Temnomastax Rehn & Rehn, 1942

Olivier, Renan S., Pujol-Luz, Cristiane V. A. & Graciolli, Gustavo, 2019, Review of Temnomastax Rehn & Rehn, 1942 (Orthoptera, Caelifera, Eumastacidae, Temnomastacinae), Zootaxa 4593 (1), pp. 1-78 : 9-16

Temnomastax Rehn & Rehn, 1942

Masyntes [in part] Burr 1899: 95 (identification key), 273 (original description); Burr 1904: 15 (identification key), 17 (species of the genus); Bruner 1911: 6 (identification key for species).

Temnomastax Rehn & Rehn 1942: 12 (identification key for species), 9 (original description). Type-species, Masyntes tigris Burr (orig. des., Burr, 1899: 276); Rehn 1948: 77, 85, 116, 136 (citation); Rehn & Grant Jr. 1958: 302 (citation), 315 (description of phallic complex); Descamps 1971: 102 –104 (comparative analyzes of morphology), 1973a: 219, 220 (citation), 1973b: 964 (citation), 1979: 144 (citation); Otte 1994: 48 (citation); Xiangchu et al. 1996: 839 (citation); Rowell & Flook 1998: 154 (phylogeny); Mojica & Fagua 2006: 206, 210 (misidentification); Matt et al. 2008: 46, 47, 49, and 50 (phylogeny); Silva et al. 2014: 1504 (citation); Olivier 2014: 457 (citation), 458 (identification key for species), 461 (citation), 2017: 235 (citation); Olivier & Aranda 2017: 2 (citation), 2018: 269 (citation), 270 (identification key for five species).

Etymology. From Greek. Tέµνω, ‘stump’, in allusion to the relatively truncate fastigium in dorsal view + µάσταξ (mastax), ‘jaws’, ‘what chews’, commonly used as suffix for Eumastacidae genera.

Diagnosis. Small to medium size (♂ 14.84 mm– 23.90 mm, ♀ 19.44 mm– 28.94 mm). Unlike the Eumastacopini , Temnomastax has a very protruding fastigium, without two submammilliform nodes and the ectophallus is completely membranous, without spines, apophysis or sclerification. From Eutemnomastax differs by the following combination of features: brachypterous or macropterous ( T. beni ). A complete black ring in the ⅓ distal of metafemur (absent in T. beni only). Presence of a dorsal spine on the outer margin of the 1 st metatarsomere (absent in T. beni only). Lateral regions of abdominal tergites with blue maculae. 7 th and 8 th male abdominal tergites and 7 th female abdominal tergite with distinct color, usually yellow or red. Male cercus laterally compressed or curved inward (tapered in T. beni only). LPjEp directed backward and downward. Endophallic plate with slight curvature in lateral view. Posterior margin of female subgenital plate subtriangular or arched with spines on the posterior margin (absent in T. hamus ).

Redescription. Male. Head: Lateral margins from ventral margin of eyes to mandibular joints parallel to (narrow) or divergent from (subtriangular) each other in frontal view ( Figure 4 ). Antennae with 12 antennomeres, an antennal organ in the 4 th antennomere from the apex to the base. Fastigium protruding in lateral view ( Figure 5 ) rounded and laterally little expanded in dorsal view. Eyes ellipsoid, from slightly to prominently protruding in frontal view ( Figures 4 and 5 ). Facial median carinae well marked, not continuous with lateral margins of fastigium. Fastigium-occipital junction convex in lateral view (concave in T. beni ). Cervical membrane with two distinct dorsal carinae. Thorax: Pronotum: Slighty sellate in lateral view ( Figures 6 A–H). Posterior margin of pronotal disk rounded or centrally fissured ( Figures 6 a–h). Central carina well marked, lateral carinae absent and dorsal-lateral junction rounded. Ventroposterior angles of lateral lobes acute and somewhat directed backward. Wings: Brachypterous or macropterous; tegmina elliptical or ellipsoidal with rounded apex ( Figure 7 A–G). Venation simple, shorter tegmina with few and more sinuous veins, larger tegmina with bifurcations on the R or M veins ( Figures 7 A–G). Membranous wings always present, with two to five lobes when extended (remigium + anal lobes) ( Figures 7 a–g). Venation simple with at most six anal veins. Shorter membranous wings often with two or more vestigial anal veins. Largest wings with Rs2 present ( Figures 7a, e ). Legs: Profemora and mesofemora with two distinct dorsal carinae, both ending in small apical spines. Metafemora with three dorsal carinae and four ventral carinae; five apical spines present, three in the end of each dorsal carina and two laterally below these three. Dorsal face of metatibiae with small outer spines, inner spines small proximally, large medially, and medium distally; apex of metatibia with two inner spurs and two outer spurs, the dorsal larger than the ventral ones and the inner dorsal larger than the outer dorsal. A dorsal spine on the outer margin of the 1 st metatarsomere always present (absent in T. beni ). Metatibiae and metatarsomeres more hirsute than remainder of the thorax and legs. Tarsal claws unequal, without internal serration. Pulvilli well developed. Abdomen: Posterior-ventral region of the 9 th and 10 th abdominal tergites rounded and slightly directed backward ( Figures 8 A–H). Posterior-dorsal margin of the 10 th abdominal tergite centrally arched (concave) and with a membranous tissue connected to epiproct ( Figures 8 a–h). Epiproct entirely narrow ( T. descampsi sp. nov. only) or trigonal with acute apex, lateral margins folded downward in different degrees and a central membranous area on the proximal region ( Figures 8 a–h). Cerci sub-cylindrical (in T. beni only), laterally compressed in the medio-apical region or completely compressed laterally; apex with varying degrees of inward curvature ( Figures 9 A–G). Distal region of subgenital plate membranous with variable rugosity and from absent to three carinae present in axial view, posterior margin more sclerotized and with different degrees of backward projection (slightly, moderately or strongly) ( Figures 8 A–H). Pallium evident in dorsal view ( Figures 8 a–h). Cerci, epiproct and paraprocts more hirsute than other parts of abdomen. Phallic complex ( Figures 10 and 11 ): Epiphallus: Large central membranous area presents (MbEp), slightly concave in the antero-medial region when viewed dorsally. Lateral portions (LREp) more sclerotized, not connected in the anterior region and with different shapes in the medio-posterior region in dorsal view. Posterior region rectilinear, curved upward or downward ( T. descampsi sp. nov.) in lateral view. Lateral projections (LPjEp) directed backward and downward, apex curved inward. True lophi absent. Spines absent. Endophallus ( Figure 12 ): Endophallic plate (EnP) Ŋ-shaped, outer margin of the anterior region bi- or trilobed, inner arc circular or in some cases seemingly more triangular, branches almost parallel, slightly convergents or divergents in ventral view, with slight curvature in lateral view, outer margins of lateral branches irregular and sometimes with marked ripples; insertion of the spermatophore sac in the anterior portion of the endophallic plate; ejaculatory sacs with distinct shapes, in few cases slightly project forward or commonly projected backward. Ectophallus: Large, globose, completely membranous and with two lobes well evident in ventroaxial view. Ectophallic sclerite (EcS) from slightly to much projected backward, without sclerification, spines or apophysis and with medio-basal portion covered by part of the posterior bilobed portion. Genital aperture (GnAp) dorsal-apical.

Female. Antennae with 12 antennomeres, but shorter than in male. Fastigium little more expanded laterally than in male in dorsal view. Brachypterous with wings shorter than in male, proportionally to the body length. Posterior-ventral regions of the 8 th abdominal tergite smooth or with cuticular striations ( Figure 13 ). Epiproct linguiform with narrow or rounded apex, carinae on the dorsal face present and with different shapes ( Figure 14 ). Length of paraprocts shorter or equal to the epiproct. Cerci sub-cylindrical, tapered and shorter than the epiproct ( Figures 9H and 14 ). Subgenital plate: lateral regions of subgenital plate partially covered ( Figures 15B, E, F, G, H ) or not ( Figures 15A, C, D ) by the ventral portions of the 8 th abdominal tergite; posterolateral regions of subgenital plate exceeding or not the length of the 8 th abdominal tergite in lateral view ( Figure 13 ); posterior margin of subgenital plate subtriangular or arched (concave or convex), spines present or absent (in T. hamus ); posterolateral regions of subgenital plate smooth or with cuticular striations ( Figure 15 ). Ovipositor: soil-type. Dorsal valves: Dorsal outer margin with larger teeth in the medio-apical regions; ventral face almost rectilinear in lateral view. Ventral valves: Dorsal face sinuous, with a median concavity in lateral view; ventral outer margins with four robust teeth in the medio-apical region, except by the apical tooth ( Figure 13 ).

Chromatic characters. Live specimen (except for T. beni ): Lateral regions of head with metallic blue stripes between the posterior-ventral margins of the eyes and occipital suture; small portions above the mandibular joints with the same color. Occiput with two metallic blue longitudinal maculae. Margin of fastigium ranging among black, yellow or metallic blue. Scape and pedicel yellow. 1 st antennomere of flagellum yellow and the others black. Eyes black. Frons, clypeus, and labrum yellow (Tigris group) or frons metallic blue and clypeus and labrum yellow (Latens group). Mouthparts yellow; in some species the central regions of mandibles on the outer face present blue maculae; mandibular joints black. Dorsal carinae of cervical membrane yellow and sometimes contiguous regions with the same color. Cervical membrane predominantly black with ventral portions of lateral regions blue. Pronotal disk with a wide longitudinal yellow stripe (usually in Tigris group) or sometimes with a mixture of black and blue maculae and yellow spots (usually in Latens group). Lateral lobes of pronotum with a wide dorsal black stripe and a narrow ventral metallic blue stripe; some species (e.g. T. hamus ) with small yellow maculae on ventroposterior regions ( Figure 16A ). Scutellum yellow. Thoracic tergites with small dorsal regions black and other parts metallic blue. Thoracic sternites yellow (usually in Tigris group) or metallic blue (usually in Latens group). Tegmina opaque, black with blue and yellow veins on dorsal surface. Membranous wings blackish, sometimes with light marks in some cells. Metafemur dark yellow, ⅓ distal with a complete black ring circled by two yellow rings, apex black ( Figure 16B ). Dorsal and ventrolateral carinae of metafemur black and ventro-central carinae yellow. Metatibiae blackish, ⅓ proximal with a small lighter portion (yellowish) distally. Tarsi and pulvilli dark yellow. Tarsal claws yellow, apex black. Abdominal tergites black with yellow spots dorsally and large metallic blue maculae laterally; 7 th and 8 th abdominal tergites of male and 7 th abdominal tergite of female predominantly yellow or red (in T. monnei sp. nov. only) ( Figure 16B ); 8 th abdominal tergite of female black. Abdominal sternites yellow (1 st abdominal sternite bluish in some species). Epiproct black. Cerci black or dark brown. Male subgenital plate yellow with black details in the lateral regions of posterior margins. Female subgenital plate yellow or red (in T. monnei sp. nov. only), posterior-medial region with a darker shade in some species. Ovipositor yellow, yellowgreenish or reddish, apex of the teeth dark brown or black.

Egg. Bacilliform; anterior extremity not uniformly rounded and with remnants of attachment to the ovaries ( Figure 17A ); posterior extremity rounded with an annular constriction on the ¼ distal ( Figures 17B , C—arrows); micropyle not visible (possibly absent?); exochorion orange, completely covered by hexagonal chorionic sculptures of different sizes with protruding edges and concave center ( Figures 17b, c , c’). Measurements (mm) (n=71): Height: 4.17–5.07 (4.63), Greater width: 0.79–1.23 (0.99). Disposition of eggs: An egg centrally and other arranged obliquely around this in lateral view. The eggs are held together by the lateral walls, which presents a sticky translucent substance. Egg pods are not enveloped by foam ( Figure 17A ).

Distribution. Temnomastax is widely distributed in the Cerrado domain. In Brazil, it occupies the region between south of Pará and Mato Grosso do Sul and south-central of Minas Gerais and west of Mato Grosso. Also occurs in Paraguay, north of Argentina, and southeast and north of Bolivia.

Remarks. Chromatic characters in dry specimens ( Figures 18 and 19 ): After mounting of specimens with entomological pin, some chromatic characters may change. These changes vary according to the method of killing, such as alcohol, ethyl acetate or low temperature. Herein we present, in general, the changes noticed among all studied specimens.

Regions of the body with black color, such as eyes, mandibular joints, tegmen and wings, and some portions of pronotum, often become brown or dark-brown. Regions with uniform yellow color, such as face (in species belonging to Tigris group), 7 th and 8 th abdominal tergites of males and 7 th of females (in almost all species), abdominal sternites, and subgenital plate all have the color preserved. Regions with red color (exclusively for T. monnei sp. nov.), such as female subgenital plate, and 8 th and 9 th (subgenital plate) male abdominal sternites which become orange or light-red. When the yellow color presents as spots, for example, on the dorsal face of abdomen and pronotum, they may become pallid or whitish. Some specimens belonging to the Latens group have pronotal disk with a mixture of colors (black, blue, and yellow spots). Additionally, they may have a lighter longitudinal band similar to that observed in the Tigris group, but not too yellow and well delimited. Predominant metallic blue regions, such as lateral regions of head, parts of pronotum, thorax, and abdomen barely maintain their color and usually become light-yellow or whitish. Species that belong to the Latens group have regions with metallic blue, such as face and thoracic sternites, but may lose the metallic luster and darken.

Color of nymphal instars relative to host plants: Nymphs are commonly found on leaves of small-sized leguminous plants (young plants). The green color of the nymph during the 1 st and 2 nd instars is very similar to the leaflets of host plants ( Fabaceae ). The position of legs when resting, overlapping the leaflets, makes it difficult to see them, so that they go unnoticed ( Figures 20A, E ). In addition to their green coloration, developing nymphs of 3 rd and 4 th instars have a light brown stripe on the dorsum that matches the leaf petiole of leguminous plants with the same color and is thus used for camouflage ( Figures 20 B–D). Subadult grasshoppers have a well-marked pattern of maculae on the lateral regions of body, mainly in the abdominal tergites; however, the colors are still not the same as those of adult individuals, and they exhibit more yellowish and light tones.

Chromatic characters in T. beni: The coloration of Temnomastax is one of the most important features for its identification; however, in T. beni , this pattern is not maintained, and for this reason, the chromatic description of this species is given together with its morphological description.

Morphological characters—Temnomastax vs. Eutemnomastax: When describing Eutemnomastax, Descamps (1979) highlighted several characteristics that separated this genus from Temnomastax , one being the presence of spines on the posterior margin of female subgenital plate. Alone, the presence of spines cannot be used as a consistent character to separate these two genera since of the nine Temnomastax species, only T. hamus has a completely smooth posterior margin of subgenital plate. In addition to considering the presence or absence of spines, a more important and consistent character is the shape of the posterior margin of female subgenital plate, which is always subtriangular in Eutemnomastax and either subtriangular or arched in Temnomastax ( Figure 15 ).

Figures

FIGURE 4. Head of males and females in frontal view. A—Temnomastax borellii (Giglio-Tos, 1897); B—Temnomastax hamus Rehn & Rehn, 1942; C—Temnomastax latens Rehn & Rehn, 1942; D—Temnomastax monnei Olivier sp. nov.; E— Temnomastax descampsi Olivier sp. nov.; F—Temnomastax otavioi Olivier sp. nov.; G—Temnomastax ricardoi Descamps, 1973; H—Temnomastax tigris (Burr,1899). Scale bar = 2 mm (Female E—MNHN-EO-CAELIF 4052, photo by Simon Poulain, CNRS-MNHN).

FIGURE 5. Head of males and females in lateral view. Black spot = absence of vision plane. A—Temnomastax borellii (Giglio-Tos, 1897); B—Temnomastax hamus Rehn & Rehn, 1942; C—Temnomastax latens Rehn & Rehn, 1942; D— Temnomastax monnei Olivier sp. nov.; E—Temnomastax descampsi Olivier sp. nov.; F—Temnomastax otavioi Olivier sp. nov.; G—Temnomastax ricardoi Descamps, 1973; H—Temnomastax tigris (Burr, 1899). Scale bar = 2 mm.

FIGURE 6. Pronotum of males and females in lateral (uppercase letters) and dorsal (lowercase letters) views. Black spots = absence of vision plane. A—Temnomastax borellii (Giglio-Tos, 1897); B—Temnomastax hamus Rehn & Rehn, 1942; C— Temnomastax latens Rehn & Rehn, 1942; D—Temnomastax monnei Olivier sp. nov.; E—Temnomastax descampsi Olivier sp. nov.; F—Temnomastax otavioi Olivier sp. nov.; G—Temnomastax ricardoi Descamps, 1973; H—Temnomastax tigris (Burr,1899). Scale bar = 1 mm.

FIGURE 7. Tegmen (uppercase letters) and membranous wings (lowercase letters) of males of Temnomastax Rehn & Rehn, 1942. A—Temnomastax latens Rehn & Rehn, 1942; B—Temnomastax hamus Rehn & Rehn, 1942; C—Temnomastax ricardoi Descamps, 1973; D—Temnomastax borellii (Giglio-Tos, 1897); E—Temnomastax tigris (Burr, 1899), specimen from Bodoquena region; F—Temnomastax descampsi Olivier sp. nov.; G—Temnomastax otavioi Olivier sp. nov. Ra—anterior radius; Rp—posterior radius; Sc-cell—formed by the division and posterior union of the subcosta (Sc). Apostropheintraspecific variation. Scale bar = 2 mm.

FIGURE 8. Abdominal apex of males in lateral (uppercase letters) and dorsal (lowercase letters) views. A—Temnomastax borellii (Giglio-Tos, 1897); B—Temnomastax hamus Rehn & Rehn, 1942; C—Temnomastax latens Rehn & Rehn, 1942; D— Temnomastax monnei Olivier sp. nov.; E—Temnomastax descampsi Olivier sp. nov.; F—Temnomastax otavioi Olivier sp. nov.; G—Temnomastax ricardoi Descamps, 1973; H—Temnomastax tigris (Burr, 1899). Scale bar = 2 mm.

FIGURE 9. Cercus of male (A–G) and female (H) of Temnomastax Rehn & Rehn, 1942. Columns from left to right show dorsal view, external lateral view, ventral view, and internal lateral view. Black spots = absence of vision plane. A— Temnomastax borellii (Giglio-Tos, 1897); B and H—Temnomastax hamus Rehn & Rehn, 1942; C—Temnomastax latens Rehn & Rehn, 1942; D—Temnomastax descampsi Olivier sp. nov.; E—Temnomastax otavioi Olivier sp. nov.; F—Temnomastax ricardoi Descamps, 1973; G—Temnomastax tigris (Burr, 1899). Scale bar = 0.5 mm.

FIGURE 10. Phallic complexes of species belonging to Latens group. Uppercase letters—dorsal view, lowercase letterslateral view. A—Temnomastax borellii (Giglio-Tos, 1897); B—Temnomastax hamus Rehn & Rehn, 1942; C—Temnomastax latens Rehn & Rehn, 1942; D—Temnomastax monnei Olivier sp. nov. Scale bar = 1 mm (for dorsal view only).

FIGURE 11. Phallic complexes of species belonging to Tigris group. Uppercase letters—dorsal view, lowercase letterslateral view, apostrophe—dorsal view of the posterior region of epiphallus inclined 45° downward. A—Temnomastax descampsi Olivier sp. nov.; B—Temnomastax otavioi Olivier sp. nov.; C—Temnomastax ricardoi Descamps, 1973; D— Temnomastax tigris (Burr, 1899). MbOrEp—Membranous ornamentation on lateral projections of epiphallus. Scale bar = 1 mm (for dorsal view only).

FIGURE 12. Schematic drawings of the endophallic plate in ventral view (A–I) and endophallus in lateral view (a–h). A— Temnomastax borellii (Giglio-Tos, 1897); B—Temnomastax hamus Rehn & Rehn, 1942; C—Temnomastax latens Rehn & Rehn, 1942; D—Temnomastax monnei Olivier sp. nov.; E—Temnomastax descampsi Olivier sp. nov.; F—Temnomastax otavioi Olivier sp. nov.; G—Temnomastax ricardoi Descamps, 1973; H—Temnomastax tigris (Burr, 1899). I and i— Intraspecific variation in the anterior outer margin of the endophallic plate in T. tigris; I—specimens from Corumbá region; ispecimens from Bodoquena region. Scale bar = 0.5 mm.

FIGURE 13. Abdominal apex of females in lateral view. A—Temnomastax borellii (Giglio-Tos, 1897); B—Temnomastax hamus Rehn & Rehn, 1942; C—Temnomastax latens Rehn & Rehn, 1942; D—Temnomastax monnei Olivier sp. nov.; E— Temnomastax descampsi Olivier sp. nov.; F—Temnomastax otavioi Olivier sp. nov.; G—Temnomastax ricardoi Descamps, 1973; H—Temnomastax tigris (Burr, 1899). Scale bar = 2 mm. (E—MNHN-EO-CAELIF 4052, photography by Simon Poulain, CNRS-MNHN).

FIGURE 14. Abdominal apex of females in dorsal view (cerci, paraprocts, and epiproct). A—Temnomastax borellii (Giglio- Tos, 1897); B—Temnomastax hamus Rehn & Rehn, 1942; C—Temnomastax latens Rehn & Rehn, 1942; D—Temnomastax monnei Olivier sp. nov.; E—Temnomastax descampsi Olivier sp. nov.; F—Temnomastax otavioi Olivier sp. nov.; G— Temnomastax ricardoi Descamps, 1973; H—Temnomastax tigris (Burr, 1899). Scale bar = 1 mm. (E—MNHN-EO-CAELIF 4052, photography by Simon Poulain, CNRS-MNHN).

FIGURE 15. Female subgenital plates in ventral view. A—Temnomastax borellii (Gilio-Tos, 1897); B—Temnomastax hamus Rehn & Rehn, 1942; C—Temnomastax latens Rehn & Rehn, 1942; D—Temnomastax monnei Olivier sp. nov.; E— Temnomastax descampsi sp. nov.; F—Temnomastax otavioi Olivier sp. nov.; G—Temnomastax ricardoi Descamps, 1973; H— Temnomastax tigris (Burr, 1899), specimen from Corumbá region. Scale bar = 2 mm.

FIGURE 16. A—Female of Temnomastax cf. hamus Rehn & Rehn, 1942 from Morro Agudo, São Paulo, Brazil; note the yellow macula in the ventroposterior region on lateral lobe of the pronotum. B—Couple of Temnomastax hamus Rehn & Rehn, 1942 from Campo Grande, Mato Grosso do Sul, Brazil, in captivity; note the different body dimensions between the sexes and the distinct color (yellow) on the 7 th and 8 th abdominal tergites of male and 7 th abdominal tergite of female. (Photography by Marcos Cesar Campis (A) and Lívia Cordeiro (B)).

FIGURE 17. Eggs of Temnomastax Rehn & Rehn, 1942. Photos A and B were obtained under a conventional stereomicroscope, and C was obtained under a Scanning Electron Microscope (SEM). A—egg pod; B and C—individualized egg; b, c, and c’—details of the chorionic sculptures. Arrows indicate the annular constrictions on the ¼ posterior extremities. Zoom: C (30x), c (80x), and c' (430x). Scale bars: A and B = 1 mm, b = 0.2 mm, C = 500 µm, c = 200 µm, and c’ = 50 µm. (SEM photos by Instituto Oswaldo Cruz–RJ).

FIGURE 18. Habitus of species belonging to Latens group. A—Temnomastax borellii (Giglio-Tos, 1897); B—Temnomastax hamus Rehn & Rehn, 1942; C—Temnomastax latens Rehn & Rehn, 1942; D—Holotype and Allotype of Temnomastax monnei Olivier sp. nov. Scale bar = 1cm.

FIGURE 19. Habitus of species belonging to Tigris group. A—Paratype and Allotype of Temnomastax descampsi Olivier sp. nov.; B—Paratype and Allotype of Temnomastax otavioi Olivier sp. nov.; C—Temnomastax ricardoi Descamps, 1973; D— Temnomastax tigris (Burr, 1899) from Corumbá region. Scale bar = 1 cm. (Female A—MNHN-EO-CAELIF 4052, photography by Simon Poulain, CNRS-MNHN).

FIGURE 20. Nymphs of Temnomastax Rehn & Rehn, 1942 on host plants (Fabaceae) in the state of Mato Grosso do Sul, Brazil. A and E—Temnomastax ricardoi Descamps, 1973 on Anadenanthera falcata from Ribas do Rio Pardo City; B, C, and D—Temnomastax hamus Rehn & Rehn, 1942 on Anadenanthera falcata from Terenos City.