Leveraging female genitalic characters for generic and species delimitation in Nilomantis Werner, 1907 and Ilomantis Giglio-Tos, 1915 (Mantodea, Nilomantinae)

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Praying mantis species description and higher-level taxonomy has historically relied on external morphological characters, male genitalic characters, and geographic distribution. For the first time, female genitalic characters were used to delimit genus- and species-level boundaries using two genera with a complex taxonomic history, NilomantisWerner, 1907 and IlomantisGiglio-Tos, 1915. We present comprehensive morphological evidence for the validity of Ilomantis, currently a junior synonym of Nilomantis and describe a new species, Ilomantis ginsburgae sp.n. We provide a key to the species of Nilomantis and Ilomantis based on genitalic and external morphological character states. High-resolution images, illustrations of morphological characters, and measurement data are also presented. We conclude that female genital characters have significant utility in praying mantis taxonomy and should be incorporated as an additional data source to complement traditional taxonomic and genetic data.

Leveraging female genitalic characters for generic and species delimitation in Nilomantis Werner, 1907 and Ilomantis Giglio-Tos, 1915 (Mantodea, Nilomantinae)

in Insect Systematics & Evolution

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References

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Figures

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    Ilomantis thalassina. (A) Female, dorsal habitus; (B) female, ventral habitus; (C) male, dorsal habitus; (D) male, ventral habitus. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1876312x.

  • View in gallery

    Ilomantis ginsburgae sp.n. (A) Holotype female, dorsal habitus; (B) holotype female, ventral habitus; (C) allotype male, dorsal habitus; (D) allotype male, ventral habitus. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1876312x.

  • View in gallery

    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/1876312x.

  • View in gallery

    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://booksandjournals.brillonline.com/content/journals/1876312x.

  • View in gallery

    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/1876312x.

  • View in gallery

    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/1876312x.

  • View in gallery

    Species distribution map for Nilomantis and Ilomantis species. Examined Nilomantis and Ilomantis specimens, as well as additional Nilomantis taxa listed in Roy & Leston (1975), have been incorporated. Collecting localities without coordinates were georeferenced using Google Earth. Ilomantis species are distributed within Madagascar; Nilomantis species are distributed across the Sahara-Sahel region of Africa, including Ghana, the Central African Republic, and Saudi Arabia. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1876312x.

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    Illustrations of anterior perspective of the Ilomantis and Nilomantis head (scale bar=1 mm). (A) Ilomantis thalassina, female; (B) Ilomantis thalassina, male; (C) Ilomantis ginsburgae sp.n., female; (D) Ilomantis ginsburgae sp.n., male; (E) Nilomantis floweri, female; (F) Nilomantis floweri, male. (B, D, E, F) Ilomantis species feature a cosinusoidal vertical margin, whereas Nilomantis species feature a relatively straight vertical margin. Gray markings on the ocellar tubercle and within the corridors of the frontal sulci represent the presence of reddish-brown markings. LF, lower frons.

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    Male genitalic complex of Ilomantis thalassina. Abbreviations: an, anterior apodeme; ap, anterior process; bm, right arm; L1, L2, L4B, a sub-sclerite of the dorsal sclerotization of the left phallomeric complex; L4A, the ventral sclerotization of the left phallomeric complex; loa, lobo membranoso; paa, apical process; pda, processo distale; pia, piastra ventrale; pva, processo ventrale sclerificato; R1, a sub-sclerite of the right phallomere; R3, a sub-sclerite of the right phallomere. Red lettering indicates structures; black lettering indicates characters. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1876312x.

  • View in gallery

    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 L4B. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1876312x.

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    Illustrations of dorsal perspective of the Ilomantis pronotum (scale bar = 1 mm). (A) Ilomantis thalassina, female; (B) Ilomantis thalassina, male; (C) Ilomantis ginsburgae sp.n., female; (D) Ilomantis ginsburgae sp.n., male. Ilomantis thalassina exhibits a medial keel that fully traverses the pronotum (A and B); Ilomantis ginsburgae sp.n. exhibits a medial keel that originates in the mid-prozone and concludes at the posterior pronotal margin (C and D). MK, medial keel.

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    Example of yellow stripe that originates in the posterior half of the ocellar tubercle and traverses the thorax of both Nilomantis and Ilomantis species. Species shown: Ilomantis ginsburgae sp.n., male. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1876312x.

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    Illustrations of ventral perspective of the Ilomantis cervix (scale bar = 1 mm). (A) Ilomantis thalassina, female; (B) Ilomantis thalassina, male; (C) Ilomantis ginsburgae sp.n., female; (D) Ilomantis ginsburgae sp.n., male. The postcervical plate and exposed thoracic membrane are contained within the boundaries of the intercervical sclerites and the episternum, with either a broad (Ilomantis thalassina, A and B) or narrow (Ilomantis ginsburgae sp.n., C and D) hexagonal shape. Abbreviations: eps, episternum; etm, exposed thoracic membrane; ics, intercervical sclerite; lcs, lateral cervical sclerite; pcpl, postcervical plate; tss, T-shaped sclerite.

  • View in gallery

    Oblique ventral view of the furcasternal tubercle (FT) located medially at the base of the T-shaped sclerite, posterior to the prothoracic coxae. Arrow indicates FT. Species shown: Nilomantis floweri, male. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1876312x.

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