The fifth family of the true crickets (Insecta: Orthoptera: Ensifera: Grylloidea), Oecanthidae defin. nov.: phylogenetic relationships and divergence times Author Campos, Lucas Denadai De Author Dias, Pedro Guilherme Barrios De Souza Author Audino, Jorge Alves Author Desutter-Grandcolas, Laure Author Nihei, Silvio Shigueo text Zoological Journal of the Linnean Society 2023 2022-11-02 197 4 1034 1077 http://dx.doi.org/10.1093/zoolinnean/zlac066 journal article 10.1093/zoolinnean/zlac066 8c2036d0-8307-4b09-8c8b-c5eeea9d030a 0024-4082 7803427 4BB4333D-64F0-4485-9C2B-47546ECFE65F IDENTIFICATION KEY TO OECANTHIDAE SUBFAMILIES, SUPERTRIBES AND TRIBES 1. Fastigium truncated in frontal and lateral views ( Fig. 5B ; Supporting Information, Fig. S1G ), male anterolateral region of metanotum not inflated, tarsal claws inner margin serrulated ( Fig. 5C ; Supporting Information, Fig. S6 Gb), TIII bearing six or more inner subapical spurs (Supporting Information, Fig. S6 Gb) ............................................................................................................................................... Euscyrtinae - Fastigium not truncated in frontal and lateral views (Supporting Information, Fig. S1A–C, I ), male anterolateral region of metanotum inflated (Supporting Information, Fig. S4C–E ), TIII bearing maximum six inner subapical spurs (except some Podoscirtinae taxa bearing more than six), claws inner margins not serrulated (Supporting Information, Fig. S6 Ga) ..............................................................................................2 2. Ventral inner apical spur of TIII reduced or absent (Supporting Information, Fig. S6E ), FWs lateral field with an angle less than 90° related to dorsal field in posterior view (Supporting Information, Fig. S2J2 ), ovipositor straight in lateral view (Supporting Information, Fig. S4 Jb), tip of ovipositor dorsal valves forked (Supporting Information, Figs S4 Ja, S5C). Male genitalia: rami two or more times longer than pseudepiphallic sclerite ............................................................................................................ Oecanthinae …4 - Ventral inner apical spur of TIII developed (Supporting Information, Fig. S6D ), FWs lateral field with an angle ~90° related to dorsal field in posterior view (Supporting Information, Fig. S2J 1 ), ovipositor up or downcurved in lateral view (Supporting Information, Fig. S4K, L ), tip of ovipositor dorsal valves single. Male genitalia: rami slightly longer or shorter than pseudepiphallic sclerite ...............................................3 3. Posterior margin of pronotum convex in the middle (Supporting Information, Fig. S4 Aa). TIII with 5/5 subapical spurs or more. Ovipositor not flattened (Supporting Information, Fig. S4J, L ), apex of dorsal valves covering ventral valves laterally and sometimes ventrally ( Fig. 9B, C ; Supporting Information, S5A, D, E, G), generally strongly sclerotized, laterals strongly serrulate, laterals of apex of ventral valves generally smooth. Male genitalia: inner margin of LLophi not membranous; ectophallic fold single or bilobate, posterior projection of endophallic sclerite, when present, bilobate ( Fig. 9A ); endophallic apodeme, when present, flattened laterally ( Fig. 9A , 12E ; Supporting Information, Fig. S9A )........... Podoscirtinae …6 - Posterior margin of pronotum entirely convex (Supporting Information, Fig. S4 Ab). TIII with 5/4 subapical spurs (4/ 3 in Perutrella and 4/ 4 in some Paroecanthini ). Ovipositor flattened dorso-ventrally ( Fig. 16B ; Supporting Information, Fig. S4K ), apex of dorsal valves above ventral valves ( Fig. 16C ; Supporting Information, Fig. S5B, F, H ), same sclerotization as the entire ovipositor, laterals slightly or not serrulate; laterals of apex of ventral valves generally serrulate. Male genitalia: inner margin of LLophi generally membranous ( Figs 18C , 19D ; Supporting Information, Fig. S7A ); ectophallic fold single, posterior projection of endophallic sclerite, when present, single; endophallic apodeme, when present, not flattened laterally (Supporting Information, Fig. S8L–N ).................................................................................... Tafaliscinae …12 4. Specimens hypognathous ( Fig. 8B ); TI and TII ventral inner apical spurs present (Supporting Information, Fig. S5O, P ); FIII proximal portion wider than distal portion in lateral view ( Figs S4K, S 6A ); first tarsomere of leg III with dorsal spines ( Fig. 12C, D ); tarsal claws not bifurcated ( Fig. S6 Ga). Tip of ventral valves of ovipositor forked. Male FW: mirror not occupying more than half of FW (Supporting Information, Fig. S2E, G ), with one dividing vein; apical field present (Supporting Information, Figs S5J, S 6C, D ). Male genitalia: ectophallic arc curved posteriorly; ventral valves present, larger than the entire genitalia (Supporting Information, Fig. S9C ); endophallic sclerite two times or longer than pseudepiphallic sclerite (Supporting Information, Fig. S7D ). Neotropical distribution ........................................................... Diatrypidi /Diatrypini - Specimens prognathous ( Fig. 6A ; Supporting Information, S1K); TI and TII ventral inner apical spurs absent; FIII proximal portion similarly thin to distal portion in lateral view; first tarsomere of leg III without spines ( Fig. 7B, C ); tarsal claws bifurcated ( Fig. 6C ; Supporting Information, Fig. S6 Gc). Tip of ventral valves of ovipositor not forked. Male FW (when developed): large mirror occupying almost or more than half of FW, with two dividing veins ( Fig. 6B ; Supporting Information, Fig. S3H ); apical field absent or almost no discernible (Supporting Information, Fig. S3H ). Male genitalia: ectophallic arc curved anteriorly (Supporting Information, Fig. S8G ); ventral valves poorly developed or absent; endophallic sclerite short (Supporting Information, Fig. S8G ). Worldwide distribution .................................................... Oecanthidi …5 5. Cerci short, shorter than FII (Supporting Information, Fig. S4G ); ventral outer apical spur of TII present; subapical spurs of TIII absent ( Fig. 7A ); ventral outer and inner apical spurs of TIII absent (2/2) ( Fig. 7B, C ), first tarsomere apical spurs of leg III absent. Male genitalia: distal prolongation of ectophallic arc absent ................................................................................................................................................ Xabeini - Cerci long, longer than FII (Supporting Information, Fig. S4H ); ventral outer apical spur of TII absent; subapical spurs of TIII present, generally 3/3; ventral outer and inner apical spurs of TIII present (3/3); first tarsomere apical spurs of leg III present. Male genitalia: distal prolongation of ectophallic arc present (Supporting Information, Fig. S8G ) ................................................................................................. Oecanthini 6. Median and lateral ocelli generally aligned ( Fig. 11A ; Supporting Information, Fig. S1B ); fifth article of maxillary palpi upcurved 90° ( Figs 11B , 12B ; Supporting Information, Fig. S1 Lc). Male FWs: PCu vein curved more than 90°, sinuous ( Figs 14B , 15B ; Supporting Information, Fig. S3 Bb, G); CuPa not parallel to proximal region of PCu ( Figs 14B , 15B ; Supporting Information, Fig. S3G ); harp veins, when present, parallel to PCu vein ( Figs 14B , 15B ; Supporting Information, Fig. S3G ). New world distribution ........................................................................................................................................................ Hapithidi …7 - Median and lateral ocelli not aligned; fifth article of maxillary palpi slightly upcurved (Supporting Information, Fig. S1 Lb). Male FWs: PCu vein curved 90°, not sinuous ( Fig. 10C ; Supporting Information, Fig. S3 Ba, F); CuPa parallel to proximal region of PCu ( Fig. 10C ; Supporting Information, Fig. S3F ); harp veins, when present, diagonal ( Fig. 10C ; Supporting Information, Fig. S3F ). Old world distribution .................................................................................................................................................. Podoscirtidi …10 7. Veins of lateral field parallel to dorsal field (Supporting Information, Fig. S2 Ia). Male FWs: stridulatory apparatus absent (Supporting Information, Fig. S3A ); apical field not delimited .........................................8 - Veins of lateral field perpendicular to dorsal field (Supporting Information, Fig. S2 Ib). Male FWs: stridulatory apparatus present; apical field delimited (Supporting Information, Fig. S3F, G ) .....................9 8. Eyes posterior margin concave in lateral view ( Fig. 12B ; Supporting Information, Fig. S1 Eb), with one horizontal band or without bands; TI outer tympana generally present, inner absent; first tarsomere apical spurs of leg III longer than the tarsomere ( Fig. 12C, D ). Male genitalia: ectophallic arc not connected ( Fig. 12E ; Supporting Information, Fig. S9A ); endophallus U-shaped ( Fig. 12E ; Supporting Information, Fig. S9A ), endophallic apodeme two or more times longer than PsP ( Fig. 12E ; Supporting Information, Fig. S9A ); rami not connected......................................................................................................... Aphonomorphini - Eyes posterior margin straight in lateral view (Supporting Information, Fig. S1 Ea); eyes with two or three horizontal bands ( Figs 11A , 13A ; Supporting Information, Fig. S1F ); TI outer tympana absent, inner present; apical spurs of first tarsomere of leg III generally same size or shorter than the tarsomere. Male genitalia: ectophallic arc connected ( Fig. 13B ; Supporting Information, Fig. S8I ); endophallus not U-shaped, endophallic apodeme same shorter or slightly longer than PsP; rami connected ( Fig. 13B ; Supporting Information, Fig. S8I ) .................................................................................................................... Cearacesaini 9. Median and lateral ocelli distant, not connected. FWs lateral and dorsal field same-sized (Supporting Information, Fig. S2H 1 ). Pronotum caudal margin entirely convex ( Fig. 14A ; Supporting Information, Fig. S4 Ab). Dorsal valves of ovipositor lateral margins slightly serrulate, same colour as the entire ovipositor. Male genitalia: MedLophi present ( Fig. 14C ; Supporting Information, Fig. S8C ); anterior margin of pseudepiphallic sclerite not folded; ectophallic arc reduced or absent ............................................ Hapithini - Median and lateral ocelli remarkably close, sometimes connected ( Fig. 15C ; Supporting Information, Fig. S1B, D ). FWs lateral field shorter than dorsal field (Supporting Information, Fig. S2H2 ). Dorsal valves of ovipositor lateral margins strongly serrulate, darker than the entire ovipositor (Supporting Information, Fig. S5A, E ). Pronotum caudal margin convex on the middle ( Figs 15A ; Supporting Information, Fig. S4 Aa). Male genitalia: MedLophi absent; anterior margin of pseudepiphallic sclerite folded dorsally ( Fig. 15D ; Supporting Information, Fig. S8F ); ectophallic arc present ....................................................... Phyllogryllini 10. Male FWs without stridulatory apparatus, only longitudinal veins, anal area not delimited; male genitalia PsP not cylindrical; endophallus asymmetric; apex of dorsal valves of ovipositor generally wider than entire ovipositor in dorsal and ventral views................................................................................ Aphonoidini - Male FWs generally bearing stridulatory apparatus sometimes harp veins or mirror lacking (or both), anal area delimited; male genitalia PsP frequently cylindrical; endophallus frequently symmetrical; apex of dorsal valves of ovipositor generally same width of the entire ovipositor in dorsal and ventral view .......11 11. TI inner tympanum usually not covered by a sclerotized tab; pronotum DD not flattened in lateral view; male genitalia generally capsular (Supporting Information, Fig. S7F ); arc and ventral projection of ectophallic invagination short, endophallic sclerite well developed .............................................Podoscirtini - TI inner tympanum covered by a sclerotized tab; pronotum DD not flattened in lateral view; male genitalia not capsular; arc and ventral projection of ectophallic invagination elongated ( Fig. 10E ), endophallic sclerite sometimes regressed or absent ............................................................................................ Truljaliini 12. Body not robust, pronotum longer than wide in dorsal view. Apex of ovipositor pointed or rounded ( Fig. 16C ; Supporting Information, Fig. S5F ). Male genitalia: MedLophi absent; endophallus flattened dorso-ventrally (Supporting Information, Fig. S8H ) or strongly reduced (Supporting Information, Fig. S8B ), lateral margins not folded; endophallic apodeme absent ......................................... Paroecanthidi …13 - Body robust, pronotum as wide as long or wider than long in dorsal view. Apex of ovipositor truncated ( Fig. 19C ; Supporting Information, Fig. S5B, H ). Male genitalia: MedLophi present; endophallus not flattened, lateral margins folded ( Fig. 19D ; Supporting Information, Fig. S8L ); endophallic apodeme generally present .................................................................................................................................. Tafaliscidi /Tafaliscini…14 13. TI tympana present. FWs covering abdomen partially or totally, stridulatory apparatus present; PCu vein curved, generally bisinuous, rarely sinuous (Supporting Information, Fig. S3C ); apical field delimited. Structures of male genitalia generally regressed, mainly from ectophallic invagination and endophallus (except Adenophallusia and Ectotrypa ) (Supporting Information, Fig. S8B ) .................... Paroecanthini …19 - TI tympana absent. FWs absent (Supporting Information, Fig. S2A ), only covering the metanotum (Supporting Information, Fig. S2B ) or covering the abdomen with longitudinal veins ( Fig. 18A ), stridulatory apparatus absent; PCu vein not curved or absent; apical field, when present, not delimited. Structures of male genitalia not regressed ................................................................................................ Neometrypini …25