OECANTHIDAE BLANCHARD, 1845 DEFIN.

Campos, Lucas Denadai De, Dias, Pedro Guilherme Barrios De Souza, Audino, Jorge Alves, Desutter-Grandcolas, Laure & Nihei, Silvio Shigueo, 2023, The fifth family of the true crickets (Insecta: Orthoptera: Ensifera: Grylloidea), Oecanthidae defin. nov.: phylogenetic relationships and divergence times, Zoological Journal of the Linnean Society 197 (4), pp. 1034-1077 : 1047-1049

FAMILY OECANTHIDAE BLANCHARD, 1845 DEFIN. NOV.

Oecanthites Blanchard, 1845: 245, Saussure, 1874: 428, Saussure, 1878: 590.

Oecanthiens Saussure, 1874: 427, Saussure, 1878: 534. Oecanthinae Saussure, 1897: 251 , Scudder, 1987: 62, Azam, 1901: 104, Kirby, 1906: 109, Zeuner, 1939: 212, Walker, 1966: 265, Otte & Alexander, 1983: 374, Gorochov, 1986: 851–858, Toms & Otte, 1988: 471, Nickle, 1992: 195, Otte, 1994: 85, Rentz, 1996: 147, Otte & Perez-Gelabert, 2009: 491, Gorochov, 2015: 31–34, Collins & Carson, 2014: 170, Chintauan-Marquier et al., 2016: 73, Rentz & Su, 2019: 231.

Oecanthidae Brunner von Wattenwyl, 1873: 164 , Brunner von Wattenwyl, 1882: 420, Bruner, 1916: 395, Chopard, 1949: 673, Chopard, 1951: 198, Vickery, 1977: 11, Chopard, 1968: 427, Kevan, 1982: 2, Desutter, 1987: 223, Coray & Lehmann, 1998: 94.

Type genus: Oecanthus Serville, 1831 .

Distribution: Worldwide.

Diagnosis: Crickets inhabiting trees and shrubs (a few exceptions registered in caves, see Xabeini ), small to large size; apterous, brachypterous or with developed wings (with or without stridulatory apparatus). Latero-anterior regions of metanotum generally inflated (Supporting Information, Fig. S4C, D, E ), sometimes with bristles. TIII with five or more inner apical spurs (except Oecanthidi ); first and second tarsomeres of legs I and II same-sized with pulvillum (Supporting Information, Fig. S5J, L ) and second tarsomere of leg III with pulvillum, second tarsomere of TI and TII flattened dorso-ventrally (except some Oecanthidi ); inner apical spurs three times or more longer than outer apical spurs of TIII (Supporting Information, Fig. S6D ); apex of ovipositor generally with projections, protuberances and/or lateral margins serrulate, sometimes smooth ( Euscyrtinae ) ( Fig. 5D ), ovipositor generally upcurved, sometimes straight or downcurved. Male genitalia: median region of

THE FIFTH FAMILY OF TRUE CRICKETS 1049

ectophallic arc not connected ( Fig. S7A ) (except Cearacesaini ).

Differential diagnosis: Oecanthidae are separated from Mogoplistidae by the presence of subapical spurs on serrulated hind tibiae (subapical spurs lacking in Mogoplistidae ). They are separated from Trigonidiidae by their serrulated TIII, head shape, lack of strong setae on the head and body, number of apical spurs on TI and TIII and male pseudepiphallic sclerite (see: Desutter-Grandcolas et al., 2021). They are separated from Phalangopsidae by the different number of spurs on TIII, FIII longer than TIII, the second tarsomere flattened (except in some Phaloriinae ) and presence of pulvillum. Finally, they are also excluded from Gryllidae by the shape of MedLophi and LLophi, presence of serrulation above the subapical spurs on TIII, head pubescent, the fastigium shape (wider in Gryllidae ), antennal scape not reduced, first and second tarsomeres same-sized (first tarsomere longer than the second in Gryllidae ), presence of pulvillum (also present in Eneopterinae ) ( Table 6 ).

Included subfamilies: Euscyrtinae Gorochov, 1985 , Oecanthinae Blanchard, 1845 defin. nov., Podoscirtinae Saussure, 1878 defin. nov., Tafaliscinae Desutter, 1988 defin. nov.

Remarks: Oecanthidae are mainly composed of crickets that live in plants, from long grasses to tall trees. The apex of their ovipositor, generally strongly sclerotized, bears serrulation and/or structures adapted to ovipositing inside leaves and under the bark of trees. Their forewings are diverse, and several groups are characterized by their acoustic apparatus or nonacoustic communication structures. This newly erected family was considered as the clade F in ChintauanMarquier et al. (2016) and was split between the subfamily Oecanthinae and the Podoscirtinae Subfamily Group by Cigliano et al. (2022). Based on the phylogenetic hypotheses presented above, with solid support and the morphological synapomorphies ( Table 4 ; Supporting Information, Fig. S10 ), we have enough evidence to elevate this lineage to family level ( Fig. 3 ; Supporting Information, Figs S13 , S 14 ) and separate it from Gryllidae defin. nov.

Figures

Figure 3. Time-calibrated phylogenetic tree of Oecanthidae. Divergence-time analysis under Bayesian inference based on four DNA markers and six fossil calibrating internal nodes (red asterisks). Red numbers are the mean ages of the nodes. Bars indicate 95% highest posterior density intervals for nodes of interest. Posterior probabilities different from 1 are indicated in black numbers on nodes. Inset A, lineages-through-time plot.

Figure 4. Distribution maps of Oecanthidae subfamilies and supertribes.

Figure 5. A, Euscyrtus aff. bipunctatus, male, dorsal habitus. Proturana subapterus: B, head and pronotum, lateral view; C, claw; D, ovipositor; dorsal view. Scales: 1mm.

Figure 6. A, Neoxabea breƲipes, head dorsal view. Oecanthus sp.: B, male, pronotum and FW; C, claw. Scales: 1mm.

Figure 7. Neoxabea breƲipes. A, hind tibia and tarsi; B, hind tibia distal margin and tarsi, inner view; C, hind tibia distal margin and tarsi, outer view. Scales: 1mm. Abbreviations: see Material and methods.

Figure 10. A, Fryerius sp., male, dorsal habitus; B, Munda aff. asyrinx, male, dorsal habitus; C, Truljalia hibinonis, male, pronotum and FW, dorsal view; D, Madasumma melanotum, male genitalia, lateral view. Scales: 1mm. Abbreviations: see Material and methods.

Figure 13. A, Cearacesa sp., male; B, Cearacesa cearensis, male genitalia, dorsal view. Scale: 1mm. Abbreviations: see Material and methods.

Figure 14. A, H. (Hapithus) Ʋagus, male, dorsal habitus; B, Stenogryllus sp., male, pronotum and FW, dorsal view; C, H. (Hapithus) sp., male genitalia, lateral view. Scales: 1mm. Abbreviations: see Material and methods.