Listroscelidinae

Chamorro-Rengifo, Juliana & Lopes-Andrade, Cristiano, 2014, The phallus in Tettigoniidae (Insecta: Orthoptera: Ensifera): revision of morphology and terminology, and discussion on its taxonomic importance and evolution, Zootaxa 3815 (2), pp. 151-199 : 181-183

Listroscelidinae . Incertae sedis. Gen. 2 sp. ( Figs 28–29 , 107 ).

Meconematinae . Aprosphylini . Pseudosaga maraisi Naskrecki. Meconematini . Xiphidiopsis Redtenbacher : X. amnicola Gorochov ; X. platycerca Bey-Bienko. Xizicus fascipes (Bey-Bienko) . Incertae sedis. Exoteratura kerinci (Gorochov) , with the distal part of the phallus bearing a pair of pubescent and weakly sclerotized areas. Grigoriora Gorochov ; G. beybienkoi Gorochov ; G. beybienkoi Gorochov.

Phaneropterinae . Ducetiini . Ducetia borealis Gorochov & Kang ; Prohimerta sichuanensis Gorochov & Kang. Incertae sedis. Gen. 3 sp. ( Figs 38–39 , 112 ); Gen. 4 sp. ( Figs 40–41 , 113 ); Gen. 5 sp. ( Figs 42–43 , 114 ). Phyllopterae group. Itarissa sp. ( Fig. 110 ). Steirodontini . Stilpnochlora acanthonotum Nickle ; Stilpnochlora quadrata (Scudder) ; Stilpnochlora azteca (Saussure,); Stilpnochlora thoracica (Serville) .

Phyllophorinae . Phyllophorella queenslandica Rentz, Su & Ueshima ; Siliquofera grandis (Blanchard) , in which cerci are comparatively longer.

Pseudophyllinae . Polyancistrini . S. alieniphaga ( Figs 64–65 , 125 ). Teleutiini . Chibchella nigrospecula Montealegre-Z. & Morris; Stetharasa exarmata Montealegre-Z. & Morris. Teleutias sp. ( Figs 70–71 , 128 ). Pleminiini . Acanthodis sp. ( Figs 58–59 , 122 ).

Zaprochilinae . Kawanaphila Rentz. Zaprochilus Caudell.

Second hypothetical morphological correlation

Males with curved or angulated cerci resembling hooks, or cerci with secondary components such as inner spines, usually accompanied by sclerite(s) TS. The sclerite(s) TS can be two longitudinal sclerotized bars, an entire TS with a bifurcated apex or a transversal bridge with two free projections. Such combination occurs in:

Conocephalinae . Agraeciini . Agraecia , all species. Anelytra Redtenbacher : A. compressa Shi & Qiu ; A. jinghonga Shi & Qiu ; A. spinia Shi & Qiu. Anthracites nakanaiensis Naskrecki & Rentz ; Dectinomima Caudell , all species. Eschatoceras sp. ( Figs 10–11 , 97 ). Gonatacanthus gahavisuka Naskrecki & Rentz ; Ingrischia macrocephala Naskrecki & Rentz. Microsalomona Karny , all species. Pandanagraecia Naskrecki & Rentz , all species. Philmontis Willemse : P. lobatus Naskrecki & Rentz ; P. nigrofasciatus Willemse ; Pseudonicsara fascifrons Naskrecki & Rentz ; Salomona richardsi Naskrecki & Rentz ; Scytocera smaragdifrons Naskrecki & Rentz ; Spinisternum castaneipictus Willemse ; Sphyrometopa Carl , all species. Trichophallus tricuspis Naskrecki & Rentz ; Uchuca Giglio-Tos , possibly all species. U. ferreirai ( Figs 12–13 , 98 ). Conocephalini . Conocephalus , possibly all species. Conocephalus sp.1 ( Figs 6–9 , 99–101 ). Copiphorini . Borinquenula Walker & Gurney , all species. Neoconocephalus Karny , possibly all species, Neoconocephalus spp. ( Figs 20–21 , 103–104 ). Panacanthus Walker , possibly all species.

Bradyporinae . Ephippigerini . Ephippiger ephippiger (Fiebig) . Bradyporus Charpentier , possibly all species.

Listroscelidinae . Terpandrini . Neobarrettia Rehn : N. hakippah Cohn ; N. sinaloae (Rehn & Hebard) . Incertae sedis. C. viridis ( Figs 22–25 , 105 ). Listroscelis Serville all species. Monocerophora Walker , possibly all species. Gen. 1 sp. ( Figs 26–27 , 106 ).

Meconematinae . Meconematini . Decma Gorochov : D. elefani Gorochov ; D. thai Gorochov. Kuzicus Gorochov , apparently all species. Pseudokuzicus tamdao Gorochov , in which the single ti is bifurcated at apex. Teratura darevskyi Gorochov , with cerci similar to those of Xiphidiopsis but bearing sclerites TS.

Phaneropterinae . Ducetiini . Abaxisotima multipunctata (Kang & Yang) . Paraducetia paracruciata Gorochov & Kang. Incertae sedis. Brinckiella Chopard , Paraxantia Liu & Kang. Stylomolpa angustipennis Karny. Mirollini. Mirollia rostellum Gorochov. Odonturini . Cohnia andeana (Hebard) . Viadanini . Anaulacomera sp. ( Figs 34–37 , 111 ).

Tettigoniinae . Drymadusini . Atlanticus testaceus (Scudder) . Delodusa humeralis (Uvarov) ; Exodrymadusa inornata (Uvarov) , a monospecific genus. Leptodusa harzi (Karabag) . Mixodusa Stolyarov , possibly all species. Novadrymadusa Dermirsoy, Salman & Sevgili , two species included in the genus. Phytodrymadusa Ramme , possibly all species. Scotodrymadusa philbyi (Uvarov) . Glyphonotini . Cyrtophyllicus chlorum Hebard. Nedubini . Aglaothorax Caudell : A. diminutiva (Rentz & Birchim A. morsei (Caudell) . Neduba Walker : N. castanea (Scudder) ; N. macneilli Rentz & Birchim ; N. sierranus (Rehn & Hebard) . Platycleidini . Anterastes Brunner von Wattenwyl , all species. Clinopleura Scudder , all species. Decorana buxtoni (Uvarov) . Decticita Hebard , all species. Eremopedes Scudder : E. cryptoptera (Rehn & Hebard) ; E. scudderi Cockerell. Idiostatus hermannii (Thomas) ; Inyodectes pallidus Rentz & Birchim ; Pediodectes nigromarginatus (Caudell) ; Steiroxys strepens Fulton. Pholidopterini . Eupholidoptera Maran. Parapholidoptera Maran , all species. Tettigoniini . Ateloplus notatus Scudder. Capnobotes Scudder : C. arizonensis (Rehn) ; C. occidentalis (Thomas) ; C. attenuatus Rentz & Birchim ; C. unodontus Rentz & Birchim ; C. granti Rentz & Birchim. Idionotus Scudder , possibly all species.

Pseudophyllinae . Cocconotini . Schedocentrus sp. ( Figs 48–49 , 116 ). Teleutiini . Teleutias : T. fasciatus Brunner von Wattenwyl ; T. akratonos Montealegre-Z. & Morris.

Third hypothetical morphological correlation

Simple cerci and a single sclerite TS or a single enlarged process ti. In most cases, the tip of TS or ti remains exposed beyond the abdominal apex. Such combination occurs in:

Phaneropterinae . Holochlorini . Stictophaula Hebard : S. armata Ingrisch ; S. spinosolaminata (Brunner von Wattenwyl) . Incertae sedis. Tamdaopteron major Gorochov , with the tip of cerci curved outward, apparently not adapted for lateral grasping of females.

Pseudophyllinae . Eucocconotini . Panoploscelis specularis Beier. Leptotettigini . L. crassicerci ( Figs 56 – 57 , 120 ); Leptotettix sp. ( Figs 44–47 , 121 ); Gen. 8 sp. ( Figs 54 – 55 , 119 ). Pleminiini . Gen. 9 sp. ( Figs 60 – 61 , 123 ); Gongrocnemis sp. ( Figs 62 – 63 , 124 ).

Variations to the proposed hypothetical morphological correlations

Copiphorini . Eriolus sp. with two sclerites TS in an inverted position and simple cerci ( Figs 19 , 102 ).

Meconematinae . P. ocraceovittata , with inverted sclerites TS but cerci simple ( Fig. 108 ). Phaneropterinae . Dysoniini . Machima sp. ( Fig. 109 ), which has almost entirely membranous phallus in combination with cerci that are apparently adapted to hook. Incertae sedis. Gen. 6 sp. with membranous phallus, excepting for the sclerotization on lobe dl ( Fig. 82 ) and cerci slightly curved ( Fig. 115 ). Pseudophyllinae . Eucocconotini . G. vorax ( Figs 50–51 , 117 ), which bears two enlarged sclerites TS, but cerci in Gnathoclita species are not adapted for grasping ( De Souza et al. 2011). The TS in this species are very different from those of other species bearing paired sclerites TS, because their apices exceed the length of the phallus. Leptotettigini . Gen. 7 sp. ( Figs 52–53 , 118 ), without sclerites TS but with curved cerci. Pterochrozini . In Porphyromma sp. the phallus bears a sclerite TS with divided apex ( Fig. 66 ), not projected over the lobe dl, and cerci are not adapted for grasping ( Fig. 126 ). Porphyromma ocellata and P. ocellata have simple cerci as in most Pterochrozini ( Figs 126, 127 ), and the phallus bears spiny microstructures throughout the surface of the lobe dl ( Figs 68–69 ).

Figures

FIGURES 26–29. Images of phalli of Listroscelidinae species, incertae sedis. 26–27. Gen.1 sp., dorsal (26) and ventral (27) views, with a diagram showing fused sclerites TS and VS in 26. 28–29. Gen.2 sp., posterior (28) and ventral (29) views. Scale bars: 1.00 mm.

FIGURES 97–128. Drawing silhouettes of cerci of examined male specimens (Figs 6–71). See Appendix 2 for suprageneric classification. 97. Eschatoceras sp. 98. Uchuca ferreirai (Piza). 99. Conocephalus sp.1. 100. Conocephalus sp.2. 101. Conocephalus sp.3. 102. Eriolus sp. 103. Neoconocephalus sp.1. 104. Neoconocephalus sp.2. 105. Cerberodon viridis Perty. 106. Gen.1 sp. 107. Gen.2 sp. 108. Phlugis ocraceovittata Piza. 109. Machima sp. 110. Itarissa sp. 111. Anaulacomera sp. 112. Gen.3 sp. 113. Gen.4 sp. 114. Gen.5 sp. 115. Gen.6 sp. 116. Schedocentrus sp. 117. Gnathoclita vorax (Stoll). 118. Gen.7 sp. 119. Gen.8 sp. 120. Leptotettix crassicerci Piza. 121. Leptotettix sp. 122. Acanthodis sp. 123. Gen.9 sp., 124. Gongrocnemis sp. 125. Spinapecta alieniphaga Naskrecki & Lopes-Andrade. 126. Porphyromma sp. 127. Pterochroza ocellata (Linnaeus). 128. Teleutias sp. Scale bars: 1.00 mm.

FIGURES 38–43. Images of phalli of Phaneropterinae species, incertae sedis. 38–39. Gen.3 sp., dorsal (38) and ventral (39) views. 40–41. Gen.4 sp., dorsal (40) and ventral (41) views, respectively. 42–43. Gen.5 sp., dorsal (42) and ventral (43) views. Scale bars: 1.00 mm.

FIGURES 60–65. Images of phalli of Pseudophyllinae species, showing examined Pleminiini (60–63) and Polyancistrini (64–65). 60–61. Gen.9 sp., dorsal (60) and ventral (61) views. 62–63. Gongrocnemis sp., dorsal (62) and ventral (63) views, with one diagram showing the sclerite TS and the VS (62) and another showing portion of the ventral lobe lw.vl (63). 64–65. Spinapecta alieniphaga Naskrecki & Lopes-Andrade, dorsal (64) and ventral (65) views, respectively; single sclerite TS hidden between the two processes ti. Scale bars: 1.00 mm.

FIGURES 66–71. Images of phalli of Pseudophyllinae species, showing examined Pterochrozini. 66–67. Porphyromma sp., dorsal (66) and ventral (67) views; the arrow indicates a secondary portion over the dl with conspicuous spines, that are not part of the sclerite TS, and we have not determined whether this portion is part of the process ti or an additional component on the dl. 68–69. Pterochroza ocellata (Linnaeus), dorsal (68) and posterior (69) views. 70–71. Teleutias sp., posterior (70) and anterior (71) views. Scale bars: 1.00 mm.

FIGURES 54–59. Images of phalli of Pseudophyllinae species, showing examined Leptotettigini (54–57) and Pleminiini (58–59). 54–55. Gen.8 sp., dorsal (54) and ventral (55) views, respectively, with a diagram showing portions of the ventral lobes lw.vl and up.vl in 54. 56–57. Leptotettix crassicerci Piza, dorsal (56) and ventral (57) views, respectively. 58–59. Acanthodis sp., dorsal (58) and ventral (59) views, respectively. Scale bars: 1.00 mm.

FIGURES 10–15. Images of phalli of Conocephalinae species, showing examined Agraeciini (10–13) and Conocephalini (14–15). 10–11. Eschatoceras sp., dorsal (10) and ventral (11) views, with a diagram showing the fold fdl in 10. 12–13. Uchuca ferreirai (Piza), dorsal (12) and ventral (13) views. 14–15. Conocephalus sp.2, dorsal (14) and ventral views (15). Scale bars: 1.00 mm.

FIGURES 6–9. Phallus of Conocephalus sp.1 (Conocephalinae, Conocephalini). 6. Image of dorsal view. 7. Image of ventral view. 8. Diagram of the left half of dorsal view, and detail showing the lobe ddl and one sclerite TS. 9. Diagram of the right half of ventral view, and detail showing the lobe vdl and one sclerite VS. Scale bar: 1.00 mm.

FIGURES 16–21. Images of phalli of Conocephalinae species, showing examined Conocephalini (16–17) and Copiphorini (18–21). 16–17. Conocephalus sp.3, dorsal (16) and ventral (17) views. 18–19. Eriolus sp., dorsal (18) and ventral (19) views, respectively. 20–21. Neoconocephalus sp.1., posterior (20) and ventral (21) views, with a diagram showing portions of the ventral lobes up.vl and lw.vl in 20. Scale bars: 1.00 mm.

FIGURES 22–25. Phallus of Cerberodon viridis Perty (Listroscelidinae, incertae sedis). 22. Image of dorsal view. 23. Image of ventral view. 24. Diagram of the left half of dorsal view, and detail showing the two sclerites TS and two scleritesVS, all four sclerites are unfused, the boundaries between them are indicated by the black dotted line. 25. Diagram of the right half of ventral view. Scale bar: 1.00 mm.

FIGURES 34–37. Phallus of Anaulacomera sp. (Phaneropterinae,Viadanini). 34–35. Images of dorsal (34) and ventral (35) views, respectively. 36. Diagram of the left half of dorsal view. 37. Diagram of the right half of ventral view. Scale bar: 1.00 mm.

FIGURES 48–53. Images of phalli of Pseudophyllinae species, showing examined Cocconotini (48–49), Eucocconotini (50–51) and Leptotettigini (52–53). 48–49. Schedocentrus sp., dorsal (48) and ventral (49) views. 50–51. Gnathoclita vorax (Stoll), dorsal (50) and ventral (51) views. 52–53. Gen.7 sp., dorsal (52) and ventral (53) views, respectively. Scale bars: 1.00 mm.

FIGURES 44–47. Phallus of Leptotettix sp. (Pseudophyllinae, Leptotettigini). 44–45. Images of dorsal (44) and ventral (45) views, respectively. 46. Diagram of the left half of dorsal view, and detail showing the entire sclerite TS. 47. Diagram of the right half of ventral view. Scale bar: 1.00 mm.

FIGURES 79–82. Components and microstructures in phalli of Phaneropterinae species, showing examined Dysoniini (79) and Viadanini (80–82). 79. Machima sp., showing microstructures on process ti. 80–81. Anaulacomera sp., showing part of lobe dl and processes ti (80), part of fold df and lobe dl, and microstructures on ti (81). 82. Gen.6 sp., showing amorphous sclerotization on lobe dl. Scale bars: 0.05 mm (79, 81–82), 0.20 mm (80).