Phaloriinae Gorochov, 1985

Desutter-Grandcolas, Laure, 2015, Phalangopsidae crickets from Tropical Africa (Orthoptera, Grylloidea), with descriptions of new taxa and an identification key for African genera, Zootaxa 3948 (3), pp. 451-496 : 454-456

Phaloriinae Gorochov, 1985

Phaloriini Gorochov, 1985: 19 . Phaloriinae , Gorochov 1996: 55.

Type genus. Phaloria Stål, 1877 .

The Phaloriinae have been originally described by Gorochov (1985) as a tribe of the subfamily Podoscirtinae . Later, Gorochov (see Gorochov 2003 and references therein) considered the Phaloriini as a subfamily close to a Cachoplistinae group comprising the Cachoplistini Saussure, 1877 and the Homoeogryllini Gorochov, 1986 , within the « Phalangopsinae subfamily group »: he then separated the Phaloriinae from the Cachoplistinae by two characters: « the ability of its representatives to powerful jumps (plesiomorphy) and the presence of more or less widened lobes on 2nd tarsal segments (autapomorphy of the Phaloriinae associated with obligatory or almost obligatory phylophilous mode of life) » ( Gorochov 2003, p. 84). These characters revealed useless in a comparative frame, as the ability to jump could probably apply to most of the cricket clade, while widened second tarsal segments are found in all the crickets living on plants and could at best be a homoplasy, showing a parallel evolution related to common habitat.

The phylogenetic evidence did not support a close relationships between the Phaloriinae and a " Cachoplistinae group" ( Chintauan-Marquier et al. 2013, 2015): the Phaloriinae were found monophyletic and robust, provided they include taxa previously not classified in the subfamily. Up to early 2015, nine genera were listed in the Phaloriinae according to the OSF, three of which subdivided into two to six subgenera. They were distributed into two tribes: the Phaloriini comprised Ceyloria Gorochov, 1996 , Gorochovius Xie, Zheng & Li, 2004 , Phaloria Stål, 1877 , Pseudotrigonidium Chopard, 1915 , Strophiola Uvarov, 1940 , Sumatloria Gorochov, 2003 , Trellius Gorochov, 1988 , Tremellia Stål, 1877 , and Vescelia Stål, 1877 , while the Subtiloriini Gorochov, 2003 included Heterotrypus , Subtiloria , Schizotrypus and Kameruloria . One additional genus belongs to the Phaloriinae clade: Megacris Desutter-Grandcolas, 2009 from the Vanuatu, actually described as a Phaloriinae . In the present paper, three additional new genera (six new species) of Phaloriinae are described, Afrophaloria Desutter-Grandcolas , n. gen., included in the phylogenetic analysis as ' Phaloriinae ngen-Tan', Upupagryllus Desutter-Grandcolas , n. gen. and Phasmagryllus Desutter-Grandcolas , n. gen. Three new species of Kameruloria are also described. Afrophaloria Desutter-Grandcolas , n. gen. and Upupagryllus Desutter-Grandcolas , n. gen. are the first apterous Phaloriinae ever described. Taking into account these genera and the troglobitic Megacris greatly increases the ecological and morphological diversity of the subfamily.

Extended morphological definition. Small to medium species (except the large Megacris ). Head. Head longer than wide. Eyes moderately to highly protruding dorsally (less so laterally). Fastigium rounded laterally, squared, not separate from the vertex, as wide or slightly wider than the scapes, with three equidistant or almost equidistant ocelli. Scapes longer than wide. Maxillary palpi not very elongate (except in Phasmagryllus Desutter- Grandcolas, n. gen.); joint 5 widened regularly toward apex, its apex most often truncated apically nearly straight (oblique in Afrophaloria Desutter-Grandcolas , n. gen. and Phasmagryllus Desutter-Grandcolas , n. gen.). Pronotum. Transverse. Legs. TI most often with one inner and one outer tympana, but outer tympanum sometimes absent (both tympana lacking in Afrophaloria Desutter-Grandcolas , n. gen. and Upupagryllus subapterus Desutter- Grandcolas, n. sp.); tympana variously developed, with three different patterns: inner tympanum the widest, or outer tympanum the widest, or both tympana small and almost equal in size; at tympana level, TI shape almost not modified, or greatly inflated and rounded, or just higher in lateral view. TI with two apical spurs, ventral. TII with three apical spurs, the outer dorsal spur missing. TIII with three pairs of apical spurs, the median the longest on outer side (except in Phasmagryllus Desutter-Grandcolas , n. gen.), the dorsal the longest on inner side; four pairs of subapical spurs, slightly alternate, thin; two different patterns of subapical spur relative lengths: In type 1, inner subapical spurs all much shorter than TIII dorsal apical spur, and nearly equal in length, although subapical spurs 2 and 3 may be slightly longer than subapical spurs 1 and 4. In type 2, inner subapical spurs growing longer toward TIII apex, the spur 4 being the smallest and the spur 1 the longest, with spurs 2 and 3 intermediate in size; the longest subapical spur shortly smaller than TIII dorsal apical spur (see Desutter-Grandcolas 2009, Fig. 5 G). TIII dorsally narrowed, often but not always making a kind of keel undulating between the subapical spurs; serrulation very sparse, most often lacking between the subapical spurs (more numerous in Megacris ); when present between the subapical spurs, spines most often aligned in one row only, located on the tibial keel; above subapical spurs, two rows of sparse spines, more numerous on outer side than on inner side. Second tarsomeres flattened and somewhat expanded (except in Afrophaloria Desutter-Grandcolas , n. gen., Phasmagryllus Desutter-Grandcolas , n. gen., Upupagryllus Desutter-Grandcolas , n. gen. and Megacris ). Basitarsomeres III with one row of spines (two rows in Megacris ).

Male. No evident glandular structures on metanotum and tergites (except perhaps in Phasmagryllus Desutter- Grandcolas, n. gen., Fig. 8 F). FWs well developed and longer than the abdomen (except in Afrophaloria Desutter- Grandcolas, n. gen. and Upupagryllus subapterus Desutter-Grandcolas , n. sp.; unknown in Megacris ). When present, HWs most often longer than FWs. Stridulatory apparatus complete. Dorsal field venation: harp with numerous veins, either oblique, parallel and regularly distributed over the file length, or more rounded and irregularly distributed over file length; mirror large, crossed by two transverse veins, one attached to the anterior part of the mirror, and the other attached to the posterior part of the mirror, the two veins being parallel over most of their length, or not (only one vein in Phasmagryllus Desutter-Grandcolas , n. gen., Fig. 9 B, and Upupagryllus alatus Desutter-Grandcolas , n. sp., Fig. 13 K); a wide, transverse cell running posteriorly to the mirror; diagonal vein bifurcate anteriorly (except in Phasmagryllus Desutter-Grandcolas , n. gen.); chords 1 and 2 well separate from chord 3, and fused quite posteriorly (except in Phasmagryllus Desutter-Grandcolas , n. gen., where chords 2 and 3 are fused posteriorly, Fig. 9 B); CuP present and reaching the first harp veins (shorter in Upupagryllus alatus Desutter-Grandcolas , n. sp., Fig. 13 K, and lacking in Phasmagryllus Desutter-Grandcolas , n. gen., Fig. 9 B); apical field present, more or less developed, most often with several clearly marked cell alignments (but reduced in Phasmagryllus Desutter-Grandcolas , n. gen., Fig. 9 B, and Upupagryllus alatus Desutter-Grandcolas , n. sp., Fig. 13 K). Lateral field well developed, with numerous oblique, parallel veins (except in Phasmagryllus Desutter- Grandcolas, n. gen.). Supra anal plate smooth, or with thick denticles (see Gorochov's papers). Subgenital plate well-developed, except in Afrophaloria n. gen.; apex straight or variously bisinuate.

Male genitalia. Very diverse, often hypertelic and asymmetrical. Pseudepiphallus with a particular upper part, located on the membrane connecting the phallic complex to the paraprocts and partly sclerotized as a transverse paired or impaired sclerite; rami partly invaginated, separate from the upper part of pseudepiphallus, but connected to its lower "normal" part. Pseudepiphallic parameres variable. Ectophallic apodemes well-developed; arc most often not complete (but present at least in Kameruloria p.p., Fig. 6 , and Afrophaloria Desutter-Grandcolas , n. gen., Fig. 3 ). Ectophallic fold most often hypertelic (ectophallic fold simple and symmetrical in Upupagryllus Desutter- Grandcolas, n. gen. and Phasmagryllus Desutter-Grandcolas , n. gen.); ventral margins often large and asymmetrical. Endophallic sclerite and apodeme, and ectophallic dorsal valves variable. Dorsal cavity lacking.

Female. FWs and HWs most often well developed; lacking in apterous Afrophaloria Desutter-Grandcolas , n. gen. and Upupagryllus Desutter-Grandcolas , n. gen.; short and corneous in Megacris and Phasmagryllus Desutter- Grandcolas, n.gen. Subgenital plate transverse, the apex sinuate. Ovipositor apex laterally flattened and smooth (most Subtiloriini , Upupagryllus Desutter-Grandcolas , n. gen.), more or less globulous ( Tremellia , Pseudotrigonidium ), or thicker with various ornementation (few big teeth, numerous small teeth, or apex small and acute with transverse crests).

Female genitalia. Copulatory papilla variable.

Remark. In two females of Afrophaloria Desutter-Grandcolas , n.gen., a spermatophore was present in the female genitalia. The ampoula of the spermaphore was inside of a membranous pouch below the aperture of the spermathecal duct. This was observed also in one female of Kameruloria nigricornis Desutter-Grandcolas , n. sp.

Figures

FIGURE 3. Afrophaloria Desutter-Grandcolas, n. gen. Male genitalia, dorsal (A, D, G), ventral (B, E, H), lateral (C, F, I) in: Afrophaloria amani Desutter-Grandcolas, n. sp. (A – C), Afrophaloria hempae Desutter-Grandcolas, n. sp. (D – F) and Afrophaloria apiariensis Desutter-Grandcolas, n. sp. (G – I). Abbreviations: see page 3.

FIGURE 5. Kameruloria Gorochov, 2003. A – C, maxillary palpi; D – E, dorsal field of male forewing; F, male supra anal plate; G – H, male subgenital plate; I – J, male genitalia, details of pseudepiphallic processes, of Kameruloria nigricornis Desutter- Grandcolas, n. sp. (A, D, F, G, I), Kameruloria gabonensis Desutter-Grandcolas, n. sp. (B, H, J) and Kameruloria trimaculata Desutter-Grandcolas, n. sp. (C, E). Scale bars 1 mm.

FIGURE 6. Kameruloria Gorochov, 2003. Male genitalia in dorsal (A, D, G), ventral (B, E, H) and lateral (C, F, I) views of: Kameruloria nigricornis Desutter-Grandcolas, n. sp. (A – C), Kameruloria gabonensis Desutter-Grandcolas, n. sp. (D – F) and Kameruloria trimaculata Desutter-Grandcolas, n. sp. (G – I). Abbreviations, see page 3.

FIGURE 8. Phasmagryllus elegans Desutter-Grandcolas, n. sp., male holotype. A, dorsal view, scale 1 mm; B, head and pronotum, dorsal view; C, face and maxillary palpi; D, hindtibia, dorsal view; E, right forewing, dorsal view; F, metanotum and tergite structures, dorsal view; G, subgenital plate, ventral view; H, subgenital plate and right forewing, lateral view; I, supra anal plate, dorsal view. Arrows point to the left pseudepiphallic distal process, which is extremely long.

FIGURE 9. Phasmagryllus elegans Desutter-Grandcolas, n. sp. A, maxillary palpus, female allotype; B, C, venation of dorsal and lateral fields of male holotype FWs; D-F, male holotype subgenital plate, ventral (D), right side (E) and left side (F), with projections of asymmetrical pseudepiphallic processes; G, H, venation of dorsal and lateral fields of female allotype right FW; I, female allotype subgenital plate, ventral view; J, female allotype copulatory papilla. Scale bars 1 mm.

FIGURE 13. Upupagryllus Desutter-Grandcolas, n. gen., A – I, U. subapterus Desutter-Grandcolas, n. sp., A, maxillary palpus; B, apex of female ovipositor; C – I, female copulatory papilla, dorsal (C, F), ventral (D, G), lateral (E, H, I), in female fn 5, 23. v. 1995 (C – E), female fn 13, 17. v. 1995 (F – H) and female fn 19, 19. v. 1995 (I). J – P, U. alatus Desutter-Grandcolas, n. sp.: J, maxillary palpus; K, L, dorsal and lateral field of male holotype right FW; M, female subgenital plate; N – P, female copulatory papilla, dorsal (N), ventral (O) and lateral (P) views. Scale bars 1 mm.