239">Ilomantis</emphasis>
</taxonomicName>
features a dilation that is approximately rectangular in shape (compare 
<figureCitation box="[224,296,690,717]" captionStart="Fig. 3" captionStartId="7.[98,131,1511,1533]" captionTargetBox="[147,1016,360,1484]" captionTargetId="figure@7.[147,1016,360,1485]" captionTargetPageId="7" captionText="Fig. 3. Female genitalic complex of Ilomantis shown in the lateral perspective. (A) Ilomantis thalassina; (B) Ilomantis ginsburgae sp. n. Abbreviations: AC, apical cleft; AL, apical lobe; GA, gonapophysis VIII; GL, gonoplac; MO, medial outgrowth; RTD, rectangular dilation. This figure is published in colour in the online edition of this journal, which can be accessed via http: // booksandjournals. brillonline. com / content / journals / 1876312 x." httpUri="https://zenodo.org/record/269842/files/figure.png" pageId="30" pageNumber="239">
Figs 
<number box="[281,296,691,717]" pageId="30" pageNumber="239" value="3.0">3</number>
</figureCitation>
and 
<figureCitation box="[360,378,691,717]" captionStart="Fig. 4" captionStartId="8.[106,139,1540,1562]" captionTargetBox="[155,1024,390,1513]" captionTargetId="figure@8.[155,1024,390,1515]" captionTargetPageId="8" captionText="Fig. 4. Female genitalic complex of Nilomantis shown in the lateral perspective. (A) Nilomantis floweri; (B) Nilomantis edmundsi. Abbreviations: KLP, knob-like projection; RD, round dilation. This figure is published in colour in the online edition of this journal, which can be accessed via http: // booksand journals. brillonline. com / content / journals / 1876312 x." httpUri="https://zenodo.org/record/269843/files/figure.png" pageId="30" pageNumber="239">
<number box="[360,378,691,717]" pageId="30" pageNumber="239" value="4.0">4</number>
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). Furthermore, in 
<taxonomicName box="[601,724,690,716]" class="Insecta" family="Iridopterygidae" genus="Nilomantis" kingdom="Animalia" order="Mantodea" pageId="30" pageNumber="239" phylum="Arthropoda" rank="genus">
<emphasis box="[601,724,690,716]" italics="true" pageId="30" pageNumber="239">Nilomantis</emphasis>
</taxonomicName>
the apex of gonapophysis IX (GP) is acuminate, whereas in 
<taxonomicName box="[465,568,725,751]" class="Insecta" family="Iridopterygidae" genus="Ilomantis" kingdom="Animalia" order="Mantodea" pageId="30" pageNumber="239" phylum="Arthropoda" rank="genus">
<emphasis box="[465,568,725,751]" italics="true" pageId="30" pageNumber="239">Ilomantis</emphasis>
</taxonomicName>
the apex of GP is mammiform to approximately rounded in shape (compare 
<figureCitation box="[514,583,760,787]" captionStart="Fig. 5" captionStartId="9.[98,131,1328,1350]" captionTargetBox="[147,1016,131,1303]" captionTargetId="figure@9.[147,1016,131,1303]" captionTargetPageId="9" captionText="Fig. 5. Female genitalic complex of Ilomantis shown in the ventral perspective with natural and spread positioning. (A) Ilomantis thalassina, natural; (B) Ilomantis thalassina, spread; (C) Ilomantis ginsburgae sp. n., natural; (D) Ilomantis ginsburgae sp. n., spread. Abbreviations: AC, apical cleft; AL, apical lobe; MO, medial outgrowth; MT, medial tine; VF, gonocoxa VIII; GA, gonapophysis VIII; GP, gonapophysis IX II; GL, gonoplac. This figure is published in colour in the online edition of this journal, which can be accessed via http: // booksandjournals. brillonline. com / content / journals / 1876312 x." httpUri="https://zenodo.org/record/269844/files/figure.png" pageId="30" pageNumber="239">
Figs 
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</figureCitation>
and 
<figureCitation box="[641,658,760,787]" captionStart="Fig. 6" captionStartId="10.[106,139,1438,1460]" captionTargetBox="[155,1024,131,1412]" captionTargetId="figure@10.[155,1024,131,1412]" captionTargetPageId="10" captionText="Fig. 6. Female genitalic complex of Nilomantis shown in the ventral perspective with natural and spread positioning. (A) Nilomantis floweri, natural; (B) Nilomantis floweri, spread; (C) Nilomantis edmundsi, natural; (D) Nilomantis edmundsi, spread. This figure is published in colour in the online edition of this journal, which can be accessed via http: // booksandjournals. brillonline. com / content / journals / 1876312 x." httpUri="https://zenodo.org/record/269845/files/figure.png" pageId="30" pageNumber="239">
<number box="[641,658,760,787]" pageId="30" pageNumber="239" value="6.0">6</number>
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). These character states were able to consistently and unambiguously distinguish the genera, which, when compared against external morphological characters, male genitalic characters, and geographic distribution, enabled us to resurrect 
<taxonomicName box="[521,624,864,890]" class="Insecta" family="Iridopterygidae" genus="Ilomantis" kingdom="Animalia" order="Mantodea" pageId="30" pageNumber="239" phylum="Arthropoda" rank="genus">
<emphasis box="[521,624,864,890]" italics="true" pageId="30" pageNumber="239">Ilomantis</emphasis>
</taxonomicName>
as a valid genus with confidence.
</paragraph>
<paragraph blockId="30.[106,1074,517,1653]" pageId="30" pageNumber="239">
We likewise examined the female genital complexes of 
<taxonomicName box="[748,880,898,924]" class="Insecta" family="Iridopterygidae" genus="Tropidomantis" kingdom="Animalia" order="Mantodea" pageId="30" pageNumber="239" phylum="Arthropoda" rank="species" species="thalassina">
<emphasis box="[748,880,898,924]" italics="true" pageId="30" pageNumber="239">I. thalassina</emphasis>
</taxonomicName>
and 
<taxonomicName box="[935,1073,898,924]" class="Insecta" family="Iridopterygidae" genus="Ilomantis" kingdom="Animalia" order="Mantodea" pageId="30" pageNumber="239" phylum="Arthropoda" rank="species" species="ginsburgae" status="sp.n.">
<emphasis box="[935,1073,898,924]" italics="true" pageId="30" pageNumber="239">I. ginsburgae</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[106,160,934,960]" pageId="30" pageNumber="239" rank="species">sp.n.</taxonomicNameLabel>
as a site for potential interspecific diagnostic characters. We found that gonapophysis VIII (GA) and GL feature compelling interspecific character states that proved successful in delimiting the species. Generally, 
<taxonomicName box="[654,873,1002,1028]" class="Insecta" family="Iridopterygidae" genus="Ilomantis" kingdom="Animalia" order="Mantodea" pageId="30" pageNumber="239" phylum="Arthropoda" rank="species" species="thalassina">
<emphasis box="[654,873,1002,1028]" italics="true" pageId="30" pageNumber="239">Ilomantis thalassina</emphasis>
</taxonomicName>
features broader, more robust gonapophyses and gonoplacs than 
<taxonomicName box="[620,752,1037,1063]" class="Insecta" family="Iridopterygidae" genus="Ilomantis" kingdom="Animalia" order="Mantodea" pageId="30" pageNumber="239" phylum="Arthropoda" rank="species" species="ginsburgae" status="sp.n.">
<emphasis box="[620,752,1037,1063]" italics="true" pageId="30" pageNumber="239">I. ginsburgae</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[758,810,1038,1064]" pageId="30" pageNumber="239" rank="species">sp.n.</taxonomicNameLabel>
(compare 
<figureCitation box="[925,989,1037,1063]" captionStart="Fig. 3" captionStartId="7.[98,131,1511,1533]" captionTargetBox="[147,1016,360,1484]" captionTargetId="figure@7.[147,1016,360,1485]" captionTargetPageId="7" captionText="Fig. 3. Female genitalic complex of Ilomantis shown in the lateral perspective. (A) Ilomantis thalassina; (B) Ilomantis ginsburgae sp. n. Abbreviations: AC, apical cleft; AL, apical lobe; GA, gonapophysis VIII; GL, gonoplac; MO, medial outgrowth; RTD, rectangular dilation. This figure is published in colour in the online edition of this journal, which can be accessed via http: // booksandjournals. brillonline. com / content / journals / 1876312 x." httpUri="https://zenodo.org/record/269842/files/figure.png" pageId="30" pageNumber="239">
Figs 
<number box="[975,989,1037,1063]" pageId="30" pageNumber="239" value="3.0">3</number>
</figureCitation>
and 
<figureCitation box="[1043,1059,1037,1064]" captionStart="Fig. 5" captionStartId="9.[98,131,1328,1350]" captionTargetBox="[147,1016,131,1303]" captionTargetId="figure@9.[147,1016,131,1303]" captionTargetPageId="9" captionText="Fig. 5. Female genitalic complex of Ilomantis shown in the ventral perspective with natural and spread positioning. (A) Ilomantis thalassina, natural; (B) Ilomantis thalassina, spread; (C) Ilomantis ginsburgae sp. n., natural; (D) Ilomantis ginsburgae sp. n., spread. Abbreviations: AC, apical cleft; AL, apical lobe; MO, medial outgrowth; MT, medial tine; VF, gonocoxa VIII; GA, gonapophysis VIII; GP, gonapophysis IX II; GL, gonoplac. This figure is published in colour in the online edition of this journal, which can be accessed via http: // booksandjournals. brillonline. com / content / journals / 1876312 x." httpUri="https://zenodo.org/record/269844/files/figure.png" pageId="30" pageNumber="239">
<number box="[1043,1059,1037,1064]" pageId="30" pageNumber="239" value="5.0">5</number>
</figureCitation>
). Within 
<taxonomicName box="[196,328,1072,1098]" class="Insecta" family="Iridopterygidae" genus="Ilomantis" kingdom="Animalia" order="Mantodea" pageId="30" pageNumber="239" phylum="Arthropoda" rank="species" species="thalassina">
<emphasis box="[196,328,1072,1098]" italics="true" pageId="30" pageNumber="239">I. thalassina</emphasis>
</taxonomicName>
, GA features a pronounced medial outgrowth (MO); GA anterodorsal margin is moderately sclerotized, with the sclerotized region concluding just posterior to the “heel” of GA. The apical cleft (AC) of GL is distinctly divergent, the resultant lobes blunt apically; the ventral and dorsal margins of GL are rugose, sculpturate (see 
<figureCitation box="[232,297,1210,1237]" captionStart="Fig. 3" captionStartId="7.[98,131,1511,1533]" captionTargetBox="[147,1016,360,1484]" captionTargetId="figure@7.[147,1016,360,1485]" captionTargetPageId="7" captionText="Fig. 3. Female genitalic complex of Ilomantis shown in the lateral perspective. (A) Ilomantis thalassina; (B) Ilomantis ginsburgae sp. n. Abbreviations: AC, apical cleft; AL, apical lobe; GA, gonapophysis VIII; GL, gonoplac; MO, medial outgrowth; RTD, rectangular dilation. This figure is published in colour in the online edition of this journal, which can be accessed via http: // booksandjournals. brillonline. com / content / journals / 1876312 x." httpUri="https://zenodo.org/record/269842/files/figure.png" pageId="30" pageNumber="239">
Fig. 
<number box="[284,297,1211,1237]" pageId="30" pageNumber="239" value="3.0">3</number>
</figureCitation>
A). However, in 
<taxonomicName box="[491,631,1210,1236]" class="Insecta" family="Iridopterygidae" genus="Ilomantis" kingdom="Animalia" order="Mantodea" pageId="30" pageNumber="239" phylum="Arthropoda" rank="species" species="ginsburgae" status="sp.n.">
<emphasis box="[491,631,1210,1236]" italics="true" pageId="30" pageNumber="239">I. ginsburgae</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[640,696,1211,1237]" pageId="30" pageNumber="239" rank="species">sp.n.</taxonomicNameLabel>
, MO of GA is reduced; and GA anterodorsal margin is less sclerotized than in 
<taxonomicName box="[626,758,1245,1271]" class="Insecta" family="Iridopterygidae" genus="Ilomantis" kingdom="Animalia" order="Mantodea" pageId="30" pageNumber="239" phylum="Arthropoda" rank="species" species="thalassina">
<emphasis box="[626,758,1245,1271]" italics="true" pageId="30" pageNumber="239">I. thalassina</emphasis>
</taxonomicName>
, with the region of sclerotization terminating just prior to the apical lobes (AL). The AC of GL is significantly reduced, with the lobe of the dorsal margin appearing to converge with the ventral lobe, resulting in a slightly notched, acuminate apex; GL ventral and dorsal margins are smooth (see 
<figureCitation box="[288,352,1384,1410]" captionStart="Fig. 3" captionStartId="7.[98,131,1511,1533]" captionTargetBox="[147,1016,360,1484]" captionTargetId="figure@7.[147,1016,360,1485]" captionTargetPageId="7" captionText="Fig. 3. Female genitalic complex of Ilomantis shown in the lateral perspective. (A) Ilomantis thalassina; (B) Ilomantis ginsburgae sp. n. Abbreviations: AC, apical cleft; AL, apical lobe; GA, gonapophysis VIII; GL, gonoplac; MO, medial outgrowth; RTD, rectangular dilation. This figure is published in colour in the online edition of this journal, which can be accessed via http: // booksandjournals. brillonline. com / content / journals / 1876312 x." httpUri="https://zenodo.org/record/269842/files/figure.png" pageId="30" pageNumber="239">
Fig. 
<number box="[338,352,1384,1410]" pageId="30" pageNumber="239" value="3.0">3</number>
</figureCitation>
B).
</paragraph>
<paragraph blockId="30.[106,1074,517,1653]" lastBlockId="31.[98,1067,134,1652]" lastPageId="31" lastPageNumber="240" pageId="30" pageNumber="239">
Although the number of specimens were limited, we examined intraspecific variation within 
<specimenCount box="[240,345,1453,1479]" pageId="30" pageNumber="239" type="female">
<number box="[240,255,1453,1479]" pageId="30" pageNumber="239" value="2.0">2</number>
females
</specimenCount>
of 
<taxonomicName box="[382,514,1453,1479]" class="Insecta" family="Iridopterygidae" genus="Ilomantis" kingdom="Animalia" order="Mantodea" pageId="30" pageNumber="239" phylum="Arthropoda" rank="species" species="thalassina">
<emphasis box="[382,514,1453,1479]" italics="true" pageId="30" pageNumber="239">I. thalassina</emphasis>
</taxonomicName>
and 
<specimenCount box="[570,674,1453,1479]" pageId="30" pageNumber="239" type="female">
<number box="[570,585,1453,1479]" pageId="30" pageNumber="239" value="3.0">3</number>
females
</specimenCount>
of 
<taxonomicName box="[712,850,1453,1479]" class="Insecta" family="Iridopterygidae" genus="Ilomantis" kingdom="Animalia" order="Mantodea" pageId="30" pageNumber="239" phylum="Arthropoda" rank="species" species="ginsburgae" status="sp.n.">
<emphasis box="[712,850,1453,1479]" italics="true" pageId="30" pageNumber="239">I. ginsburgae</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[858,913,1454,1480]" pageId="30" pageNumber="239" rank="species">sp.n.</taxonomicNameLabel>
; variation was most frequently observed in the apical lobe (AL) when viewed from the lateral perspective, ranging from rounded, to blunted, to angled in shape. However, this might be due to AL being a membranous structure, which can be actively or passively moved by an insect, and which might also slightly change shape when submerged within ethanol. Furthermore, AL is a lobe-like apical dilation of GA that is folded toward the external surface along the ventral margin, which could lend AL a varied appearance. We found that the overall shape of the female genitalia was more uniform in comparison to the intraspecific variation of character states observed in male genitalia, specifically in the overall length and width of the apical process (ap) of L 
<number box="[741,756,239,265]" pageId="31" pageNumber="240" value="4.0">4</number>
B, as well as the shape and sclerotization of the anterior margin of ap (see 
<figureCitation box="[616,695,273,299]" captionStart="Fig. 10" captionStartId="14.[106,139,1247,1269]" captionTargetBox="[155,1024,131,1220]" captionTargetId="figure@14.[155,1024,131,1222]" captionTargetPageId="14" captionText="Fig. 10. Male genitalic complex of Ilomantis ginsburgae sp. n. (A) Right phallomere, ventral sclerotization of the left phallomeric complex, and dorsal (i. e., “ left ”) sclerotization of the left phallomeric complex (from left to right, respectively); (B, C, D) examples of shape and sclerotization variability present within the anterior process (ap) of L 4 B. This figure is published in colour in the online edition of this journal, which can be accessed via http: // booksandjournals. brillonline. com / content / journals / 1876312 x." httpUri="https://zenodo.org/record/269849/files/figure.png" pageId="31" pageNumber="240">
Fig. 
<number box="[665,695,273,299]" pageId="31" pageNumber="240" value="10.0">10</number>
</figureCitation>
) of the one 
<taxonomicName box="[828,958,273,299]" class="Insecta" family="Iridopterygidae" genus="Ilomantis" kingdom="Animalia" order="Mantodea" pageId="31" pageNumber="240" phylum="Arthropoda" rank="species" species="thalassina">
<emphasis box="[828,958,273,299]" italics="true" pageId="31" pageNumber="240">I. thalassina</emphasis>
</taxonomicName>
male and four 
<taxonomicName box="[153,292,308,334]" class="Insecta" family="Iridopterygidae" genus="Ilomantis" kingdom="Animalia" order="Mantodea" pageId="31" pageNumber="240" phylum="Arthropoda" rank="species" species="ginsburgae" status="sp.n.">
<emphasis box="[153,292,308,334]" italics="true" pageId="31" pageNumber="240">I. ginsburgae</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[299,353,308,334]" pageId="31" pageNumber="240" rank="species">sp.n.</taxonomicNameLabel>
males examined.
</paragraph>
<paragraph blockId="31.[98,1067,134,1652]" pageId="31" pageNumber="240">
An examination of the female genital complexes of 
<taxonomicName box="[708,910,342,368]" class="Insecta" family="Iridopterygidae" genus="Nilomantis" kingdom="Animalia" order="Mantodea" pageId="31" pageNumber="240" phylum="Arthropoda" rank="species" species="floweri">
<emphasis box="[708,910,342,368]" italics="true" pageId="31" pageNumber="240">Nilomantis floweri</emphasis>
</taxonomicName>
(
<emphasis box="[927,942,343,368]" italics="true" pageId="31" pageNumber="240">n</emphasis>
= 
<number box="[960,970,343,369]" pageId="31" pageNumber="240" value="1.0">1</number>
) and 
<taxonomicName class="Insecta" family="Iridopterygidae" genus="Nilomantis" kingdom="Animalia" order="Mantodea" pageId="31" pageNumber="240" phylum="Arthropoda" rank="species" species="edmundsi">
<emphasis italics="true" pageId="31" pageNumber="240">N. edmundsi</emphasis>
</taxonomicName>
(
<emphasis box="[218,233,378,403]" italics="true" pageId="31" pageNumber="240">n</emphasis>
= 
<number box="[252,262,377,403]" pageId="31" pageNumber="240" value="1.0">1</number>
) led to the addition of new characters with which to delimit the species. In particular, 
<taxonomicName box="[250,359,412,438]" class="Insecta" family="Iridopterygidae" genus="Nilomantis" kingdom="Animalia" order="Mantodea" pageId="31" pageNumber="240" phylum="Arthropoda" rank="species" species="floweri">
<emphasis box="[250,359,412,438]" italics="true" pageId="31" pageNumber="240">N. floweri</emphasis>
</taxonomicName>
features a gonocoxa VIII (VF) with a rounded internal margin; which, in 
<taxonomicName box="[212,352,446,472]" class="Insecta" family="Iridopterygidae" genus="Nilomantis" kingdom="Animalia" order="Mantodea" pageId="31" pageNumber="240" phylum="Arthropoda" rank="species" species="edmundsi">
<emphasis box="[212,352,446,472]" italics="true" pageId="31" pageNumber="240">N. edmundsi</emphasis>
</taxonomicName>
, is approximately angular (see 
<figureCitation box="[699,766,446,473]" captionStart="Fig. 6" captionStartId="10.[106,139,1438,1460]" captionTargetBox="[155,1024,131,1412]" captionTargetId="figure@10.[155,1024,131,1412]" captionTargetPageId="10" captionText="Fig. 6. Female genitalic complex of Nilomantis shown in the ventral perspective with natural and spread positioning. (A) Nilomantis floweri, natural; (B) Nilomantis floweri, spread; (C) Nilomantis edmundsi, natural; (D) Nilomantis edmundsi, spread. This figure is published in colour in the online edition of this journal, which can be accessed via http: // booksandjournals. brillonline. com / content / journals / 1876312 x." httpUri="https://zenodo.org/record/269845/files/figure.png" pageId="31" pageNumber="240">
Fig. 
<number box="[749,766,446,473]" pageId="31" pageNumber="240" value="6.0">6</number>
</figureCitation>
). From LH, VF is without projection in 
<taxonomicName box="[258,371,481,507]" class="Insecta" family="Iridopterygidae" genus="Nilomantis" kingdom="Animalia" order="Mantodea" pageId="31" pageNumber="240" phylum="Arthropoda" rank="species" species="floweri">
<emphasis box="[258,371,481,507]" italics="true" pageId="31" pageNumber="240">N. floweri</emphasis>
</taxonomicName>
, whereas VF features a distinct knob-like projection that is directed dorsad (see 
<figureCitation box="[328,394,516,542]" captionStart="Fig. 4" captionStartId="8.[106,139,1540,1562]" captionTargetBox="[155,1024,390,1513]" captionTargetId="figure@8.[155,1024,390,1515]" captionTargetPageId="8" captionText="Fig. 4. Female genitalic complex of Nilomantis shown in the lateral perspective. (A) Nilomantis floweri; (B) Nilomantis edmundsi. Abbreviations: KLP, knob-like projection; RD, round dilation. This figure is published in colour in the online edition of this journal, which can be accessed via http: // booksand journals. brillonline. com / content / journals / 1876312 x." httpUri="https://zenodo.org/record/269843/files/figure.png" pageId="31" pageNumber="240">
Fig. 
<number box="[377,394,516,542]" pageId="31" pageNumber="240" value="4.0">4</number>
</figureCitation>
). The apical lobe (AL) of 
<taxonomicName box="[683,792,516,542]" class="Insecta" family="Iridopterygidae" genus="Nilomantis" kingdom="Animalia" order="Mantodea" pageId="31" pageNumber="240" phylum="Arthropoda" rank="species" species="floweri">
<emphasis box="[683,792,516,542]" italics="true" pageId="31" pageNumber="240">N. floweri</emphasis>
</taxonomicName>
is heavily ciliated, while in 
<taxonomicName box="[128,269,550,576]" class="Insecta" family="Iridopterygidae" genus="Nilomantis" kingdom="Animalia" order="Mantodea" pageId="31" pageNumber="240" phylum="Arthropoda" rank="species" species="edmundsi">
<emphasis box="[128,269,550,576]" italics="true" pageId="31" pageNumber="240">N. edmundsi</emphasis>
</taxonomicName>
AL cilia were not observed across the majority of its surface (see 
<figureCitation box="[999,1066,550,577]" captionStart="Fig. 4" captionStartId="8.[106,139,1540,1562]" captionTargetBox="[155,1024,390,1513]" captionTargetId="figure@8.[155,1024,390,1515]" captionTargetPageId="8" captionText="Fig. 4. Female genitalic complex of Nilomantis shown in the lateral perspective. (A) Nilomantis floweri; (B) Nilomantis edmundsi. Abbreviations: KLP, knob-like projection; RD, round dilation. This figure is published in colour in the online edition of this journal, which can be accessed via http: // booksand journals. brillonline. com / content / journals / 1876312 x." httpUri="https://zenodo.org/record/269843/files/figure.png" pageId="31" pageNumber="240">
Figs 
<number box="[1051,1066,551,577]" pageId="31" pageNumber="240" value="4.0">4</number>
</figureCitation>
and 
<figureCitation box="[147,164,585,612]" captionStart="Fig. 6" captionStartId="10.[106,139,1438,1460]" captionTargetBox="[155,1024,131,1412]" captionTargetId="figure@10.[155,1024,131,1412]" captionTargetPageId="10" captionText="Fig. 6. Female genitalic complex of Nilomantis shown in the ventral perspective with natural and spread positioning. (A) Nilomantis floweri, natural; (B) Nilomantis floweri, spread; (C) Nilomantis edmundsi, natural; (D) Nilomantis edmundsi, spread. This figure is published in colour in the online edition of this journal, which can be accessed via http: // booksandjournals. brillonline. com / content / journals / 1876312 x." httpUri="https://zenodo.org/record/269845/files/figure.png" pageId="31" pageNumber="240">
<number box="[147,164,585,612]" pageId="31" pageNumber="240" value="6.0">6</number>
</figureCitation>
). Lastly, the ventral margin of the gonoplac (GL) is rugose and the dorsal margin is relatively smooth in 
<taxonomicName box="[358,468,620,646]" class="Insecta" family="Iridopterygidae" genus="Nilomantis" kingdom="Animalia" order="Mantodea" pageId="31" pageNumber="240" phylum="Arthropoda" rank="species" species="floweri">
<emphasis box="[358,468,620,646]" italics="true" pageId="31" pageNumber="240">N. floweri</emphasis>
</taxonomicName>
, whereas both the ventral and dorsal margins of GL in 
<taxonomicName box="[129,269,654,680]" class="Insecta" family="Iridopterygidae" genus="Nilomantis" kingdom="Animalia" order="Mantodea" pageId="31" pageNumber="240" phylum="Arthropoda" rank="species" species="edmundsi">
<emphasis box="[129,269,654,680]" italics="true" pageId="31" pageNumber="240">N. edmundsi</emphasis>
</taxonomicName>
are relatively smooth (see 
<figureCitation box="[567,634,654,681]" captionStart="Fig. 4" captionStartId="8.[106,139,1540,1562]" captionTargetBox="[155,1024,390,1513]" captionTargetId="figure@8.[155,1024,390,1515]" captionTargetPageId="8" captionText="Fig. 4. Female genitalic complex of Nilomantis shown in the lateral perspective. (A) Nilomantis floweri; (B) Nilomantis edmundsi. Abbreviations: KLP, knob-like projection; RD, round dilation. This figure is published in colour in the online edition of this journal, which can be accessed via http: // booksand journals. brillonline. com / content / journals / 1876312 x." httpUri="https://zenodo.org/record/269843/files/figure.png" pageId="31" pageNumber="240">
Fig. 
<number box="[617,634,655,681]" pageId="31" pageNumber="240" value="4.0">4</number>
</figureCitation>
).
</paragraph>
<paragraph blockId="31.[98,1067,134,1652]" pageId="31" pageNumber="240">
The examined specimens were collected between 
<number box="[709,753,689,715]" pageId="31" pageNumber="240" value="116.0">116</number>
to 
<number box="[800,829,689,715]" pageId="31" pageNumber="240" value="13.0">13</number>
years ago, with the majority collected around 
<number box="[408,437,724,751]" pageId="31" pageNumber="240" value="50.0">50</number>
years ago. With material of this age, desiccation and damage from drawer movement, museum environmental conditions (e.g., fluctuations in temperature or humidity), and specimen handling may occur. While we took great pains to ensure that the entire female and male genital complex of each specimen was removed intact, we were unable to isolate the spermatheca, vestibular floor, and the laterosternal shelf of the female genitalia. This might be due to the fact that the genitalia of these species are membranous, and when coupled with specimen age, more susceptible to being destroyed over time or while submerged in potassium hydroxide, a solution frequently used to clear insect genitalia. While we were not able to access these structures within the female genital complex of 
<taxonomicName box="[635,737,1036,1062]" class="Insecta" family="Iridopterygidae" genus="Ilomantis" kingdom="Animalia" order="Mantodea" pageId="31" pageNumber="240" phylum="Arthropoda" rank="genus">
<emphasis box="[635,737,1036,1062]" italics="true" pageId="31" pageNumber="240">Ilomantis</emphasis>
</taxonomicName>
and 
<taxonomicName box="[793,915,1036,1062]" class="Insecta" family="Iridopterygidae" genus="Nilomantis" kingdom="Animalia" order="Mantodea" pageId="31" pageNumber="240" phylum="Arthropoda" rank="genus">
<emphasis box="[793,915,1036,1062]" italics="true" pageId="31" pageNumber="240">Nilomantis</emphasis>
</taxonomicName>
, we were still able to find characters on the gonapophyses, gonoplacs, and gonocoxae, which unambiguously delimited the genera and the species.
</paragraph>
<paragraph blockId="31.[98,1067,134,1652]" pageId="31" pageNumber="240">
Our investigation of mantodean female genitalia revealed a novel character system that successfully resurrected 
<taxonomicName box="[417,518,1174,1200]" class="Insecta" family="Iridopterygidae" genus="Ilomantis" kingdom="Animalia" order="Mantodea" pageId="31" pageNumber="240" phylum="Arthropoda" rank="genus">
<emphasis box="[417,518,1174,1200]" italics="true" pageId="31" pageNumber="240">Ilomantis</emphasis>
</taxonomicName>
as a valid genus, delimited the species contained within both 
<taxonomicName box="[237,338,1209,1235]" class="Insecta" family="Iridopterygidae" genus="Ilomantis" kingdom="Animalia" order="Mantodea" pageId="31" pageNumber="240" phylum="Arthropoda" rank="genus">
<emphasis box="[237,338,1209,1235]" italics="true" pageId="31" pageNumber="240">Ilomantis</emphasis>
</taxonomicName>
and 
<taxonomicName box="[393,514,1209,1235]" class="Insecta" family="Iridopterygidae" genus="Nilomantis" kingdom="Animalia" order="Mantodea" pageId="31" pageNumber="240" phylum="Arthropoda" rank="genus">
<emphasis box="[393,514,1209,1235]" italics="true" pageId="31" pageNumber="240">Nilomantis</emphasis>
</taxonomicName>
(i.e., 
<emphasis box="[577,828,1209,1235]" italics="true" pageId="31" pageNumber="240">
<taxonomicName box="[577,710,1209,1235]" class="Insecta" family="Iridopterygidae" genus="Ilomantis" kingdom="Animalia" order="Mantodea" pageId="31" pageNumber="240" phylum="Arthropoda" rank="species" species="thalassina">I. thalassina</taxonomicName>
, 
<taxonomicName box="[720,828,1209,1235]" class="Insecta" family="Iridopterygidae" genus="Nilomantis" kingdom="Animalia" order="Mantodea" pageId="31" pageNumber="240" phylum="Arthropoda" rank="species" species="floweri">N. floweri</taxonomicName>
</emphasis>
and 
<taxonomicName box="[883,1022,1209,1235]" class="Insecta" family="Iridopterygidae" genus="Nilomantis" kingdom="Animalia" order="Mantodea" pageId="31" pageNumber="240" phylum="Arthropoda" rank="species" species="edmundsi">
<emphasis box="[883,1022,1209,1235]" italics="true" pageId="31" pageNumber="240">N. edmundsi</emphasis>
</taxonomicName>
), as well as described a new 
<collectingCountry box="[360,464,1244,1270]" name="Madagascar" pageId="31" pageNumber="240">Malagasy</collectingCountry>
praying mantis species (i.e., 
<taxonomicName box="[782,920,1244,1270]" class="Insecta" family="Iridopterygidae" genus="Ilomantis" kingdom="Animalia" order="Mantodea" pageId="31" pageNumber="240" phylum="Arthropoda" rank="species" species="ginsburgae" status="sp.n.">
<emphasis box="[782,920,1244,1270]" italics="true" pageId="31" pageNumber="240">I. ginsburgae</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[926,982,1244,1270]" pageId="31" pageNumber="240" rank="species">sp.n.</taxonomicNameLabel>
). These results underscore the utility of female genitalia in 
<taxonomicName box="[691,806,1278,1304]" class="Insecta" family="Mantodea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Mantodea" pageId="31" pageNumber="240" phylum="Arthropoda" rank="order">Mantodea</taxonomicName>
taxonomy, and subsequently demonstrate the need for thorough examination of female genital complexes as sites of taxonomic characters. Furthermore, by describing both male and female genitalic characters (when available), the ability to diagnose females in externally homogenous genera might be resolved within 
<taxonomicName box="[638,757,1417,1443]" class="Insecta" family="Mantodea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Mantodea" pageId="31" pageNumber="240" phylum="Arthropoda" rank="order">Mantodea</taxonomicName>
. For this study, traditional taxonomic data, such as external morphology, male genitalic characters, and geographic distribution, supported the results of the investigation of the female genitalic character system. As such, it is our hope that future taxonomic descriptions of praying mantises assess female genitalic characters in addition to traditional taxonomic treatments. A more thorough investigation of female genitalia across a wide taxon sampling might provide a new motif from which to build a novel taxonomic character system.
</paragraph>
</subSubSection>
</treatment>
</document>