The Paroecanthini crickets (Orthoptera: Grylloidea: Gryllidae: Oecanthinae) from French Guiana

ABSTRACT The crickets of the tribe Paroecanthini Gorochov, 1986 (Orthoptera, Grylloidea, Gryllidae, Oecanthinae) present in the collections of the Muséum national d'Histoire naturelle, Paris, are studied, including those collected during the “La Planète Revisitée – Mitaraka 2015″ biological survey. Two species are redescribed, i.e., Tafalisca elongata elongata (Chopard, 1912) n. comb. and Brazitrypa maroniensis (Chopard, 1930) n. comb. Nine species new to science are also described, i.e., Adenophallusia legendrei n. sp., Adenophallusia aratayensis n. sp., Brazitrypa cornuta n. sp., Cylindrogryllus (Apterotrypa) mitarakensis n. sp., Cylindrogryllus (Apterotrypa) guyanensis n. sp., Perutrella septentrionalis n. sp., Stenoecanthus planixiphus n. sp., Tafalisca hugeli n. sp. and Tafalisca ansoi n. sp. The diversity of the Paroecanthini in French Guiana is discussed considering the huge morphological diversity observed in this tribe. The identification key presented elsewhere for Brazilian Paroecanthini (Campos et al. 2020) is extended to Guyanese species.


INTRODUCTION
With more than 16 000 species described (Gargominy et al. 2017), French Guiana, located in northern South America, includes two ecoregions: Guianan Lowland moist forests and Guianan Highlands moist forests (Dinerstein et al. 2017). Over the last decade, the knowledge on insect diversity in this region increased due to the work of taxonomists and the amateur community (Touroult et al. 2018). However, the real richness of French Guiana is far from being fully known (around 20%) (Brûlé & Touroult 2014).
Orthoptera is relatively well-studied in French Guiana (Brûlé & Touroult 2014), but a strong unbalance exists in our knowledge of orthopteran clades. Although Guianese grasshoppers (Caelifera) have been intensively studied and are quite well-known nowadays, as shown by Pocco & Cigliano (2020), few studies and records are available for crickets (Grylloidea). The most abundant species of crickets living in the leaf litter or on tree trunks, and thus relatively easy to collect during the day or at night, have been described only in the 1990's (Desutter-Grandcolas 1992a, b, 1993, apart from the classic works of Chopard (1912Chopard ( , 1920. Things are worse for species living in the canopy, which are very badly known, from the point of view of their systematics and of their biology. As an example, within the Neotropical tribe Paroecanthini Gorochov, 1986, which includes 17 genera and 119 species in the Neotropical region, only two genera and four species are presently recorded from the region (Cigliano et al. 2020). The diversity of this group is clearly underestimated, considering the great diversity of the Neotropical region, the poor sampling of these nocturnal crickets and their probable sensitivity to environmental degradation (Campos et al. 2020).
Here, we studied the Paroecanthini of the Muséum national d'Histoire naturelle (MNHN), Paris, collected in French Guiana over the last 30 years, including during the "Planète revisitée" 2015 Expedition in the Mitaraka area, the large unexplored southwestern corner of French Guiana, close to Suriname and Brazil.
Paroecanthini are presently divided into two subtribes (Paroecanthina Gorochov, 1986and Tafaliscina Desutter, 1988) (Cigliano et al. 2020, which representatives are usually found on leaves of different heights of shrubs and trees during the night (Fig. 1). The morphological diversity of this clade is remarkably huge for such a small cricket clade: from small and slender to large and robust bodies; male genital structures regressed or not; hindlegs long or short, with or without spines; all species, however, show an ovipositor flattened dorso-ventrally (Desutter 1988;Gorochov 2017), except perhaps Stenoecanthus Chopard, 1912, which classification is still uncertain (see below).
The most notable diversity of paroecanthine crickets lays in their forewings, which are deeply involved in communication modalities in crickets: they can be absent or reduced (Cylindrogryllus Saussure, 1878), elongated with longitudinal veins (Tafalisca Walker, 1869 andBrazitrypa Gorochov, 2011), with or without a complete stridulatory apparatus, with significant differences between the shape of the veins (e.g. Adenophallusia de Mello, 1990;Paroecanthus Saussure, 1859;Selvagryllus Otte, 2006;Veredatrypa Campos, 2020). This diversity most certainly shows a wide diversity of behaviors, which are totally unexplored, regarding their modalities of communication (acoustic, vibratory), use of habitats, and sexual behavior. Thus, these crickets may be excellent models to test hypotheses about the evolution of morphology, communication, and life habits (Campos et al. 2020), provided their diversity are well-sampled and their phylogeny attested on a sound ground.
Although we use here the definition of the Paroecanthini tribe by Cigliano et al. (2020), with two subtribes, we follow the phylogenetic results of Chintauan-Marquier et al. (2013, 2016 and classify the Paroecanthini in the Oecanthinae, not in the Podoscirtinae Saussure, 1878 (see also Campos et al. 2020): the relationships between Oecanthus Serville, 1831 and the Paroecanthini are actually well-supported in the molecular phylogeny presented by Chintauan-Marquier et al. (2013, 2016, and supported by further phylogenetic analyses (Campos pers. obs.). The genus Stenoecanthus Chopard, 1912, presently classified within the Podoscirtinae, is consequently moved to the Oecanthinae, but not yet attributed to a definite tribe: its position will be tested with a wide-scale phylogeny of the "Podoscirtinae clade F" of Chintauan-Marquier et al. (2013, 2016 using both molecular and morphological characters. ZOOSYSTEMA • 2020 • 42 (20) Many species of Paroecanthina are available in the MNHN collection. They will not be described here as they do not increase our knowledge of Paroecanthini morphological diversity.

MATERIAL AND METHODS
The specimens were analyzed, compared and described using a Wild M3Z stereomicroscope. Drawings of male genitalia and forewings were also made under a Wild M3Z stereomicro-scope coupled with a camera lucida. The external morphology photographs were taken with a Canon 60D camera with a 100 mm and a 65 mm 1-5x macro lens attached, using the software Helicon Remote. The photographs of male genitalia and female copulatory papilla were photographed immersed in hand sanitizer (Su 2016), under a Nikon SMZ1500 stereomicroscope with a Canon 60D attached.
Male phallic complexes were removed and treated with aqueous solution 10% KOH for few hours to remove muscular tissues and to clarify sclerites and membranes, stored in vial with glycerin and pinned with the respective specimen. The female copulatory papillae were also dissected, stored in small vials with glycerin and pinned 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). The taxa described here have been compared with all Paroecanthini genera (available in MNHN collection), except the genera Eubezverkhovia Gorochov & Izerskyy, 2020 and Mexitrypa Gorochov, 2011 for which only the original description and images were available.
Distribution data of Paroecanthini species from French Guiana were plotted on a map and edited in the software Quantum-gis 3.4 (QGIS Development Team, 2020).

AbbreviAtions
General morphology I, II, III anterior, median, posterior (leg, tarsomere); DD dorsal disc of pronotum; F femur; FW forewing; iad, iam, iav dorsal, median, ventral apical spurs of hind tibia on inner side; L lateral lobe of pronotum; oad, oam, oav dorsal, median and ventral apical spurs of hind tibia on outer side; T tibia; TIII subapical and apical spurs formula indicated inner/outer respectively, counted from distal spurs upwards.

type mAteriAl
The holotypes and allotypes will be deposited in MNHN. Paratypes will be deposited in MNHN and MNRJ as designed in "material examined" and "type material". The specimens collected during the "Our Planet Revisited" Mitaraka 2015 Survey in the Tumuc Humac Mountains, French Guiana ( Fig. 1) Pronotum. DD slightly wider than long, with short bristles, light brown, surrounded by a thin line medium to dark brown; with two dark brown median maculae horizontally drop-shaped in dorsal view. DD cephalic margin slightly concave; caudal margin slightly convex, dark brown ( Fig. 2A, C). LL light brown; ventro-cephalic angle rounded; ventro-caudal angle gradually ascendant in lateral view (Fig. 2B, D).
Wings. FWs covering entire abdomen; FWs somewhat translucent, light brown, with dark brown veins; lateral margin of dorsal field dark brown ( Fig. 2A-D). HWs longer than FWs in dorsal and lateral views.
Legs. Legs I and II with yellowish bristles. FI and FII light brown, apical margin dark brown. TI and TII with small protuberance on dorsal side of proximal region, without bristles (arrow on Fig. 2B); tympana absent. TI and TII medium to dark brown. TI with three apical spurs: two ventral same-sized; one dorsal, inner, almost as long as ventral ones. TII with two ventral spurs, same-sized, two dorsal spurs smaller than ventral ones. FIII longer than TIII; with yellowish bristles, light brown, reddish-brown to dark brown apically; dorsal part with a median dark brown macula, outer side divided by a dark brown stripe (Fig. 2B, D). TIII reddish-brown to dark brown, with yellowish bristles. TIII subapical spurs 5/4, with one spine between each inner and outer pair of successive spurs, three or four spines above uppermost outer subapical spur, two or three spines above uppermost inner subapical spurs. TIII apical spurs 3/3, longer on inner side; inner apical spurs: dorsal longest (iad), median shorter than dorsal (iam), ventral smallest (iav) (iad>iam>iav); outer apical spurs: median longest (oam), dorsal slightly shorter (oad), ventral almost as long as dorsal (oav) (oam>oad>oav). TIII subapical spurs, spines and apical spurs reddish-brown, apex dark brown almost black (Fig. 2B, D). Basitarsus dorsal spines 3/1, apical spine the longest; inner apical spur longer than basitarsus, outer apical spur slightly smaller than outer apical spur. Basitarsus reddish-brown.
Abdomen. Tergites slightly pubescent, medium to dark brown. Cerci pubescent, dark brown, with medium to dark brown setae. Supra anal plate with yellowish bristles, dark brown; posterior margin almost straight (Fig. 2G).
Female genitalia (Fig. 3D-F). Copulatory papilla longer than wide, somewhat cylindrical, with a ventral aperture; posterior half lateral margins slightly curved inwards in dorsal and ventral views, posterior margin rounded in dorsal and ventral views. Anterior margin convex in dorsal view; with a median projection flattened dorso-ventrally, curved upwards in lateral view. description General morphology. Body. Medium to large size, general coloration reddish-brown with black brown spots, covered by bristles.
Head. Occiput and vertex covered by small bristles, reddishbrown. Fastigium as long as wide, pubescent, reddish-brown ( Fig. 5A-C). Three ocelli, median very reduced, almost in the same line as lateral ones in frontal view; lateral ocelli rounded. Frons median region covered by yellowish bristles, reddishbrown (Fig. 5C). Eyes almost as long as wide in lateral view, marginal ommatids light brown, others dark brown almost black ( Fig. 5B, C). Antennal scape longer than wide, inner margin with yellowish bristles, light reddish-brown; antennomeres yellowish with some isolated dark brown ones. Gena reddishbrown in frontal and lateral views. Mandibles medium brown, apex darker. Epistomal suture, clypeus, and labrum medium brown (Fig. 5C). Maxillary palpi pubescent, covered by yellowish bristles; articles reddish-brown, apex light brown; article 5 larger than other articles; articles 5, 4, and 3 almost same-sized.
Wings. FWs covering the entire abdomen; covered by tiny bristles, light brown with several dark brown almost black spots, veins light to medium brown. Lateral field with dark brown veins (Fig. 5A, B). HWs longer than FWs in dorsal and lateral views.
Legs. Legs I and II with yellowish bristles. FI and FII light reddish-brown, apical margins darker. TI and TII with small area not covered by bristles, lighter that TI on dorsal side of proximal region; inner and outer tympana absent. TI and TII dark brown. TI with three apical spurs: two ventral samesized; one dorsal, inner, longer than ventral spurs. TII with three apical spurs: two ventral, same-sized; one dorsal, inner, longer than ventral spurs. FIII longer than TIII; with yellowish bristles, light reddish-brown, darker apically (Fig. 5E). TIII dark brown, with yellowish bristles. TIII subapical spurs 5/4, with three spines between each inner and outer pair of successive spurs, sometimes two spines between first two spurs; seven spines above subapical spurs on both inner and outer TIII margins; apex of TIII subapical spurs slightly curved. TIII apical spurs 3/3, longer on inner side; inner apical spurs: dorsal the longest (iad), median shorter than dorsal (iam),   description General morphology. Body. Medium to large size, general coloration reddish-brown, covered by small bristles.
Head. Occiput and vertex covered by small bristles, reddishbrown. Fastigium as long as wide, pubescent, reddish-brown ( Fig. 8A-C). Three ocelli, the median very reduced, almost in the same line as lateral ones in frontal view; lateral ocelli rounded. Frons median region covered by brownish bristles, reddish-brown (Fig. 8C). Eyes almost as long as wide in lateral view, dorsal ommatids light brown, the others reddish-brown to medium brown ( Fig. 8B, C). Antennal scape longer than wide, inner margin with bristles dark brown, light reddishbrown; antennomeres dark yellow. Gena reddish-brown in frontal and lateral views. Mandibles medium brown, apex darker. Epistomal suture and clypeus medium brown; labrum dark brown (Fig. 8C). Maxillary palpi pubescent, covered by yellowish bristles; articles light reddish-brown, darker apically; article 5 shorter than articles 3 and 4.
Wings. FWs covering the entire abdomen; covered by tiny yellowish bristles; medium brown with light brown veins. Lateral field with light brown veins. (Fig. 8A, B). HWs longer than FWs in dorsal and lateral views.
Abdomen. Tergites slightly pubescent, medium brown. Cerci pubescent, medium brown, marbled dark brown. Supra anal plate dark brown crossed horizontally by a medium light brown band; posterior margin rounded (Fig. 5F).

Male genitalia (Figs 6, 7)
. Pseudepiphallus: pseudepiphallic sclerite apex slightly upcurved in lateral view, anterior margin concave in dorsal view. LLophi thinner than MLophi in ventral view, their apex curved inwards, inner margin of anterior region somewhat rounded in ventral view. MLophi very short, shorter than LLophi in dorsal view, posterior margin rounded. PsP well sclerotized, inclined inwards in ventral view, shorter than pseudepiphallic sclerite, not surpassing its posterior margin; posterior half larger than anterior, inner margins rounded. R elongated, flattened laterally, longer than pseudepiphallic sclerite, anterior half wider than posterior half. Ectophallic invagination: EctAp surpassing the anterior margin of pseudepiphallic sclerite, almost straight in lateral view. Arc not complete, directed posteriorly in dorsal and ventral views; ventral projections of ectophallic invagination shorter than EctAp in ventral view. EctF membranous almost no discernible. Endophallus: EndSc longer than wide, shorter than EctAp, posterior margin rounded, lateral margins folded ventrally; EndAp shorter than EndSc.

Female
Unknown.

Female
Unknown.

Wings.
FWs not covered by bristles, light to medium brown, somewhat translucent; veins light brown (Fig. 11A, C).

Male
Morphology. Metanotum with two median projections rounded in dorsal view; antero-lateral regions inflated, without bristles. First abdominal tergite with a depression resembling an inverted heart in dorsal view (Fig. 11F). FWs covering the abdomen; PCu curved on anterior region, bearing a stridulatory file with c. 56 stridulatory teeth on ventral side. Harp crossed by three diagonal veins, connected to CuPa, second and third connected apically; CuPb short; mirror divided on the middle by a curved vein (Fig. 11A).
Lateral field with c. 12 parallel veins, perpendicular to dorsal field in lateral view. Supra anal posterior margin rounded; plate bearing a median spine, its base enlarged; dark brown, (Fig. 11G, H). Subgenital plate longer than wide, posterior margin straight, strongly pubescent; medium brown, with two longitudinal dark brown bands. Bristles yellow (Fig. 11I).
Male genitalia (Figs 12A-C, 13). Phallic complex not glandular (different from type species). Pseudepiphallus: pseudepiphallic sclerite general shape triangular in dorsal and ventral views, upcurved in lateral view, anterior margin almost straight in dorsal view; MLophi absent. LLophi curved inwards in dorsal view; apex outer lobe longer than inner lobe; apex of both lobes pointed. PsP almost as long as LLophi, not surpassing their tip; forming two lobes, one inner, one posterior; tips of posterior and inner lobes rounded. R elongated, long as pseudepiphallic sclerite, flattened laterally, anterior region curved inwards. Ectophallic invagination: EctAp shorter than LLophi, curved inwards in dorsal and ventral views; surpassing the anterior margin of pseudepiphallic sclerite.
Arc not complete, flattened dorso-ventrally, not connected region slightly curved posteriorly in dorsal and ventral views; ventral projections of ectophallic invagination very short, almost no discernible. EctF membranous, posterior margin rounded. Endophallus: EndSc weakly sclerotized, longer than wide, oval in dorsal and ventral views, anterior and posterior margins rounded.
Female genitalia (Fig. 12D-F). Copulatory papilla longer than wide, downcurved in lateral view, not open ventrally; posterior margin with a median rounded projection in dorsal view; anterior margin almost straight.
Head. Occiput and vertex covered by light bristles, medium brown. Fastigium as long as wide, pubescent, medium brown ( Fig. 14A, C, E). Ocelli absent. Frons with some yellowish bristles on median region, medium to dark brown (Fig. 14E). Eyes longer than wide in lateral view, ommatids dark yellow with some sparse black ones. Antennal scape longer than wide, inner margin with yellowish bristles, medium to light brown, antennomeres medium brown. Gena medium brown in frontal view, light brown in lateral view. Mandibles medium brown, apex darker (Fig. 14E). Epistomal suture dark yellow; clypeus and labrum light to medium brown (Fig. 14E). Maxillary palpi pubescent with light bristles; articles light brown; articles 3-5 almost same-sized; article 5 clavate.

Wings.
FWs covered by tiny bristles, somewhat translucent, medium brown; veins medium brown.

Male
Morphology. FWs covering abdomen but not supra anal plate; PCu vein curved on anterior region, bearing a stridulatory file. Harp crossed by four diagonal veins, connected to CuPa, third and fourth connected apically; CuPb short; mirror divided on the middle by a curved vein (Fig. 14A).
Lateral field with c. 13 parallel veins, perpendicular to dorsal field in lateral view. Supra anal plate bearing a median spine, posterior margin rounded; dark brown, median region black (Fig. 14F, G). Subgenital plate posterior margin straight, strongly pubescent; medium brown, bristles yellow (Fig. 14H).
Male genitalia (Figs 15A-C, 16). Phallic complex not glandular (different from type species). Pseudepiphallus: pseudepiphallic sclerite general shape triangular in dorsal and ventral views, longer than in Adenophallusia legendrei n. sp., slightly upcurved in lateral view, anterior margin concave in dorsal view. MLophi absent. LLophi slightly curved inwards in dorsal view; outer lobe longer than inner lobe; apex of outer lobe rounded, apex of inner lobe pointed. PsP almost same size as LLophi, not surpassing their tip; with two lobes, outer and inner; tips of outer lobe rounded, tip of inner lobe rounded; outer lobe longer than inner lobe. R elongated, shorter than pseudepiphallic sclerite, flattened laterally, anterior region curved inwards. Ectophallic invagination: EctAp almost same size as LLophi, almost straight in dorsal and ventral views; surpassing the anterior margin of pseudepiphallic sclerite. Arc not complete, flattened dorso-ventrally, median apex slightly curved posteriorly in dorsal and ventral views; ventral projections of ectophallic invagination short, as long as half Arc. EctF membranous, posterior margin rounded. Endophallus: EndSc weakly sclerotized, longer than wide, oval in dorsal and ventral views, anterior and posterior margins rounded.
Female genitalia (Fig. 15D-E). Copulatory papilla longer than wide, cylindrical, almost straight in lateral view, without ventral aperture; posterior half thinner than anterior, unpigmented; posterior margin straight, anterior margin convex in dorsal view.

Measurements (mm)
Male ( redescription General morphology Body. Medium size, general coloration medium to light brown, body covered by bristles, except FWs and HWs ( Fig. 17A-C).
Head. Occiput and vertex with bristles, medium brown. Fastigium as long as wide, pubescent, medium brown. Frons light brown, with dark brown band under the eyes (Fig. 17D). Eyes longer than wide in lateral view, ommatids black (Fig. 17B). Antennal scape almost as long as wide, light brown; antennomeres medium brown. Gena light brown in frontal and lateral views. Mandibles medium brown, with dorsal maculae dark brown in frontal view. Epistomal suture light brown; clypeus medium brown; labrum medium brown, darker ventrally (Fig. 17D).
HWs as long as FWs in dorsal and lateral views.
Female genitalia (Fig. 17E-G). Copulatory papilla straight in lateral view, without ventral aperture; posterior margin acuminated. Anterior margin convex in dorsal view and pointed in ventral view.

Male
Unknown.

Measurements (mm)
Female ( Head. Occiput and vertex with bristles, medium brown. Fastigium as long as wide, pubescent, medium brown ( Fig. 18A, C, E). Frons with yellowish bristles, light brown, with medial medium brown macula, and a dark brown band under the eyes (Fig. 18E). Eyes longer than wide in lateral view, ommatids black, dorsal ommatids whitish (Fig. 18B, E). Antennal scape longer than wide, inner margin with yellow bristles, light brown; antennomeres light brown. Gena light brown in frontal and lateral views. Mandibles yellowish brown, apex darker. Epistomal suture and clypeus light brown; labrum medium brown (Fig. 18E). Maxillary palpi slightly pubescent, article 3 longer than articles 4 and 5, article 5 clavate, light brown.
Wings. FWs covering entire abdomen; FWs somewhat translucent, light brown, with dark brown veins (Fig. 18A-D). HWs slightly longer than FWs in dorsal and lateral views.

Male
Morphology. Metanotum with a trapezoid elevation on the middle; antero-lateral regions inflated without bristles. First abdominal tergite with two anteromedial projections, its tips rounded (Fig. 18F). FW dorsal field bearing 7-8 parallel veins, without stridulatory apparatus; PCu vein not curved inwards on anterior region (Fig. 18A), without stridulatory teeth ventrally. Lateral field with c. 8 parallel veins, parallel to dorsal field, medium brown. Subgenital plate longer than wide, posterior margin rounded, strongly pubescent; medium to dark brown with yellow bristles. (Fig. 18H).
Female genitalia (Fig. 19D-F). Copulatory papilla curved downwards in lateral view, without a ventral aperture; posterior margins acuminated, whitish in dorsal view. Anterior margin concave in dorsal view.

Legs.
Legs I and II with yellow and brown bristles. FI and FII light brown. TI and TII light brown. TI with three apical spurs: two ventral same-sized; one dorsal, inner, longer than ventral ones. TII with two ventral spurs same-sized, two dorsal spurs smaller than ventral ones. FIII longer than TIII; with short bristles, light brown (Fig. 21B, D). TIII light brown, with yellowish bristles; region of spurs insertion medium brown. TIII subapical spurs 5/4, with two spines between each inner and outer pair of successive spurs, uppermost spurs with one spine between them, eight or nine spines above the subapical spurs. TIII apical spurs 3/3, longer on inner side; inner apical spurs: dorsal longest (iad), median shorter than dorsal (iam), ventral smallest (iav) (iad>iam>iav); outer apical spurs: median longest (oam), dorsal slightly shorter (oad), ventral almost same size of dorsal (oav) (oam>oad>oav). TIII subapical spurs and spines and apical spurs apex curved, light brown, base medium to dark brown (Fig. 21A-D). Basitarsus dorsal spines 3/1, apical spine the longest; inner apical spur shorter than basitarsus, outer and inner apical spurs samesized. Basitarsus light brown.
Female genitalia (Fig. 22D-F). Copulatory papilla slightly curved downwards in lateral view, without ventral aperture; posterior margin whitish in dorsal view. Anterior margin concave in dorsal view.

Cylindrogryllus (Apterotrypa
description General morphology Body. Small size, general coloration medium to dark brown, body covered by bristles. Head. Occiput and vertex with dark brown bristles, medium brown. Fastigium wider than long, pubescent, medium brown ( Fig. 24A, C, E). Ocelli absent. Frons smooth, light brown (Fig. 24E). Eyes longer than wide in lateral view, ommatids black. Antennal scape longer than wide, inner margin with bristles brown, medium brown; antennomeres light brown with dark brown bands of three articles. Gena medium brown in frontal and lateral views. Mandibles medium brown. Epistomal suture dark yellow; clypeus medium brown; labrum light brown, ventral margin dark brown (Fig. 24E). Maxillary palpi slightly pubescent, article 3 longer than articles 4 and 5, article 5 clavate; articles medium brown, article 5 apex darker.
Male genitalia (Figs 25A-C, 26). Pseudepiphallus: pseudepiphallic sclerite apex straight in lateral view, anterior margin concave in dorsal view. LLophi apex pointed, shorter than PsP, inner margin bearing a membrane. PsP almost not reaching posterior margin of pseudepiphallic sclerite, with two posterior lobes; one dorsal longer, one ventral shorter. R elongated, almost same size of pseudepiphallic sclerite, flattened laterally, anterior region not curved. Ectophallic invagination: EctAp longer than LLophi, straight and inclined outwards in dorsal and ventral views; surpassing the anterior margin of pseudepiphallic sclerite. Arc not complete, inclined posteriorly in dorsal and ventral views; ventral projections of ectophallic invagination very short, shorter than EctAp. EctF membranous almost no discernible. Endophallus: EndSc strongly sclerotized, as long as wide, anterior margin rounded; lateral margins folded ventrally. EndAp short.
Female genitalia (Fig. 25D-F). Copulatory papilla straight in lateral view; dorsal side three times as long as ventral side; posterior margin with a ventral aperture, whitish in dorsal view. Anterior margin convex in dorsal view. Head. Occiput and vertex with yellow and brown bristles; occiput light brown; vertex light brown with two small dark brown maculae. Fastigium wider than long, slightly pubescent, light brown ( Fig. 27A, C, E). Lateral ocelli rounded, above antennal socket in frontal view; median ocellus absent. Frons smooth, dark brown (Fig. 27E). Eyes slightly longer than wide in lateral view, dorsal ommatids medium brown, ventral ommatids dark brown (Fig. 27B, D). Antennal scape longer than wide, inner margin with yellowish bristles, light brown, marbled dark brown; antennomeres dark brown, with some bands of three antennomeres medium brown on proximal region. Gena dark brown in frontal and lateral views. Mandibles dark brown. Epistomal suture dark brown; clypeus dark brown, margins light brown; labrum dark brown (Fig. 27E). Maxillary palpi pubescent with yellowish bristles, light brown, dark brown marbled; article 5 longer than articles 3 and 4, article 4 the smallest; article 5 dorsal margin straight, ventral margin rounded.

Measurements
Pronotum. DD wider than long, with some bristles brown; light brown with a dark brown median macula resembling an inverted "Y". DD cephalic margin slightly concave, with bristles brown; caudal margin somewhat convex, with brown bristles (Fig. 27A, C). LL dark brown; ventro-cephalic angle rounded; ventro-caudal angle gradually ascendant in lateral view (Fig. 27B, D). Wings. FWs not covered by bristles, light to medium brown, somewhat translucent; veins light, medium and dark brown ( Fig. 27A-D). HWs longer than FWs in dorsal view.
Legs. Legs I and II with yellowish and brownish bristles. FI and FII light brown, punctuated medium to dark brown. TI and TII light brown, punctuated medium to dark brown. TI with oval inner and outer tympana. TI with three apical spurs: two ventral, one dorsal, inner. TII with three apical spurs: two ventral, one dorsal, inner. FIII twice longer than TIII; light brown, with a median dark brown band on outer side, apical region dark brown (Fig. 27B, D). TIII light brown, proximal and distal regions dark brown. TIII subapical spurs 4/3, without spines between spurs, three or four spines above the spurs. TIII apical spurs 3/3, longer on inner side; inner apical spurs: dorsal (iad) and median (iam) with same length, ventral the smallest (iav) (iad=iam>iav); outer apical spurs: median longest (oam), dorsal slightly shorter (oad), ventral almost same size of dorsal (oav) (oam>oad>oav). TIII apical and subapical spurs and spines medium to dark brown ( Fig. 27A-D). Basitarsus dorsal spines 5/3, apical spine the longest; inner and outer apical spur same size, shorter than basitarsus. Basitarsus light brown.
Abdomen. Tergites slightly pubescent, medium to dark brown. Supra anal plate posterior margin rounded, dark brown. Cerci pubescent, medium brown, marbled dark brown.

Male
Morphology. Metanotum without projections; median region with cluster of bristles; antero-lateral regions inflated, with bristles; medio-posterior region somewhat elevated (Fig. 27F). FWs covering the whole abdomen; PCu vein curved on anterior region, portion close to lateral field sinuous; bearing a stridulatory file with c. 53 stridulatory teeth on ventral side. Harp crossed by four diagonal veins, connected to CuPa, first one very short, third and fourth connected apically; CuPb short; mirror divided on the middle by one vein curved; apical field developed, with seven columns of cells (Fig. 27A). Lateral field with c. 17 parallel veins, perpendicular to dorsal field in lateral view. Subgenital plate longer than wide, posterior margin almost straight, strongly pubescent; medium brown, with a dark brown median longitudinal band, with yellow bristles (Fig. 27H).
Head. Occiput and vertex with few yellowish bristles; occiput light brown with dark brown band behind eyes; vertex light brown with two median medium brown stripes. Fastigium longer than wide, light brown; antennal socket inner margin bent up in frontal view (Fig. 30A, E). Frons smooth, light brown with a dark brown median macula somewhat triangular (Fig. 30E). Eyes as long as wide in lateral view, ommatids medium brown; crossed by a dark brown horizontal line (Fig. 30B, D). Antennal scape longer than wide, inner margin with whitish bristles, light brown, outer margin dark brown; antennomeres light brown, with medium brown bands of two or three antennomeres. Gena light brown with transversal dark brown band below eyes in frontal and lateral views. Mandibles light brown. Epistomal suture dark yellow; clypeus light brown, upper margin medium to dark brown; labrum light brown (Fig. 30E). Maxillary palpi slightly pubescent with whitish bristles, light brown, articles 3-5 almost same-sized; article 5 apex upcurved.
Pronotum. DD wider than long, with some whitish bristles; light brown with two dark brown lateral short stripes on anterior region. DD cephalic margin concave; caudal margin convex wider than cephalic margin. LL light to medium brown with a dark brown median spot; ventro-cephalic angle slightly rounded; ventro-caudal angle rounded in lateral view (Fig. 30A, C). FWs not covered by bristles, light brown, translucent; veins light and medium brown (Fig. 30A, C). HWs longer than FWs in dorsal and lateral views.
Abdomen. Tergites medium to dark brown with two dark brown median spots on each tergite. Cerci pubescent, longer than TIII, light brown.

Male
Morphology. Metanotum without projections, antero-lateral regions inflated, without bristles (Fig. 30F). FWs covering entire abdomen; PCu curved on anterior region; bearing a stridulatory file ventrally with c. 49 stridulatory teeth. Harp crossed by two transverse veins, connected apically and connected to CuPa; CuPb short; mirror divided on the middle by a curved vein; apical field poorly developed, with four columns of cells (Fig. 30A). Lateral field with c. 17 parallel veins, perpendicular to dorsal field in lateral view. Supra anal plate posterior margin rounded, dark brown (Fig. 30G). Subgenital plate longer than wide, posterior margin straight, pubescent; light brown, with darker median region, bristles whitish (Fig. 30H). weakly sclerotized, posterior margin rounded. Endophallus: EndSc longer than wide, shorter than EctAp, posterior margin concave in dorsal and ventral views, antero-lateral margins folded ventrally; EndAp short.

Female
Morphology. Size and coloration as in male (Fig. 30B, D). FWs covering abdomen; bearing nine longitudinal veins, with small transverse connective veins between them; lateral margin with three dark brown spots in dorsal view. Supra anal plate light brown, median region darker, latero-posterior regions with a dark spot; posterior margin rounded (Fig. 30I) Subgenital plate wider than long, posterior margin rounded; medium brown (Fig. 30H). Last abdominal tergite posterior margin concave, with latero-posterior regions somewhat elongated and inflated in dorsal and lateral views, with bristles ( Fig. 30J, K). Ovipositor flattened dorso-ventrally, downcurved; dorsal valves laminar, laterally expanded, resembling a leaf, involving dorsal region of ventral valves, yellow (Fig. 30M); ventral valves medium brown; apex lateral margins smooth, posterior tip pointed.

DISCUSSION
the pAroecAnthini in the neotropics The Neotropical tribe Paroecanthini is relatively poorly known considering all the extension and diversity of the Neotropical region (Antonelli & Sanmartín 2011;Antonelli et al. 2018). Recently, one new genus and five new species from Brazil and French Guiana were described (Campos et al. 2020). In the present paper, we describe nine new species from French Guiana, a relatively small territory (~83 500 km²) compared to South America (c. 17 840 000 km²), or the whole Neotropical region. According to The Orthoptera Species File (OSF), the tribe comprises 17 genera and 119 species (Cigliano et al. 2020). That means that more than 10% of Paroecanthini diversity has been described in less than one year. As already mentioned for Tafaliscina, a subtribe within Paroecanthini, the diversity of this group is totally underestimated, considering their nocturnal habits, habitat (living in plants and trees canopies), and low population densities. These life habits make Paroecanthini crickets quite hard to find and collect (Campos et al. 2020); for Oecanthinae as for most tropical crickets in general, none of the passive or attractive traps usually used to collect insects is efficient, and only visual collecting is effective to find these species. Thus, we expect that many additional new taxa will be discovered in the next future.
the pAroecAnthini in french guiAnA The morphological diversity of Paroecanthini crickets is remarkable. This diversity includes several aspects of their morphology, and particularly the forewings. In French Guiana we find all the forewing diversity documented in the Paroecanthini, except for the brachyptery as displayed by subgenera Cylindrogryllus (Cylindrogryllus) Saussure, 1878 and Cylindrogryllus (Neometrypus) Desutter, 1988, known from Argentina, Brazil, and Peru. However, this does not mean that they are absent in this region. As already mentioned, this group of crickets is far from well-sampled and the real diversity of this taxa in French Guiana or even South America is far from being comprehensively understood.
Besides forewings diversity, Paroecanthini also varies in the morphology of male metanotum, with projections, bristles and concavities. These structures are involved in mating, producing nutritious secretions which are offered to the females. The female is entertained licking the secretions and sometimes biting structures of male metanotum during copulation, meanwhile the male transfers the spermatophore to female. This behavior was already described for other crickets, including Oecanthus Serville, 1831, Truljalia Gorochov, 1985(Podoscirtinae Saussure, 1878 and phalangopsid crickets (Walker & Gurney 1967;Ono et al. 2004;Prado 2006;Zefa et al. 2008). Usually, metanotal structures are covered by the forewings, as shown in some Paroecanthini crickets (e.g. some species of Tafalisca, Adenophallusia, Angustitrella). The apterous Cylindrogryllus (Apterotrypa) guyanensis n. sp. may however have also metanotal structures, as shown in Fig. 1D of a living male in which there are some structures on the metanotum, but the only holotype available has not been dissected to check this character. A similar condition appears in Cylindrogryllus (Cylindrogryllus), Cylindrogryllus (Neometrypus) and Brazitrypa.
We have also noticed similar structures on the dorsum of the first abdominal tergites in two Guianese taxa: Adenophallusia legendrei n. sp. (Fig. 11F) and Brazitrypa cornuta n. sp. (Fig. 18F). As their positions are almost similar to that of the metanotal structures, it can be hypothesized that these tergal structures could be complementary to the metanotal structures for mating. This is the first time that structures on tergites are documented in the Paroecanthini, but they have been described in phalangopsid crickets, e.g. Anomaloterga mantiqueirae De Mello & Bolfarini, 2010 and Ectecous segregatus Gorochov, 1996(Bolfarini & De Mello 2010Fernandes et al. 2016), and the Odontogryllini Yarrubura nigricephala De Mello & Campos, 2014(Campos & De Mello 2014. By contrast, the Guianese species of Tafalisca lack both metanotal/tergal structures (except Tafalisca hugeli n. sp. with metanotal structures present) and a developed stridulatory apparatus. They may have a vestigial stridulatory file in symmetrical forewings, like in Tafalisca lineatipes Bruner, 1916, which would suggest that these forewings could be used to produce some kind of weak and low sound (Campos et al. 2020), or be a source for vibrational signals, as these species do not have auditory tympana.
This diversity of forms and structures is probably related to different behaviors and life habits. However, no observations on natural history, behavior, or acoustic communication have been described so far for these species. The photo on Fig. 1F shows the mating position in Stenoecanthus planixiphus n. sp., where the female mounts the male, while the male raised its forewings and initiate copulation. These observations mean that at least some Paroecanthini exhibit a cricket mating already documented in other Oecanthinae and Podoscirtinae crickets (Bell 1980;Ono et al. 2004).

the genus StenoecanthuS
Since we consider move Paroecanthini out of Podoscirtinae (Cigliano et al. 2020), grouping it into Oecanthinae according to strong phylogenetic evidence (Chintauan-Marquier et al. 2013, 2016, the genus Stenoecanthus is consequently transferred to this latter subfamily. However, we do not place it in any tribe inside this group, as its relationships remain uncertain. The morphological features of this genus raise many questions about its systematic position. The harp veins are parallel to the PCu (stridulatory file), a putative synapomorphy of the Hapithini Gorochov, 1986 but the ectophallic fold of male genitalia is not well-developed, almost not sclerotized. Finally, the female ovipositor is flattened dorso-ventrally, as in all Paroecanthini. These important characteristics do not allow us to place this genus in one or another Oecanthinae group right now. Besides that, the female of Stenoecanthus planixiphus n. sp. has remarkable features, never described in a cricket before. The ovipositor dorsal valves are laterally expanded, dorso-ventrally flattened and reinforced with transverse veinlets, resembling a leaf, and covering the ventral valves (Fig. 30M). The shape of ovipositors is generally related to the site of oviposition (Gwynne 2001) and Stenoecanthus planixiphus n. sp. particular ovipositor may facilitate oviposition inside the parenchyma of leaves for example, as shown on a photograph of a female (Fig. 1E). Species ovipositing in fresh leaves are known in katydids, but these species usually have a laterally flattened, curved ovipositor (Gwynne 2001).
The second original feature of the female of S. planixiphus n. sp. is a morphological modification in the last abdominal tergite (Fig. 30I, J, K). These structures are expanded laterally and have some bristles, indicating that it could be a glandular structure, which function is not known today. distribution of guyAnese pAroecAnthini None of the Paroecanthini species described from French Guiana are documented in other regions, except for Tafalisca vestigialis, which is also from Brazil Northern Region, State of Para (Campos et al. 2020). The lack of sampling in the Guianese region prevent us to consider those species as endemics to French Guiana.
From the point of view of the genera, we observed very diverse situations, which call for additional sampling in the whole Neotropics. As a few examples, Tafalisca has a wide distribution in the northern Neotropics, being present in the Amazonian region, Central America, Caribbean, Mexico and south of Florida. Except the Amazon Rainforest, there are no records of this genus in other South American regions. In the same habitat, we found Brazitrypa in the Atlantic Forest. Adenophallusia shows a distribution more restricted than Tafalisca, being known presently in Northern South America only, with Adenophallusia naiguata de Mello & de Camargo e Mello, 1996 recorded from Naiguata, Venezuela, Adenophallusia legendrei n. sp. and Adenophallusia aratayensis n. sp. described herein from French Guiana. Finally, Perutrella is recorded from Peru by its type species, Perutrella originalis Gorochov, 2011, and we attest its presence in French Guiana with the description of Perutrella septentrionalis n. sp. (Fig. 33).

CONCLUSION
The classification of Oecanthinae, in which we include the Paroecanthini (Campos et al. 2020), is somewhat unorganized today. The molecular data presented in Chintauan-Marquier et al. (2013, 2016 helped to clarify the situation and lead for a complete reanalysis of the "traditional classification" of several cricket groups, considered as families/subfamilies according to the authors, i.e. Podoscirtinae, Hapithinae Gorochov, 1986, Euscyrtinae Gorochov, 19855, Pentacentrinae Saussure, 1878, and Oecanthinae (Bruner 1916Chopard 1949Chopard , 1968Desutter 1988;Gorochov 1986). The molecular studies of Chintauan-Marquier et al. (2013, 2016, who sampled the whole cricket clade using 206 terminals, were however not based on enough taxa to test the monophyly and relationships of all these groups, which were gathered in a "Clade F" without formal classificatory proposal. A phylogeny based morphological and molecular data is being developed to test the monophyly of these groups and to facilitate future evolutionary studies of this diverse clade of crickets.