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Subtribe † Pherodactylina Cadena-Castañeda & He n. subtr. Diagnosis. Slender body covered with several hairs. Head globose, slightly flattened dorsoventrally. Pronotum moderately elongated, 1.5 times as long as broad, rectangular. Forelegs fossorial type, with the distal portion of fore tibia laterally compressed, and three strong movable apical spurs arranged only on inner side of tibia. Hind tibia moderately flattened, slightly shorter than femur; with small spines on dorsal margin and six spurs apically, three inner spurs and three outer ones nearly equal in length and shorter than first tarsomere, upper inner spur significantly longest. Taxa included. † Pherodactylus Poinar, Su & Brown, 2020 (type genus) and † Burmagryllotalpa Wang et al., 2019. Remarks. This new subtribe is proposed to group fossil taxa only, in which some are originally described as mole crickets. After morphological analysis in this study, we conclude that they are true crickets with short dorsal spines on the hind tibia. Possessing serrated hind tibia is one of the characters that characterize Sclerogryllini taxa. Comparison. † Pherodactylina n. subtr. is included in Sclerogryllini, due to the structure of the hind tibia, in addition to the shape of the pronotum, which is more elongated in comparison with the field crickets of other Gryllinae tribes. The venation of the tegmina of the male of † P. rectanguli n. comb. resembles the venation of known Sclerogryllus males (Fig. 2). The same happens in the shape of the ovipositor in females, which presents a similar structure in its length with respect to the hind femur and the shape of the apex of the valves. Sclerogryllina n. stat. differs from † Pherodactylina n. subtr., due to its more robust appearance, glabrous body, and less pubescence. The species of † Pherodactylina n. subtr. are yellowish-brown in color, while Sclerogryllus species are almost black. Sclerogryllina n. stat. does not possess the fore tibiae with modification to dig, and it possesses three small apical spurs on each margin (outer and inner). On the other hand, the new tribe does not possess apical spines on the outer margin. Finally, Sclerogryllina n. stat. has conspicuously delineated hind femur chevrons traces, which does not appear in † Pherodactylina n. subtr., Published as part of Cadena-Castañeda, Oscar J., Liu, Yi-Jiao, Yu, Zhe-Yuan, Hu, Tian-Hao, Wu, Shi-Yang & He, Zhu-Qing, 2023, Are the recently described fossil Mole Crickets of Myanmar amber real gryllotalpids? (Orthoptera: Gryllotalpidae & Gryllidae), pp. 48-64 in Zootaxa 5311 (1) on page 52, DOI: 10.11646/zootaxa.5311.1.2, http://zenodo.org/record/8090321, {"references":["Poinar Jr., G., Su, Y. N. & Brown, A. E. (2020) A new genus of crickets (Orthoptera: Gryllidae) in Mid-Cretaceous Myanmar Amber. Biosis: Biological Systems, 1 (1), 33 - 38. https: // doi. org / 10.37819 / biosis. 001.01.0049"]}

† Pherodactylus rectanguli (Xu, Fang & Wang, 2020) n. comb. Figs. 1–6 = † Chunxiania fania Xu, Wang & Fang, 2022 n. syn. = † Tresdigitus gracilis Jiang, Xu, Jarzembowski & Xiao, 2022 n. syn. General description. Male (based on specimens ECNU-AM-0019 and ECUN-AM-0010, Figs. 1–3). Smallsized (8.07 mm.). Predominantly yellowish coloration with grayish and black spots. Cephalic capsule with dorsal surface (from vertex to fastigium) and upper area of frons with dark brown to black stripes; area surrounding ocelli, mandibles, lower margin of genae, and frons ocher; ocelli yellow; eyes greyish. Frons with a yellow inverted “Y”, * for the data may be doubtful due to different measurement method. starting from vertex, dividing at fastigium, and branching to the inner margin of each lateral ocellus. Pronotal disc greyish brown, with black corner rims and delimiting pronotal disc from lateral lobes; pronotal disk with two distinct large dark “eyespots”. Lateral lobes of pronotum with upper half grayish brown and lower half yellowish. All legs are yellowish with black or grayish spots, irregularly scattered, most noticeable on femora. Tegminae translucent, with veins outlined in light yellow. Abdomen yellowish covered by various black or grayish spots. Body covered with short or mid-sized fine hairs. Head as wide as pronotum, rounded and smooth; almost as wide as high in frontal view, with epistomal suture little concave. Vertex rounded. Eyes ovoid, not protruding. Lateral ocelli circular, central ocellus absent. Eyes and antennal pits located very low on face, close to epistomal suture, almost at the same level. Fastigium wide, almost four times as wide as scape. Maxillary palpi with the 4 th and 5 th longer, apical segment of labial palpi flattened. Thorax. Pronotum longer than wide, covered by several and mid-sized hairs; anterior margin concave and broader than posterior margin, with long bristles; lateral lobules rectangular, wider than high and with a rounded lateral edge, ventral margin almost straight. Wings. Tegmina short and ovoid, reaching to the second abdominal tergite. Mirror in oval shape, longer than wide, with a curved dividing vein; harp with three dividing veins; apical field reduced; lateral field two longitudinal veins from base to apex, and ten transversal veins born from the second longitudinal vein to costal margin. Hind wings no visible. Legs. Coxa and trochanter not elongated. Foreleg robust with long setae, large and oval tympana in both sides, on middle of fore tibiae; three movable dactyls on apical part of fore tibiae, first tarsomere almost as long as fore tibiae, other two tarsomere short. Mid leg slender, coxa robust, with a process partly covering trochanter; femur slightly slenderer than fore femur; tibia slightly inflated, shorter than femur, armed at apex with three inner spurs and two outer spurs. Hind leg elongated, femur strong, tibia with several small spines on dorsal side at apex. Three internal apical spurs long, and two external apical spurs short. The first and third tarsomere long and the second one short. Abdomen cylindrical; tergites with few short hairs; epiproctus subtriangular, apex rounded; subgenital plate short, wider than long, apex rounded; cerci almost as long as the two-thirds of hind tibia, covered by abundant medium-sized hairs. Female (emended description; based on specimens ECNU-AM-0002 and ECUN-AM-0024, Figs. 4, 5). Similar to the males in shape and size, differing by the following characters: the color tones are darker than the males, but keep the same pattern. Tegmina absent. Abdomen with hairs and several long bristles on dorsal side. Epiproct triangular, with rounded posterior margin. Ovipositor almost straight, apex of valves sharp, pointed, and lanceolate in dorsal view. Subgenital plate trapezoidal, apex moderately protruding. Male nymph (based on specimen ECNU-AM-0045, Fig. 6). Similar to the adult males, but no developed wings, with a more fragile and slender appearance. It differs from the adults, because the tympana are smaller in diameter than the other adult specimens examined. The small tympana in immature specimens can happen. This structure increases its size in adults. Measurements (mm). Adult male ECNU-AM-0019 (Fig. 1, 2): BL: 8.07; HL: 1.07; PL: 1.57; PW: 1.43; FWL: 2.18; FFL: 1.61; FTL: 1.04; MFL: 1.41; MTL: 1.09; HFL: 3.15; HTL: 1.82; AL: 5.14; CL: 2.26. Adult male ECNU-AM-0010 (Fig. 3): PL: 1.47; PW: 1.33; FWL: 2.05; FFL: 0.98; FTL: 1.06; Body broken, uncertain for else. Adult female ECNU-AM-0002 (Fig. 4): BL: 8.20; HL: 1.21; PL: 1.69 PW: 1.71 FFL: 1.56; FTL: 0.97; MFL: 1.91; MTL: not visible; HFL: 3.63; HTL: 2.54; AL: 4.63, OL: 1.82. Adult female ECNU-AM-0024 (Fig. 5): BL: 5.24; HL: 1.06; PL: 1.60; PW: uncertain; FFL: 1.45; FTL: 0.95; MTL: 1.42, femur and tarsus incomplete; HFL: 3.34, HTL: 2.56; AL: 3.82, CL: 2.39, OL: 1.72. Nymph ECNU-AM-0045 (Fig. 6): BL: 5.22; HL: 0.87; PL: 1.33, PW: uncertain; FFL: 1.40; FTL: 0.83; MFL: 0.97, MTL: 0.82; HFL: 2.49; HTL: 1.71; AL: 3.25; CL: 1.67. Remarks. Synonymy and new combination at the species level are justified as follows: 1) † P. micromorphus Poinar et al. 2020 is a subadult female specimen, which looks different in the shape of the pronotum in contrast to the holotype of † P. rectanguli n. comb. (a subadult male). However, when the photos of specimens are taken from different angles, an optical effect will be caused, which confuses us to consider them as different shapes. This can be happened in the additional specimens of † P. rectanguli n. comb. studied here. The fore tibia does not vary in both specimens. Only in the holotype of P. rectangulari n. comb., a bending of the ventral edge is observed, but this can happen due to dehydration during fossilization. However, we prefer to keep this species as valid, since the subadult female holotype specimen of † P. micromorphus had a wing pad, which is different from the adult females of † P. rectangulari n. comb. 2) † C. fania n. syn. is an adult male specimen. In the images of the original description, its diagnostic characters coincide with † P. rectanguli n. comb. in terms of the shape of fore tibia, apical spurs organization and shape, and location of the spurs of hind tibia. The hind tibiae are thin and moderately elongated, and the shape of pronotum and its spots coincide in these specimens. Unfortunately, the venation of † C. fania n. syn. is not observed due to preservation.Although the subgenital plate of † C. fania n. syn. dehydrated in fossilization process and coiled dorsally, it still could be observed in a triangular shape, which is similar to those of field crickets in a semicircle shape. 3) † Tresdigitus gracilis n. syn. is a nymph of indefinite sex (possibly female in penultimate or ultimate instar), not as developed as the two “species” previously synonymized. In the key presented in the original description by Jiang et al., 2022, the arrangement of head and the shape of pronotum are some of the characters proposed by the authors to distinguish the species from † P. rectanguli n. comb. The disposition of the head can vary in the way that insects die, and thus remains when fossilized. In contrast to † P. rectanguli n. comb., its head is in a normal position, locating mouthparts down (hypognathous/ hypognathous mandible). Other characters that stand out in † T. gracilis n. syn. is the moderately expanded fore tibia, and the small tympanum. However, it is common for crickets that they are different in these characters in their early stages. Through molting and growth, the tympanum becomes larger, and the tibiae attains the shape final in the last stages of insect development (Table 2; Ball and Cowan, 1978). In this study, we provide well-preserved males, females, and nymphal specimens, in which most of the morphological characters can be studied. It is applicable to verify the similarities and differences with the other recently described fossil taxa of “Mole Crickets” with those specimens. In terms of coloration, it was feasible to show that the coloration of † P. micromorphus is similar to those of the two males studied here. † P. rectanguli n. comb. has the same color pattern, but its image was darker in the original description (possibly due to a lack of brightness or light from the photographic record). For † C. fania n. syn., the same issue happened as mentioned above. All the specimens in the different stages of development have similar coloration, with two distinct large dark “eyespots” of similar shape; lower half of the anterior lobe of the yellow pronotum; shape and location of apical spurs of fore tibia; hind tibia serrulate with small spines; apex with conspicuous slender spines. The number of dorsal spines on legs is not reliable classification characters due to its intraspecific variation, which may be different from one leg to the other leg on one specimen (Cadena-Castañeda et al. 2022b). We consider † Pherodactylus micromorphus and † P. rectanguli n. comb. as valid genus and species. Both genera were described in 2020, and the original description of † P. micromorphus is available on March of 2020, while † Tresdigitus rectanguli is published on August of 2020. It is worth noting that Poinar et al. (2020) proposed a more reasonable taxonomic placement of its genus and species than the other authors, consisting with our results discussed and analyzed here. † Burmagryllotalpa longa is proposed to move under Gryllidae: Gryllinae: Sclerogryllini: Pherodactylina n. subtr. Although this species shares some morphologies with † Pherodactylus species, † B. longa completely lacks tympanum, and its pronotum is dorsally curved without spots on pronotal disc that characterize † Pherodactylus species. Thus, we consider † B. longa as valid.

† Pherodactylus Poinar, Su & Brown, 2020 = † Tresdigitus Xu, Fang & Wang, 2020 n. syn. = † Chunxiania Xu, Wang & Fang, 2022 n. syn. Diagnosis. Head round, pronotum elongate, longer than the wide, middle of pronotal disk with two distinct large dark “eyespots”, foreleg robust, with three apical spurs arranged on the inner side of fore tibia. Included species. † Pherodactylus micromorphus Poinar, Su & Brown, 2020 (type species by monotypy and original designation) and † Pherodactylus rectanguli (Xu, Fang & Wang, 2020) n. comb. Remarks. We treated genera † Tresdigitus n. syn. and Chunxiania n. syn. as synonyms of genus Pherodactylus Poinar, Su & Brown, 2020. We propose the genera and their species as synonyms or relocated: genera † Tresdigitus n. syn. and Chunxiania n. syn. as synonyms of genus Pherodactylus Poinar, Su & Brown, 2020, and † Tresdigitus gracilis Jiang, Xu, Jarzembowski & Xiao, 2022 n. syn. and † Chunxiania fania n. syn. as synonyms of † Pherodactylus rectanguli n. comb. The type specimen of each species represents different stages of the same species (Table 2), and they are all from the same locality. Besides, their morphological characters coincide with the diagnostic characters of † Pherodactylus. For additional taxonomic justifications, see † P. rectanguli n. comb. remarks., Published as part of Cadena-Castañeda, Oscar J., Liu, Yi-Jiao, Yu, Zhe-Yuan, Hu, Tian-Hao, Wu, Shi-Yang & He, Zhu-Qing, 2023, Are the recently described fossil Mole Crickets of Myanmar amber real gryllotalpids? (Orthoptera: Gryllotalpidae & Gryllidae), pp. 48-64 in Zootaxa 5311 (1) on page 52, DOI: 10.11646/zootaxa.5311.1.2, http://zenodo.org/record/8090321, {"references":["Poinar Jr., G., Su, Y. N. & Brown, A. E. (2020) A new genus of crickets (Orthoptera: Gryllidae) in Mid-Cretaceous Myanmar Amber. Biosis: Biological Systems, 1 (1), 33 - 38. https: // doi. org / 10.37819 / biosis. 001.01.0049","Wang, H., Lei, X., Zhang, G., Xu, C., Fang, Y. & Zhang, H. (2020) The earliest Gryllotalpinae (Insecta, Orthoptera, Gryllotalpidae) from mid-Cretaceous Burmese amber. Cretaceous Research, 107, 104292. https: // doi. org / 10.1016 / j. cretres. 2019.104292","Xu, C., Wang, H., Fang, Y., Jarzembowski, E. A. & Zhuo, D. (2022) Chunxiania fania: a new genus and species of mole cricket (Orthoptera: Ensifera: Gryllotalpidae) from mid-Cretaceous Kachin amber. Cretaceous Research, 134, 105 - 159. https: // doi. org / 10.1016 / j. cretres. 2022.105159","Jiang, X., Xu, C., Jarzembowski, E. A. & Xiao, C. (2022) A peculiar species of mole cricket (Orthoptera: Gryllotalpidae) from mid-Cretaceous Kachin amber. Cretaceous Research, 139, 1 - 6. https: // doi. org / 10.1016 / j. cretres. 2022.105273"]}

Cadena-Castañeda, Oscar J., Liu, Yi-Jiao, Yu, Zhe-Yuan, Hu, Tian-Hao, Wu, Shi-Yang, He, Zhu-Qing (2023): Are the recently described fossil Mole Crickets of Myanmar amber real gryllotalpids? (Orthoptera: Gryllotalpidae & Gryllidae). Zootaxa 5311 (1): 48-64, DOI: 10.11646/zootaxa.5311.1.2, URL: http://dx.doi.org/10.11646/zootaxa.5311.1.2

Tribe Sclerogryllini Gorochov, 1985 Remarks. This tribe was described by Saussure (1877) as “Légion des Scléropterites”, including Acanthoplistus Saussure, 1877, and Scleropterus Haan, 1844. Subsequently, this taxon was treated as a subfamily by Shiraki (1930), and this status was followed by Chopard (1934, 1936) and Desutter (1987). Gorochov (1985) proposed a new name Sclerogryllus for Scleropterus Haan, 1844, which has been used for Scleropterus Schoenherr, 1825 (Coleoptera). At the same time, Gorochov also established the tribal status as Sclerogryllini, with Sclerogryllus as the type genus under Gryllinae. Otte (1994) argued that according to the nomenclatural change proposed by Gorochov, the subfamily name should be changed to Sclerogryllinae. Otte used the name Sclerogryllinae for the first time, and this was followed by Storozhenko et al. (2015). Recently, Yu et al. (2022) published a complete mitochondrial genome of cricket Sclerogryllus punctatus. By phylogenetic analysis, they verified that Sclerogryllus was grouped within the Gryllinae clade. Therefore, Sclerogryllinae should be treated as a synonym of Gryllinae, and Gorochov’s tribe Sclerogryllini under Gryllinae is recommended rather than a different subfamily from field crickets. Sclerogryllini is a monogeneric tribe, with five species distributed in Asia. Rhabdotogryllus caraboides Chopard, 1954 from Guinea (Africa), is only known from its holotype female, and no additional records have been reported since its description. R. caraboides is superficially similar to Sclerogryllus females, but possessing more conspicuous dorsal spurs on the hind tibia. Thus, the homology of Rhabdotogryllus and Sclerogryllus is still a question, and Sclerogryllini probably only occurs in Asia. Here, we propose a new fossil subtribe that more closely resembles extant Sclerogryllini species in morphology than any other tribal taxon of field crickets. Additionally, a key to separate the taxa of the tribe is provided., Published as part of Cadena-Castañeda, Oscar J., Liu, Yi-Jiao, Yu, Zhe-Yuan, Hu, Tian-Hao, Wu, Shi-Yang & He, Zhu-Qing, 2023, Are the recently described fossil Mole Crickets of Myanmar amber real gryllotalpids? (Orthoptera: Gryllotalpidae & Gryllidae), pp. 48-64 in Zootaxa 5311 (1) on page 51, DOI: 10.11646/zootaxa.5311.1.2, http://zenodo.org/record/8090321, {"references":["Gorochov, A. V. (1985) On the Orthoptera subfamily of Gryllinae (Orthoptera, Gryllidae) from eastern Indochina. In: Medvedev, L. N. (Ed.), Fauna iekologiya nasekomykh Vetnama [The fauna and ecology of insects of Vietnam], 1985, pp. 9 - 17.","Saussure, H. (1877) Melanges orthopterologiques V. fascicule Gryllides. Memoires de la Societe de Physique et d'Histoire Naturelle de Geneve, 25 (1), 169 - 504.","Shiraki, T. (1930) Orthoptera of the Japanese Empire. Part I. (Gryllotalpidae and Gryllidae). Insecta Matsumurana, 4, 181 - 252.","Chopard, L. (1934) Catalogues raisonnes de la faune entomologique du Congo belge. Orthopteres - Gryllides. Annales du Musee du Congo belge Tervueren, Belgique, Zoologie, 3 (2), 4, 1 - 88.","Chopard, L. (1936) The Tridactylidae and Gryllidae of Ceylon. Ceylon Journal of Science (Biological Science), 20, 9 - 87.","Desutter, L. (1987) Structure et evolution du complexe phallique de Gryllidea (Orthopteres) et classification des genres neotropicauxde Grylloidea. Premiere partie. Annales de la Societe Entomologique de France, New Series, 23 (3), 213 - 239.","Otte, D. (1994) Crickets (Grylloidea). Orthoptera Species File. Vol. 1. Orthopterist' Society, Philadelphia, Pennsylvania, iv + 120 pp.","Storozhenko, S. Y., Kim, T. W. & Jeon, M. J. (2015) Monograph of Korean Orthoptera. National Institute of Biological Resources, Incheon, 377 pp.","Yu, Z., Xie, H., Liu, Y., Li, K. & He, Z. (2022) The complete mitochondrial genome of cricket Sclerogryllus punctatus (Orthoptera: Gryllidae) and phylogenetic analysis. Journal of Asia-Pacific Entomology, 25 (3), 1 - 9. https: // doi. org / 10.1016 / j. aspen. 2022.101933"]}