New Brazilian Tafaliscina increase the diversity of this Neotropical cricket clade (Orthoptera: Grylloidea: Gryllidae: Oecanthinae: Paroecanthini)

ABSTRACT Tafaliscina Desutter, 1988 (Grylloidea Laicharting, 1781, Gryllidae Laicharting, 1781, Oecanthinae Blanchard, 1845, Paroecanthini Gorochov, 1986) are a Neotropical cricket clade with a remarkable morphological diversity. We study here their Brazilian representatives. We describe one new genus (Veredatrypa Campos n. gen.) and three new species from Cerrado and Caatinga, i.e., Veredatrypa rosai n. gen., n. sp., V. seca n. gen., n. sp. and V. fusca n. gen., n. sp., and two new species of Tafalisca Walker, 1869 from Amazon Forest, i.e., T. duckeana n. sp. and T. vestigialis n. sp. The new genus and the new species are mainly characterized by their male phallic complexes and forewings. An identification key of Tafaliscina Brazilian genera is provided, and the diversity of the subtribe is discussed in relation to the life habits and communication modalities of Tafaliscina genera.


INTRODUCTION
Tafaliscina Desutter 1988 is a subtribe of Neotropical crickets found on leaves of different heights of shrubs and trees at night. With great morphological differences between its representatives, the subtribe shows a remarkable diversity: from small and slender to a large and robust body, male tegmina with or without stridulatory apparatus, hindlegs with long spines and with or without denticles, distal portion of ovipositor widened and flattened or somewhat rectangular in ventral view (Desutter 1988;Gorochov 2017).
The diversity of forewings in Tafaliscina crickets are particularly impressive. They can be absent or reduced ( Cylindrogryllus Saussure, 1878), elongated with longi tudinal veins (Tafalisca Walker, 1869 andBrazitrypa Gorochov, 2011), or with stridulatory apparatus completely developed (e.g. Adenophallusia de Mello, 1990 andAmblyrhethus Kirby, 1906). Moreover, some species show a stridulatory file without the usual resonant structures present in crickets, i.e., the harp and the mirror, being able to produce a sound, but not the loud, musical ones usually emitted by crickets. Auditory tympana are also variable, from absence to full development. This morphological diversity is unique in a small clade as Tafaliscina, and it raises questions not only about the utility of forewings and different related elements for communication, but also about the evolution of morphology and life habits in those Neotropical crickets.
In the present paper, we increase the knowledge of Tafaliscina diversity describing a new genus, Veredatrypa Campos n. gen. with three new species, and two new species of Tafalisca. We provide an identification key to the Brazilian genera of Tafaliscina, and discuss the diversity of the clade in relation with life habits and communication modalities.
We adopt here the familial classification derived from the molecular phylogenetic of Chintauan-Marquier et al. (2013, 2016. In particular, the family Gryllidae is restricted to one clade of the whole Grylloidea superfamily, which otherwise includes the Phalangopsidae and Trigonidiidae families. According to this topology, the Tafaliscina genera are the sister group of Oecanthus Blanchard, 1845, a result which is supported by additional phylogenetic investigations with molecular and morphological data (Campos, pers. obs.): we consequently modify the classification of Cigliano et al. (2019) to take this result into account.

MATERIAL AND METHODS
The specimens were analyzed, compared and described using a Leica EZ4 stereomicroscope. Drawings of male genitalia and forewings were made under a Leica MZ9.5 stereomicroscope coupled with a camera lucida. The photographs were taken under a Leica MZ16 stereomicroscope with a Leica DFC-420 camera, using the software Leica Application Suite LAS V4.0, with specimens immersed in ethanol 80%. Male genitalia and female copulatory papilla were photographed immersed in hand sanitizer (Su 2016). The photograph of Veredatrypa rosai n. gen., n. sp. (Fig. 1) were taken using a camera Canon T6i with a 100 mm macro lens attached. The distribution map was built with software QGIS 3.4.
Male phallic complexes were removed and treated with aqueous solution 10% KOH for 24 hours to remove muscular tissues and to clarify sclerites and membranes and stored in vial with 80% ethanol with the respective specimen. The female copulatory papillae were also dissected and stored in small vials with ethanol 80% with the respective specimen. Genitalia morphology follows Desutter (1987) and Desutter-Grandcolas (2003). Forewings venation follows Desutter-Grandcolas et al. (2017), modified by Schubnel et al. (2019).   descriPtion In addition to the characters of the genus body. Size medium. Head and pronotum general coloration reddish brown, abdomen dark brown almost black; body covered by brownish bristles.

Male genitalia
Head. Occiput and vertex yellowish brown, marmored medium brown ( Fig. 2A, C, J). Fastigium marmored medium to dark brown ( Fig. 2C). Antennal scape with few bristles on the distal margin in frontal view, light brown, with a medium to dark brown maculae on distal portion in frontal view (Fig. 2C); antennomeres light to medium brown with some isolated antennomeres dark brown. Frons light brown with a median triangular and dark brown macula in frontal view (Fig. 2C). Gena light brown, marmored medium brown in frontal and lateral views. Mandibles yellowish brown. Clypeus light brown, whitish laterally, with two median stripes medium brown; labrum whitish, lower margin yellowish brown (Fig. 2C). Maxillary palpi slightly pubescent, light brown, marmored medium brown; articles 3-5 elongated, article 3 the longest, articles 4 and 5 almost same-sized; apex of article 5 whitish (Fig. 2B, C).
Pronotum. DD yellowish brown; median maculae dark brown, almost black, one close to the cephalic margin, "T" shaped, one in the middle of DD, rounded; caudal margin medium to dark brown. DD cephalic margin almost straight, DD caudal margin slightly convex ( Fig. 2A, B).

legs.
Legs I and II pubescent. FI and FII pale brown, medium to dark brown punctuated; TI and TII darker. FIII pubescent, light brown, medium to dark brown apically; TIII medium to dark brown, yellowish brown ventrally; TI, TII, TIII composition of subapical and apical spurs same as the genus description ( Fig. 2H, I). Basitarsus III pale brown, medium brown ventrally.
Abdomen. Tergites dark brown, almost black; with two bands, thin, median, longitudinal, light brown. Sternites medium brown; each sternite with two dark points, close to the lateral margin. Cerci pilose, light to medium brown.

Male
Metanotum lighter than pronotum DD; metanotal projections: median pair as in Fig. 2Ea, caudal pair as in Fig. 2Eb; clusters of bristles as in Fig. 2E. FWs long, covering hindwings, mainly medium brown; dorsal and lateral field divided by very sclerotized vein (M + CuA vein), yellowish; anal vein area well sclerotized, yellowish; harp crossed by four diagonal veins, not connected to CuPa; first very reduced, third and fourth connected apically; CuPb short; mirror distal cell divided by a longitudinal vein; apical field reduced ( Fig outwards in ventral view; EctF anterior portion concave, forming lateral lobes in anterior and ventral views, posterior portion thinner, upcurved to dorsal, almost reaching MLophi apex, apex inflated. Endophallus: endophallic longer than wide, concave in ventral view; endophallic apodeme as long as EndSc, curved inwards in dorsal and ventral views.

Female
General color as in male. FWs not covering last four abdominal tergites, slightly translucent, medium to light brown (Fig. 2J). Supra anal plate similar to the male, lighter (Fig. 2K). Subgenital plate wider than long, medium brown; posterior margin concave forming two lateral lobes light brown (Fig. 2L). Ovipositor shorter than TIII; dark brown almost black; ovipositor apex slightly serrulated laterally; medium to dark brown (Fig. 2K, L).  descriPtion In addition to the characters of the genus: body. Small to median size (in comparison with V. rosai n. gen., n. sp. and V. fusca n. gen., n. sp); general coloration light brown, medium to dark brown spotted, body covered by sparse bristles, yellowish.
Head. Occiput and vertex light brown, with two bands close to antennal scape and eyes, dark brown (Fig. 5A, C). Antennal scape almost as long as wide, few bristles on the distal margin in frontal view, light brown, with a few medium brown spots in frontal view; antennomeres light brown. Frons and gena light brown, median brown spotted in frontal view (Fig. 5C). Clypeus light brown; superior margin with two vertical and lateral bands medium to dark brown; labrum whitish ( Fig. 5C). Maxillary palpi slightly pubescent, light brown, medium brown spotted; articles 3-5 elongated, almost same-sized.
Pronotum. DD light brown, with medium brown spots; two median maculae dark brown, one close to cephalic margin, one in middle of DD, "V" shaped; caudal margin darker than cephalic margin. DD cephalic margin almost straight, DD caudal margin slightly convex (Fig. 5A, B).
legs. Legs I and II less pubescent than in V. rosai n. gen., n. sp. FI, FII, TI and TII light brown, medium brown punctuated; FIII slightly pubescent, light brown, medium brown punctuated; TIII light brown, apical margin with dorsal maculae medium brown, circles medium brown surrounding subapical spurs; TI, TII, TIII subapical and apical spurs as in genus description (Fig. 5H, I); apical and subapical spurs light brown, apex medium brown. Basitarsus III light brown.

Male
Metanotum light brown; with two latero-anterior protuberances, whitish; without metanotal projections or cluster of bristles (Fig. 5E). FWs long for the genus, not covering last two abdominal tergites, mainly light brown; anal vein area very sclerotized, yellowish; anterior region of diagonal vein very sclerotized; harp crossed with three diagonal veins; third vein reduced, connected apically with second vein, not connected to CuPa; CuPb short; apical field reduced, less cells than V. rosai n. gen., n. sp. (Fig. 5A, D); lateral field with c. 16 diagonal, parallel veins (Fig. 5B). Supra anal plate slightly pubescent, light brown, lateral-anterior margins dark brown; anterior margin almost straight, posterior margin rounded (Fig. 5F). Subgenital plate slightly pubescent, light brown; median region lighter; posterior margin yellowish; anterior margin almost straight, posterior margin convex (Fig. 5G). Male genitalia (Figs 6; 7) Pseudepiphallus: MLophi apex pointed, LLophi very reduced, non-discernible; presence of membrane on distal margin of pseudepiphallic sclerite, with line of strong setae; anterior margin of pseudepiphallic sclerite with small "V" shaped indentation. PsP short, posterior margin slightly concave in ventral view; posterior half wider than anterior half in ventral view; R curved inwards in dorsal and ventral views, poster half flattened, anterior apex curved outwards; almost connected to pseudepiphallic sclerite. Ectophallic invagination: EctAp thin, shorter than in V. rosai n. gen., n. sp., straight in dorsal and ventral views; ventral projections of ectophallic invagination short; EctF median portion cordiform, posterior portion thinner, upcurved to dorsal, not reaching MLophi apex, apex pointed. Endophallus: EndSc longer than wide, concave in ventral view; endophallic curved outwards in dorsal and ventral views.

Female
Unknown.  descriPtion In addition to the characters of the genus body. Small to medium size; head and pronotum general coloration medium brown, abdomen dark brown almost black; body covered by brownish and yellowish bristles.
Pronotum. DD wider than long, with sparse setae, anterior half reddish-brown, posterior half medium to dark brown; DD cephalic margin almost straight, dark brown, DD caudal margin slightly convex (Fig. 8A, B).
legs. Legs I and II slightly pubescent. FI and FII light to medium brown, medium brown punctuated; TI and TII medium brown. FIII slightly pubescent, light brown, medium to dark brown apically; TIII medium to dark brown; apical and subapical spurs yellowish brown, apex and base medium to dark brown; TI, TII, TIII subapical and apical spurs same as in genus description. Basitarsus III very pubescent ventrally; medium brown.

Male
Metanotum cephalic half dark brown, caudal half yellowishbrown; with a pair of latero-anterior protuberances, p ubescent; with two pairs of metanotal projections; one pair of projections in the median line of metanotum, large, elliptical, concave anteriorly; other pair of projections close to caudal margin, median, small, apex pointed; a pair of cluster of bristles pointing to posterior margin, one single cluster, larger, pointing to anterior margin (Fig. 8E). FWs long, covering hindwings, not covering last three abdominal tergites, mainly medium brown, veins light brown; dorsal and lateral field divided by very sclerotized vein (M + CuA vein), yellowish; anal vein area not so as sclerotized as in V. rosai n. gen., n. sp. and V. seca n. gen., n. sp., connection between anal veins more sclerotized, yellowish; harp crossed with four diagonal veins, not connected to CuPa; first very reduced, fourth somewhat reduced, third and fourth connected apically; CuPb short; apical field reduced (Fig. 8A, D); lateral field with c. 14 diagonal, parallel veins (Fig. 8B). Supra anal plate slightly pubescent, yellowish-brown, lateralposterior margin dark brown; anterior margin almost straight, posterior margin rounded (Fig. 8F). Subgenital plate pubescent, anterior half median band reddish-brown, posterior half median maculae greyish brown; anterior margin almost straight, posterior margin rounded, with a median slightly indentation (Fig. 8G). portion concave, forming lateral lobes with spines in anterior and ventral views, posterior portion very thin, upcurved to dorsal, almost reaching MLophi apex, apex inflated. Endophallus: endophallic slightly wider than long, concave in ventral view; endophallic apodeme as long as EndSc, thicker than in V. rosai n. gen., n. sp. endophallic apodeme, curved inwards in dorsal and ventral views.
diAGnosis. -FWs with only parallel and longitudinal veins, not specialized for producing sounds (sometimes bearing only a stridulatory file), absence of tympanum, small protuberances in the dorsal face of fore and mid tibias (not present in all species). Male genitalia: pseudepiphallic sclerite divided in median (bilobate) and lateral lobes, apex of R curved inwards, EctF somewhat membranous, EndSc short, endophallic apodeme very short (sometimes almost no discernible).   projections; dorsal field with reticulated veins between parallel veins; TI and TII with a small, rounded and unpigmented protuberance on apical region of dorsal face, this region without bristles. Male genitalia: MLophi not surpassing LLophi of pseudepiphallus apex in dorsal view, curved outwards in dorsal view; LLophi apex curved inwards forming an angle less than 90°; basis of PsP twisted.
descriPtion General morphology body. Medium to large size; general coloration reddish-brown; body almost entirely covered by small yellowish bristles, except FWs and hindwings.

Male
Metanotum without projections, antero-lateral regions inflated, medio-posterior region somewhat elevated, central region with bristles, lighter than pronotum DD (Fig. 14E). FWs long, covering entire abdomen, medium brown, slightly translucid; left and right FWs with same color and texture; FWs dorsal field bearing a vestigial stridulatory file bearing c. 70 very reduced teeth (Fig. 14D, arrow); apparently without other veins modified to sound production; with c. 7 diagonal and parallel veins, yellow; reticulated veins between parallel veins; lateral field with ten diagonal and parallel veins light yellow (Fig. 14D). Hindwing apex surpassing FWs apex in dorsal view. Subgenital plate pubescent, anterior margin concave, posterior margin rounded, yellowish-brown (Fig. 14G).

DISCUSSION
The morphological diversity within Tafaliscina is remarkable for such a small clade within crickets. As previously mentioned, it mainly concerns the body size and shape, the hind leg armature, the shape and ornementation of female ovipositor, and the forewing development and venation. Its representatives Veredatrypa seca n. gen., n. sp.

Tafalisca duckeana n. sp.
Tafalisca vestigialis n. sp. 500 km N have body lengths ranging from 10 mm (Cylindrogryllus) until 40 mm (Tafalisca), which is quite huge for crickets: this size range could imply different habitats of activity and refuge. Tafaliscina are often found at night on leaves of bushes and trees, but some species of Tafalisca and Brazitrypa, which body shape is cylindrical and elongated, with short and heavy hind legs, have been found inside tree branches both during the day and the night (LDC & PGBSD pers. obs.).
Regarding the importance of forewings in the communication of crickets, the diversity of forewings in Tafaliscina suggests different kinds of communication within the group. Species of Cylindrogryllus are apterous or brachypterous and, when present, their forewings are not modified for sound production; they also lack the auditory tympana, which means that this genus does not use acoustic signals for mating. On the other side, males of Cylindrogryllus generally have some structures on the metanotum under the forewings, that seem to be glandular, and may be used on mating behavior (Walker & Gurney 1967;Prado 2006) and/or chemical communication, as described for Rhaphidophorids (Haley & Gray 2013).
Within Tafalisca, the forewings are symmetrical and welldeveloped, covering the entire abdomen (except for T. mexico Gorochov, 2011 andT. pallidocincta Kirby, 1890) but the veins are not modified for sound-production. In some species, as T. lineatipes Bruner, 1916 and T. vestigialis Campos, Souza-Dias & Nihei n. sp., the PCu vein is curved as a stridulatory file with teeth on the ventral side, possibly being able to produce sound, but the lack of resonant structures (harp, mirror) precludes the emission of loud calls as generally emitted by crickets. Moreover, Tafalisca species do not have auditory tympana, and they probably receive the signals in another way. Female drumming has been observed in rearing conditions (Campos & Desutter-Grandcolas in press), which could reveal vibratory communication in at least some species of this genus. The forewings of Brazitrypa species are similar to that of Tafalisca for venation, but the PCu is not curved and has no stridulatory teeth in the species described so far: these species could be unable to emit sounds. Besides developed not functional forewings for sound production, By contrast, other genera like Adenophallusia, Amblyrhethus, and Veredatrypa Campos n. gen., have forewings with stridulatory apparatus completely developed, as well as auditory tympana: these taxa should be able to a fully acoustic communication. The position of Tafaliscina in the subfamily Oecanthinae based on molecular phylogenetic evidence (Chintauan-Marquier et al. 2013, 2016 still increases the diversity of Tafaliscina direct relatives for body shape and communication modalities (see Otte 1994, Mhatre et al. 2012, 2017. Veredatrypa Campos n. gen. is closely related to the genus Amblyrhethus. The new genus was compared with all Brazilian genera of Tafaliscina: Amblyrhethus Kirby, 1906(Amblyrhethus manni Rehn, 1917, Amblyrhethus natalensis Rehn, 1917, ZOOSYSTEMA • 2020 • 42 (19) are collected mostly through casual findings on plants, top of trees, and inside tree branches. They could also be sensitive to environmental degradation (Chapin et al. 2000). Owing to its diversity mainly based on morphology and forewings, the Tafaliscina clade shows up an excellent model to clarify and understand questions about the evolution of morphology, communication and modification of life habits. Furthermore, all the diversity mentioned above about this group probably leads to a diverse repertoire of behaviors, which is totally unexplored.