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Sphenarium planum Bruner, 1906 (http://lsid.speciesfile.org/urn:lsid: Orthoptera.speciesfile.org:TaxonName:36998) Description. External morphology (Figs. 8 C, D; 9K, L): total body length ranging from 21.30 to 29.04 mm in females and 18.29 to 26.24 mm in males; antennae filiform, notably shorter in females or slightly longer than head and pronotum together in males; head subtriangular-compressed, wider than long with spherical eyes in females or subtriangular-elongated moderately longer than wide with oval eyes in males; fastigium notably reduced, less than a half the length of interocular space in females or moderately elongated, nearly half the length of interocular space in males; tegmina spatula-like in both sexes; subgenital plate of males rounded, moderately developed posteriorly; dorsal ovipositor valves lanceolate, moderately elongated towards the apex. Male genitalia: bridge of epiphallus as long as the length of lateral plates in most cases (Fig. 12 D-I). Ectophallus in dorsal view (Fig. 12 D-II) with lateral borders of ramus convergent slightly rounded; basal emargination of cingulum notably reduced; interspace between apodemal plates of cingulum widely open. Ectophallus in posterior view (Fig. 12 E) without a conspicuous sclerotized hollow in the sheath; inflections of supraramus reduced; ramus of cingulum distinctly developed ventrally; valves of cingulum with unique form notably developed posteriorly (Fig. 12 F). In lateral view of endophallus (12D-III) pseudoarch elongated loosely joined to the valves of cingulum; aedeagal valves slender, long, with smooth ventral margins and moderately rounded in the apex without apical spine; aedeagal valves and sclerites together about 2 �� fold the length of endophallic apodemes. Colouration. Ground colours vary from green, beige, brown or grey, yellow or brown. Body uniformly coloured (Fig. 9 L) or with the following colour traits: antennae often yellow; fastigium frequently brownish to reddish; lateral postocular bands yellow; dorsomedial line frequently absent, if present very narrow, restricted almost entirely to the abdomen and yellowish; dorsal shades black, grey or brown, frequently less evident in head and pronotum; lateral shades frequently absent, if present very narrow and restricted to the head; lateral bands of blotches sometimes evident and yellowish; ventral bands of pronotum if present yellowish, wide or narrow; mesonotum partially or entirely black; lateral blotches of 1st abdominal segment whitish; hind femora frequently with upper medial area dark green and lower medial area yellow; frequently knees of hind femora laterally black, dorsally brownish; frequently hind tibia yellow or brown. Diagnosis. Externally this species closely resembles S. purpurascens and S. rugosum. Generally, females of S. planum have smaller antennae than any other of its congeners. Moreover, when present, the yellow colourations of the lateral bands of blotches and the medial area of the hind femora of S. planum distinguish it from S. purpurascens. On the other hand, the male genitalia of S. planum more closely resemble that of S. rugosum, and S. crypticum sp.n., which also lack the apical spine of aedeagus. However, S. planum differs from all these species by the following combination of male genital traits: lateral borders of ramus convergent slightly rounded, basal emargination of cingulum notably reduced with a wide open interspace between the apodemal plates of cingulum, inflections of supraramus reduced, ramus of cingulum is distinctly developed ventrally, aedeagal valves moderately rounded in the apex without apical spine, and aedeagal valves and sclerites relatively long and slender. Distribution. This species is apparently restricted to the Tehuacan valley with sparse records in the surrounding mountain ranges of this valley in Puebla, Oaxaca and Veracruz, Mexico (Fig. 7 A) in elevations ranging from approximately 1200 to 2095 m a.s.l. Material examined. Lectotype m (Fig. 8 C) and paralectotype f (Fig. 8 D) from Mexico: Puebla, Tehuacan, XI (L. Bruner); designation: Rehn and Hebard (1912); location: ANSP. We could examine only the external morphology of this type material. Additional material: 10 m, 10 f, from the type locality; 45 m, 16 f, from 12 adjacent localities (Appendix Table 5). Taxonomic discussion. Bruner (1906) originally described this species from two syntypes (a male and female) erroneously specifying Tehuantepec as its type locality rather than Tehuacan as it is labelled in the type material. Bruner also recognised a close relationship between S. planum and S. purpurascens indicating that the former species differed from the latter in its more rotund form and in the absence of lateral carina of pronotum. However, these traits are ambiguous and variable within the genus. Posteriorly, Boyle (1974) identified differences in endophallic morphology between these two species, but he interpreted this differentiation as geographic variation of S. p. purpurascens and/or probable hybridization with S. p. minimum, recognising S. planum only as an intermediate form between these two later taxa. Previously, we proposed the re-establishment of S. planum as valid species mainly based on genetic evidence (Sanabria-Urb��n et al. 2015). In this study we found that S. planum shows a unique combination of male genitalia characters, a well-supported monophyly (Fig. 2), relatively high levels of interspecific genetic differentiation (Table 3). All these lines of evidence support the recognition of S. planum as a valid species., Published as part of Sanabria-Urb��n, Salom��n, Song, Hojun, Oyama, Ken, Gonz��lez-Rodr��guez, Antonio & Castillo, Ra��l Cueva Del, 2017, Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae), pp. 1-86 in Zootaxa 4274 (1) on pages 32-33, DOI: 10.5281/zenodo.804182, {"references":["Bruner, L. (1906) Insecta. Orthoptera. Vol. II. In: Godman, F. D. & Salvin, O. (Eds.), Biologia Centrali-Americana. R. H. Porter, London, pp. 199 - 207.","Rehn, J. A. G. & Hebard, M. (1912) Fixation of the single type (Lectotype) specimens of species of American Orthoptera. Proceedings of the Academy of Natural Sciences of Philadelphia, 64 (1), 60 - 183.","Boyle, W. K. (1974) A Revision of the Genus Sphenarium (Orthoptera: Pyrgomorphidae). McGill University, McGill, 214 pp.","Sanabria-Urban, S., Song, H., Oyama, K., Gonzalez-Rodriguez, A., Serrano-Meneses, M. A. & Cueva del Castillo, R. (2015) Body size adaptations to altitudinal climatic variation in neotropical grasshoppers of the genus Sphenarium (Orthoptera: Pyrgomorphidae). Plos ONE, 10 (12), 1 - 24."]}

Sphenarium purpurascens Charpentier, 1842 (http://lsid.speciesfile.org/urn:lsid: Orthoptera.speciesfile.org:TaxonName:36986) Description. External morphology (Figs. 8 A, B; 9A, B, C, D, E, F): total body length ranging from 18.01 to 30.24 mm in females and 16.04 to 28.14 mm in males. In most cases: antennae filiform, slightly shorter in females or longer than head and pronotum together in males; head subtriangular-compressed, wider than long with spherical eyes in females or subtriangular-elongated moderately longer than wide with oval eyes in males; fastigium notably reduced, less than a half the length of interocular space in females or moderately elongated, nearly half the length of interocular space in males; tegmina spatula-like; subgenital plate of males rounded, moderately developed posteriorly; dorsal ovipositor valves lanceolate, moderately elongated towards the apex. Male genitalia: bridge of epiphallus as long as the length of lateral plates in most cases (Figs. 5 A; 10A-I, D-I, G-I). Ectophallus in dorsal view (Fig 10 A-II, D-II, G-II) with lateral borders of ramus strongly concave; basal emargination of cingulum moderately developed; interspace between apodemal plates of cingulum moderately open. Ectophallus in posterior view (Fig. 10 B, E, H) without a conspicuous sclerotized hollow in the sheath; inflections of supraramus moderately developed with distal margins dorsally directed in most cases; valves of cingulum small, triangular, slightly developed (morphotypes 1 and 2; Figs. 10 C, F, respectively) or notably developed posteriorly (morphotype 3; Fig. 10 I). In lateral view of endophallus (Fig. 10 A-III, D-III, G-III) pseudoarch elongated, loosely joined to the valves of cingulum; aedeagal valves long with smooth ventral borders and an apical spine slightly longer (morphotypes 1 and 3; Fig. 10 A-III, G-III, respectively) or shorter (morphotype 2; Fig. 10 D-III) than the base of aedeagal sclerites (see Fig. 5 E); aedeagal valves and sclerites together about twice (morphotypes 1 and 3; Fig. 10 A-III, G-III, respectively) or 1�� fold (morphotype 2; Fig. 10 D-III) the length of dorsal inflections of endophallic apodemes (see Fig. 5 F). Colouration. Ground colours vary from green, beige, brown or grey (Fig. 9 A, B, C, D, E, F). Body uniformly coloured with ground colours (Fig. 9 F) or with the following colour traits: antennae black, yellow or orange; fastigium reddish or brownish; lateral postocular bands whitish or yellowish; dorsomedial line frequently present, narrow, whitish or reddish; dorsal shades frequently present, black, purple, brown or grey, covering partially (Fig. 9 E) or entirely (Fig. 9 A, C) the dorsal portion of the body; lateral shades often present; lateral bands of blotches not evident; ventral bands of pronotum generally present, wide and whitish; mesonotum partially or entirely black; light lateral blotches of 1st abdominal segment generally present and whitish; hind femora frequently with upper medial area black to brown and lower medial area whitish or yellowish; frequently knees of hind femora black laterally, brownish to reddish dorsally; hind tibia black, yellow or orange. Diagnosis. Sphenarium purpurascens mainly differs from its congeners in the following combination of male genitalia characters: lateral borders of ramus of cingulum strongly concave, inflections of supraramus moderately developed, aedeagus valves moderately long and its apical spine of aedeagus always present slightly longer or notably shorter than the base of aedeagal sclerites. Distribution. This species is distributed in elevations ranging approximately from 800 to 2700 m a.s.l. from the southern Altiplano to the Sierra Madre del Sur in the Mexican states of Guanajuato, Hidalgo, Mexico, Mexico City, Michoacan, Oaxaca, Puebla, Queretaro, Tlaxcala, and Veracruz (Fig. 7 A). The distribution of S. purpurascens is interrupted at the Tehuacan valley and somewhat delimited by the higher mountains of the Mexican Volcanic Belt, Sierra Madre Oriental and Sierra Madre del Sur. Across this geographic range the morphotype 1 of this species has the widest distribution, whereas the morphotype 2 is restricted to the central valleys of Oaxaca and the morphotype 3 is found in separate populations in the inner slopes of the Sierra Madre del Sur, Oaxaca (Fig. 7 A & C). Previously, lower and southern ranges were recognized for S. purpurascens (Kevan, 1977). However, these assumptions were based on two misidentified male nymphs from 1.0 mi W Puerto Angel, Oaxaca (resembling S. histrio nymphs); and three adult females from Las Margaritas, Chiapas, apparently corresponding to S. purpurascens but probably mislabelled. Moreover, during our fieldwork we did not find S. purpurascens beyond the above-specified ranges. Material examined. Lectotype m (Fig. 8 A) and paralectotype f (Fig. 8 B) from Mexico (V. Charpentier); designation: Kevan (1960, unpublished results); location: Berlin Zoological Museum (BZM), Berlin, Germany. We were able to examine only the external morphology of this material. Additional material: 459 m, 361 f, from 100 localities. Locality information and depositories of these examined specimens is provided in Appendix Table 5. Taxonomic discussion. Charpentier (1842) described this species apparently based on two females and males syntypes from an unspecified Mexican locality. After its original description the taxonomic status of S. purpurascens remained unchanged until Boyle (1974) and Kevan (1977) recognised it as a subspecies, S. purpurascens purpurascens. Recently, Pedraza-Lara et al. (2015) and Sanabria-Urb��n et al. (2015) proposed the elevation of this species at the species level considering principally its morphological distinctiveness. Based on the morphological, genetic and geographical cohesiveness of this species we also consider S. purpurascens as a valid species within the genus., Published as part of Sanabria-Urb��n, Salom��n, Song, Hojun, Oyama, Ken, Gonz��lez-Rodr��guez, Antonio & Castillo, Ra��l Cueva Del, 2017, Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae), pp. 1-86 in Zootaxa 4274 (1) on pages 24-25, DOI: 10.5281/zenodo.804182, {"references":["Kevan, D. K. M. (1977) The American Pyrgomorphidae (Orthoptera). Revista de la Sociedad Entomologica Argentina, 36 (1 - 4), 3 - 28.","Boyle, W. K. (1974) A Revision of the Genus Sphenarium (Orthoptera: Pyrgomorphidae). McGill University, McGill, 214 pp.","Pedraza-Lara, C., Barrientos-Lozano, L., Rocha-Sanchez, A. Y. & Zaldivar-Riveron, A. (2015) Montane and coastal species diversification in the economically important Mexican grasshopper genus Sphenarium (Orthoptera: Pyrgomorphidae). Molecular Phylogenetics and Evolution, 84, 220 - 231.","Sanabria-Urban, S., Song, H., Oyama, K., Gonzalez-Rodriguez, A., Serrano-Meneses, M. A. & Cueva del Castillo, R. (2015) Body size adaptations to altitudinal climatic variation in neotropical grasshoppers of the genus Sphenarium (Orthoptera: Pyrgomorphidae). Plos ONE, 10 (12), 1 - 24."]}

Sphenarium rugosum Bruner, 1906 (http://lsid.speciesfile.org/urn:lsid: Orthoptera.speciesfile.org:TaxonName:37030) Sphenarium barretti Bruner, 1906. Description. External morphology (Figs. 14 A, B, C; 9S, T; 15A, B, C, D): total body length ranging from 25.46 to 36.14 mm in females and from 19.8 to 35.07 mm in males; antennae filiform, slightly shorter in females or longer than head and pronotum together in males; head subtriangular-elongated nearly as wide as long in females or moderately longer than wide in males with oval eyes in both sexes; fastigium moderately elongated, nearly half the length of interocular space in both sexes; tegmina spatula-like in both sexes; subgenital plate of males rounded moderately developed posteriorly; dorsal ovipositor valves lanceolate moderately elongated towards the apex. Male genitalia: bridge of epiphallus moderately longer than the length of lateral plates (Fig. 13 D-I, G-I, J-I). Ectophallus in dorsal view (Fig. 13 D-II, G-II, J-II) with lateral borders of ramus convergent; basal emargination of cingulum moderately or not developed (morphotypes 1 and 3; Fig. 13 D-II, J-II, respectively) or notably developed (morphotype 2; Fig. 13 G-II); interspace between apodemal plates of cingulum moderately open (morphotypes 1 and 3; Fig. 13 D-II, J-II, respectively) or notably close (morphotype 2; Fig. 13 G-II). Ectophallus in posterior view (Fig. 13 E, H, K) without a conspicuous sclerotized hollow in the sheath; inflections of supraramus moderately developed with distal margins laterally or dorsally directed; valves of cingulum triangular to slightly quadrangular, relatively small (morphotypes 1 and 2; Fig. 13 E, H, respectively) or large (morphotype 3; Fig. 13 K), slightly developed posteriorly (Fig. 13 F, I, L). Endophallus in lateral view (Fig. 13 D-III, G-III, J-III) with elongated pseudoarch, loosely joined to the valves of cingulum; aedeagal valves moderately (morphotype 1; Fig. 13 D-III) or notably (morphotype 2 and 3; Fig. 13 G-III, J-III, respectively) broad, medium-sized (morphotype 1 and 3) or short (morphotype 2), with ventral margins smooth (morphotypes 1 and 2) or somewhat serrated (morphotype 3; Fig. 13 J-IV, arrow), broadly rounded apically (Fig. 13 E, I, K, arows); aedeagal valves and sclerites together varying from �� (morphotype 2; Fig. 13 G-III) to 1 �� (morphotypes 1 and 3; Fig. 13 D-III, J-III) fold the length of dorsal inflections of endophallic apodemes. Colouration. Ground colours vary from olive green to brown. Body uniformly coloured (Fig. 9 T; 15B, D) or with the following colour traits: antennae frequently black or dark brown; fastigium generally reddish; lateral postocular bands frequently present, wide and yellow; dorsomedial line frequently present, narrow and yellowish; dorsal shades often present, covering partially (Fig. 9 S) or entirely (Fig. 15 C) the dorsal part of the body, black to brown; lateral shades often present, black or dark brown, mostly evident in the abdomen (Fig. 9 S); lateral bands of blotches often present with distinct yellow colouration; ventral bands of pronotum often present and yellow; sometimes lateral carinas of pronotum highlighted with yellow (Fig. 9 S); mesonotum partially or entirely black; lateral blotches of 1st abdominal segment frequently present, whitish or yellowish; hind femora uniformly coloured or with upper medial area black and lower medial area yellowish: knees of hind femora laterally black, dorsally reddish; hind tibia frequently black. Diagnosis. Externally this species closely resembles S. macrophallicum, S. planum, S. tarascum sp.n., S. crypticum sp.n., and S. infernalis sp.n. Sometimes S. rugosum differs from all these species by its more conspicuous yellow colourations in the dorsomedial line, lateral carinas and lateral bands of blotches, in combination with darker dorsal colourations in the body (Fig. 15 C). Nevertheless, S. rugosum clearly differs from all these species by the following combination of male genitalia characters: lateral borders of ramus convergent, inflections of supraramus moderately to notably developed, and aedeagal valves moderately or notably broad, medium sized or small, broadly rounded apically. Distribution. This species is distributed in elevations ranging from 457 to 2344 m a.s.l. and is apparently confined to the eastern Balsas River Basin and the Sierra Madre del Sur in the Mexican states of Guerrero, Mexico, Michoacan, Morelos, Oaxaca, and Puebla (Fig. 7 A). The morphotype 1 of this species is widely distributed in the western portion of the Balsas River Basin; whereas the morphotype 2 is apparently restricted to inner slope of middle portion of the Sierra Madre del Sur (Fig. 7 A). The morphotype 3 is only known from three localities east of the middle portion of the Balsas River Basin (Fig. 7 A). Material examined. S. rugosum: lectotype m (Fig. 14 A) and paralectotype f (Fig. 14 B) from Mexico: Morelos, Cuernavaca, I-4-1899 (C. C. Deam); designation: Rehn and Hebard (1912); location: ANSP. S. barretti: lectotype m (Fig. 14 C), from Mexico: Guerrero, Rio Cocula, XII (O. W. Barrett); designation: Rehn and Hebard (1912); location: ANSP. We examined only external morphology of this type material. Additional material: 355 m, 201 f, from 80 localities (see Appendix Table 5). Taxonomic discussion. Bruner (1906) originally described this species based on one male and female syntypes from Cuernavaca, Morelos. He considered the dull granulose and pubescent surface of the type specimen of S. rugosum as the main evidence for separating this species from its congeners, though these traits are variable across the species and the genus. Bolivar (1909) and Hebard (1932) recognized S. rugosum as a valid species. M��rquez (1962) synonymized this species within S. purpurascens based on variable and ambiguous morphologic traits of the epiphallus and endophallus of both species. Later, Boyle (1974) and Kevan (1977) re-established S. rugosum as valid species based on more detailed analysis of its male genital morphology. Recent genetic studies (Pedraza-lara et al. 2015; Sanabria-Urb��n et al. 2015) have also supported the validity of S. rugosum. In this study we analysed several specimens of S. rugosum, some of them collected at or near the type locality of this species (L414, L435, L440, and L449; Appendix Table 5). Our phylogenetic analysis recovered S. rugosum as a paraphyletic species and we observed relatively low levels of genetic differentiation between S. rugosum and other morphologically different species including adjacent (S. crypticum sp.n., S. macrophallicum, S. purpurascens, and S. tarascum sp.n.) and distant taxa (S. minimum, S. variabile, and S. zapotecum sp.n.) (see Table 3). Nevertheless, the unique combination of morphologic traits, and restricted geographic distribution of this species supports its recognition as an independent species within the genus. Moreover, we observed considerable morphological similarity (externaly and in male genitalia), low genetic differentiation (CO1 P-distances S. rugosum and the putative new taxa Sphenarium sp. Gro1+Gro8 and Sphenarium sp.n. 6 recognized in Pedraza-Lara et al. (2015) and Sanabria-Urb��n et al. (2015) respectively. Therefore, we consider that these taxa represent part of the same species, S. rugosum (morphotype 2 and 3, respectively). Sphenarium barretti was originally described by Bruner (1906) based on an unstated number of specimens, from Rio Cocula, Guerrero. Bruner (1906) originally considered a closer relationship between S. barretti and S. purpurascens. Posteriorly, S. barretti was initially synonymised as S. purpurascens (M��rquez 1962) and later as S. rugosum (Boyle 1974; Kevan 1977). In this revision we examined several specimens collected near the type locality of this species (L428, L427 and L423; Appendix Table 5); which were similar in their male genitalia to other S. rugosum specimens (morphotype 1). Therefore, we agree in considering S. barretti as a synonym of S. rugosum., Published as part of Sanabria-Urb��n, Salom��n, Song, Hojun, Oyama, Ken, Gonz��lez-Rodr��guez, Antonio & Castillo, Ra��l Cueva Del, 2017, Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae), pp. 1-86 in Zootaxa 4274 (1) on pages 39-40, DOI: 10.5281/zenodo.804182, {"references":["Bruner, L. (1906) Insecta. Orthoptera. Vol. II. In: Godman, F. D. & Salvin, O. (Eds.), Biologia Centrali-Americana. R. H. Porter, London, pp. 199 - 207.","Rehn, J. A. G. & Hebard, M. (1912) Fixation of the single type (Lectotype) specimens of species of American Orthoptera. Proceedings of the Academy of Natural Sciences of Philadelphia, 64 (1), 60 - 183.","Bolivar, I. (1909) Orthoptera Fam. Acridiidae Subfam. Pyrgomorphinae. Genera insectorum, 90, 1 - 59.","Hebard, M. (1932) New species and records of Mexican Orthoptera. Transactions of the American Entomological Society, 58 (3), 201 - 371.","Marquez, C. (1962) Estudios de las especies del genero Sphenarium basado en sus genitalia (Acrididae; Orthoptera), con la descripcion de una nueva especie. Anales de Instituto de Biologia UNAM Serie Zoologia, 33, 247 - 258.","Boyle, W. K. (1974) A Revision of the Genus Sphenarium (Orthoptera: Pyrgomorphidae). McGill University, McGill, 214 pp.","Kevan, D. K. M. (1977) The American Pyrgomorphidae (Orthoptera). Revista de la Sociedad Entomologica Argentina, 36 (1 - 4), 3 - 28.","Sanabria-Urban, S., Song, H., Oyama, K., Gonzalez-Rodriguez, A., Serrano-Meneses, M. A. & Cueva del Castillo, R. (2015) Body size adaptations to altitudinal climatic variation in neotropical grasshoppers of the genus Sphenarium (Orthoptera: Pyrgomorphidae). Plos ONE, 10 (12), 1 - 24.","Pedraza-Lara, C., Barrientos-Lozano, L., Rocha-Sanchez, A. Y. & Zaldivar-Riveron, A. (2015) Montane and coastal species diversification in the economically important Mexican grasshopper genus Sphenarium (Orthoptera: Pyrgomorphidae). Molecular Phylogenetics and Evolution, 84, 220 - 231."]}

Sphenarium occidentalis Sanabria-Urb��n, Song & Cueva del Castillo sp.n. (http://lsid.speciesfile.org/urn:lsid: Orthoptera.speciesfile.org:TaxonName:495098) Description. External morphology (Fig. 20 K, L, M, N): total body length ranging from 32.58 to 43.67 mm in females and from 30.84 to 43.03 mm in males; antennae filiform, slightly shorter in females or longer than head and pronotum together in males; head conical notably longer than wide with oval eyes in both sexes; fastigium moderately elongated, nearly half the length of interocular space in females or notably elongated, nearly as long as the interocular space in males; tegmina strap-like in both sexes; subgenital plate of males somewhat tapered in the apex; dorsal ovipositor valves rounded or lanceolate slightly elongated towards the apex. Male genitalia: bridge of epiphallus slightly longer than the length of lateral plates (Fig. 21 J-I). Ectophallus in dorsal view (Fig. 21 J-II) small with lateral borders of ramus convergent, straight or slightly rounded; basal emargination of cingulum mostly slightly developed; interspace between the apodemal plates notably closed. Ectophallus in posterior view (Fig. 21K) with a conspicuous sclerotized hollow in the sheath moderately open; valves of cingulum arrow-like, relatively small; inflections of supraramus moderately developed whit distal borders laterally directed. Ectophallus in lateral view (Fig. 21 L) with valves of cingulum not developed posteriorly. Endophallus in lateral view (Fig. 21 J- III) with a short pseudoarch tightly joined to the valves of cingulum; aedeagal valves very small, tapered in the apex without apical spine; aedeagal valves and sclerites ranging from one half to �� the length of dorsal inflections of endophallic apodemes. Colouration. round colours vary from yellow to green. Body uniformly coloured or with the following colour traits: antennae black, gray or dark blue; fastigium blue or brown; lateral postocular bands frequently present, wide and yellowish; dorsomedial line frequently present, wide and yellowish; dorsal shades dark blue to dark green covering partially the dorsal portion of the body, frequently present in northern populations (Fig. 20 K, L) or absent in southern populations (Fig. 20 M, N); lateral shades often present, black or dark blue; lateral bands of blotches frequently present, yellow to pale orange; ventral bands of pronotum often present, wide, whitish to bluish; pronotum sometimes with white lateral carinas and small strips and dots in the posterior margin; mesonotum yellow or pale orange; frequently mesonotum and metanotum laterally white; lateral blotches of 1st abdominal segment frequently present and whitish; hind femora uniformly coloured with knees laterally black laterally, dorsally bluish or brownish. Diagnosis. This species is similar to S. histrio both externally and in male genital structures. Nevertheless, S. occidentalis sp.n. differs from S. histrio by the following combination of characters: if present lateral bands of blotches yellow to pale orange but never red, sclerotized hollow in the sheath moderately open, inflections of supraramus moderately developed whit distal borders laterally directed, and valves of cingulum although similar in form (specially to S. histrio morphotype 1) somewhat larger than in S. histrio. Distribution. This species is distributed in elevations ranging approximately from 30 to 750m a.s.l. and is apparently restricted to the western portion of the Balsas River Basin and the Pacific Cost in Michoacan and Guerrero, Mexico (Fig. 7 A). Material examined. Holotype m (Fig. 20 K) from Mexico: Michoacan, Cuchurumuco, 18.67008��N, - 101.670625��W, 241 m a.s.l., IX-22-2012 (Sanabria-Urb��n S. #P39); measurements: BS = 41.36 mm, FL = 1.66 mm, PL = 9.98 mm, HF = 19.25 mm. Paratypes from Mexico: Michoacan: 7 m, 7 f, same data as holotype; 1 m, 20 mi E Nueva Italia (on La Huacana rd.), IX-25-1959 (I. J. Cantrall & T. J. Cohn #174); 1 m, 23 mi E Nueva Italia (on La Huacana rd.), IX-25-1959 (I. J. Cantrall & T. J. Cohn #175); 1 m, 31mi S Nueva Italia (on Arteaga rd), 550 ft, IX-26-1959 (I. J. Cantrall & T. J. Cohn #182); 1 m, 11 mi SW La Huacana, 1600 ft, XII-4-1959 (T. J. Cohn #346); 1 m, 26 rd. mi NE Arteaga on Hwy. 37 (rd. mi SW Rancho Nuevo), 2060 ft, XI-3-1974 (T.J. & J. W. Cohn #121); 1 m, 3rd mi SW Arteaga church (0.1mi E Hwy. 37), 2460 ft, XI-3-1974 (T.J. & J. W. Cohn #122); 1 m, 7.2 mi NE Playa Azul (on Hwy. 37) 2.8 mi. NE La Mira Jct., 700 ft, XI-6-1974 (T.J. & J. W. Cohn #123); 1 m, 1 f, 15mi W Caleta de Campos (W playa Azul), X-9-1981 (Otte #62); 1 m, 1 f, 12mi W Caleta de Campos (W playa Azul), X-9- 1981 (Otte #63); 1 m, 1 f, 26mi n La Mira, VIII-9-1981 (Otte #59); 1 m, 1 f, 16-20km NE Rt 200 Ixtapa-Altamirano Rd., VIII-9-1981 (Otte #60); 6 M, 6 F, Las Pe��itas, 17.99 0621��N, - 102.026924��W, 36 m a.s.l., IX-23-2012 (Sanabria-Urb��n S. # P40 [L7 MS1]); 1 M, 1 f, Zicuir��n Carr. 120 Km 144, 18.881308��N, - 101.967336��W, 239 m a.s.l., IX-22-2012 (Sanabria-Urb��n S. # P38 [L8 MS1]); 1 m, 1 f, Carr. 37 D Km 236, 18.3868��N, - 101.89475��W, 197 m a.s.l., IX- 23-2012 (Sanabria-Urb��n S. # M055). Guerrero: 1 m, 5mi N Acapulco, IX-15-1940 (C. Bolivar & H. R. Roberts); 2 m, 1 f, 13mi SW Tierra Colourada, 1000 ft, XII-11-1958 (T. J. Cohn # 368); 6 m, 4 f, Rio Papagayo Carr. 200, 16.773689��N,- 99.608538��W, 56 m a.s.l., X-5-2011 (Sanabria-Urb��n S. # 16 ACA [L9 MS1]). The holotype was deposited at IBUNAM and paratypes were deposited at IBUNAM and TAMUIC. Additional material: 3 m, 3 f, same locality as holotype; 64 m, 59 m, from other 25 localities (Appendix Table 5). Taxonomic discussion. This species is closely related morphologically to S. histrio. In deed material of this new species was identified as S. m. histrio in previous studies (Boyle 1974; Kevan 1977). Nevertheless, S. occidentalis sp.n. shows a unique combination of morphologic traits, in both external and male genitalia structures, relatively high levels of interspecific genetic differentiation (Table 3) and is geographically separated from its congeners. All these lines of evidence support the recognition of S. occidentalis sp.n. as a valid species. Recently, Sanabria-Urb��n et al. (2015) identified specimens of this new species as Sphenarium sp.n. 1, whereas Pedraza-Lara et al. (2015) recognised two taxa of uncertain identity, Sphenarium sp. Gro6 and Sphenarium sp. Gro9, collected within the distribution ranges of S. occidentalis sp.n. During this revision we examined multiple specimens collected near to the localities of Sphenarium sp. Gro6 and Gro9 (e.g. L09, L309, and L307, L315; Appendix Table 5) that were undoubtedly S. occidentalis sp.n. Moreover, CO1 sequences of Sphenarium sp. Gro6 of S. occidentalis sp.n. formed a well supported (PP Ż 0.95) monophyletic group (Fig. 11). Therefore, we consider that Sphenarium sp. Gro6 and Sphenarium sp. Gro9 probably represent additional populations of S. occidentalis sp.n. Etymology. This species is named after its distribution in the occidental region of Mexico., Published as part of Sanabria-Urb��n, Salom��n, Song, Hojun, Oyama, Ken, Gonz��lez-Rodr��guez, Antonio & Castillo, Ra��l Cueva Del, 2017, Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae), pp. 1-86 in Zootaxa 4274 (1) on pages 52-54, DOI: 10.5281/zenodo.804182, {"references":["Boyle, W. K. (1974) A Revision of the Genus Sphenarium (Orthoptera: Pyrgomorphidae). McGill University, McGill, 214 pp.","Kevan, D. K. M. (1977) The American Pyrgomorphidae (Orthoptera). Revista de la Sociedad Entomologica Argentina, 36 (1 - 4), 3 - 28.","Sanabria-Urban, S., Song, H., Oyama, K., Gonzalez-Rodriguez, A., Serrano-Meneses, M. A. & Cueva del Castillo, R. (2015) Body size adaptations to altitudinal climatic variation in neotropical grasshoppers of the genus Sphenarium (Orthoptera: Pyrgomorphidae). Plos ONE, 10 (12), 1 - 24.","Pedraza-Lara, C., Barrientos-Lozano, L., Rocha-Sanchez, A. Y. & Zaldivar-Riveron, A. (2015) Montane and coastal species diversification in the economically important Mexican grasshopper genus Sphenarium (Orthoptera: Pyrgomorphidae). Molecular Phylogenetics and Evolution, 84, 220 - 231."]}

Sphenarium miztecum Sanabria-Urb��n, Song & Cueva del Castillo sp.n. (http://lsid.speciesfile.org/urn:lsid: Orthoptera.speciesfile.org:TaxonName:495094) Description. External morphology (Fig. 20 S, T): total body length ranging from 31.09 to 32.07 mm in females and from 24.87 to 33.35 mm in males; antennae ensiform, slightly shorter in females or longer than head and pronotum together in males; head conical notably longer than wide with oval eyes in both sexes; fastigium notably elongated, nearly as long as the interocular space in both sexes; tegmina strap-like in both sexes; subgenital plate of males somewhat tapered in the apex; dorsal ovipositor valves lanceolate slightly elongated towards the apex. Male genitalia: bridge of epiphallus as long or slightly longer than the length of lateral plates in most cases (Fig. 22 G-I). Ectophallus in dorsal view (Fig. 22 G-II) small with lateral borders of ramus convergent, straight; basal emargination of cingulum mostly notably developed closing the interspace between the apodemal plates. Ectophallus in posterior (Fig. 22 H) view with a conspicuous sclerotized hollow in the sheath notably closed; valves of cingulum small with form, not developed posteriorly (Fig. 22 I); inflections of supraramus notably developed laterally and dorsally whit dorsal borders evidently fused above the valves of cingulum (Fig. 22 H, arrow). Endophallus in lateral view (Fig. 22 D-III) with a short pseudoarch tightly joined to the valves of cingulum; aedeagal valves very small, sharply pointed apically, without apical spine; aedeagal valves and sclerites about half the length of dorsal inflections of endophallic apodemes. Colouration. Ground colours green or brown. Body uniformly coloured (Fig. 20 T) or with the following colour traits (Fig. 20 S): antennae dark brown; fastigium brownish; lateral postocular bands frequently present, wide and yellowish, whitish or bluish; dorsomedial line frequently present, wide and yellowish; dorsal shades brown to dark magenta, covering entirely the dorsal portion of pronotum and abdomen in most specimens; lateral shades often present, dark to light brown; lateral bands of blotches if present, cyan or distinct emerald green (Fig. 20 S, arrow); ventral bands of pronotum often present, wide, whitish to bluish; pronotum sometimes with small dots in the dorsal posterior margin; mesonotum brownish; lateral blotches of 1st abdominal segment frequently present, cyan to whitish; tegmina red or green; hind femora with medial area uniformly coloured and lower marginal area with distinct blue colouration, specially in the apex of femur; knees of hind femora darkly coloured laterally, lighter dorsally, black to bluish; hind tibia intense yellow or green. Diagnosis. S. miztecum sp.n. is very similar to its sister species S. adelinae sp.n. both in external and male genital morphology. Externally S. miztecum sp.n. differs from other Sphenarium species by its cyan or emerald green colourations in the lateral bands of blotches, when present (Fig. 20 S, arrow). The main difference in their male genital structures is that in S. miztecum sp.n. the dorsal borders of the inflections of supraramus are fused above the valves of cingulum. Distribution. This species is only known from a single locality from in the Pacific Cost of northern Oaxaca (Fig. 7 B). Additional fieldwork on this region is necessary to accurately delimit the distribution rages of this species. Material examined. Holotype m (Fig. 20 S) from Mexico: Oaxaca, ca. de Pinotepa Nacional Carr 200, 16.37195��N, - 98.177102��W, 216 m a.s.l., X-5-2011 (Sanabria-Urb��n S. # 18CPI [L18 MS1]). Paratypes from, 11 m, 6 f, same data as holotype. The holotype was deposited at IBUNAM and the paratypes were deposited at IBUNAM and TAMUIC. Taxonomic discussion. This species is closely related morphologically and genetically to S. adelinae sp.n. Nevertheless, this new species show a unique combination of morphologic traits (both in external and male genitalia structures) in combination with a well-supported monophyly and relatively high levels of genetic differentiation (Table 3). Therefore, we considered S. miztecum sp.n. as separate lineage from S. adelinae sp.n. Previously, we recognized specimens S. miztecum sp.n. as the putative new taxa Sphenarium sp.n. 3 (Sanabria- Urb��n et al. 2015). For other studies in the genus, this new species was unknown. Etymology. Named in honour of the Mixtecos, an ancient Native American people still living in the area where this species was found., Published as part of Sanabria-Urb��n, Salom��n, Song, Hojun, Oyama, Ken, Gonz��lez-Rodr��guez, Antonio & Castillo, Ra��l Cueva Del, 2017, Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae), pp. 1-86 in Zootaxa 4274 (1) on pages 60-61, DOI: 10.5281/zenodo.804182

Sphenarium borrei Bolivar, 1884 (http://lsid.speciesfile.org/urn:lsid: Orthoptera.speciesfile.org:TaxonName:37036) Sphenarium bruneri Bolivar, 1909 Description. External morphology (Figs 14 D, E, F, G; 15G, H): total body length ranging from 26.82 to 31.76 mm in females and from 21.08 to 30.62 mm in males; antennae filiform, notably shorter in females or slightly longer in males than head and pronotum together; head conical notably longer than wide and with oval eyes in both sexes; fastigium moderately elongated, nearly half the length of interocular space in females or notably elongated, nearly as long as the interocular space in males; tegmina spatula-like in both sexes; subgenital plate of males rounded moderately developed posteriorly; dorsal ovipositor valves lanceolate, notably elongated towards the apex. Male genitalia: bridge of epiphallus as long as the length of lateral plates (Fig. 16 D-I). Ectophallus in dorsal view (Fig. 16 D-II) broad at the base, with lateral borders of ramus convergent; basal emargination of cingulum notably developed towards the base reducing completely the interspace between the apodemal plates. Ectophallus in posterior view (Fig. 16 E) with a conspicuous sclerotized hollow in the sheath notably closed; inflections of supraramus notably developed anterolaterally; valves of cingulum triangular, stout and notably developed posteriorly (Fig. 16 F). Endophallus in lateral view (Fig. 16 D-III) with a short pseudoarch tightly joined to the valves of cingulum; aedeagal valves very small, tapered in the apex without apical spine; aedeagal valves and sclerites about �� the length of dorsal inflections of endophallic apodemes. Colouration. Ground colours vary from green, yellow or brown. Body uniformly coloured or with the following colour traits (Fig. 15 G, H): antennae frequently black or brown; fastigium often reddish to pinkish; lateral postocular bands frequently present, narrow, yellowish, whitish or pinkish; dorsomedial line absent; dorsal shades sometimes absent, if present almost entirely restricted to the head, metanotum and abdomen apex, brown or black; lateral shades often present, black or dark brown, principally restricted to head and pronotum; lateral bands of blotches absent; ventral bands of pronotum often present, narrow and pinkish; frequently posterior margin of pronotum pinkish; mesonotum is partially or entirely black; tegmina green, black or magenta; lateral blotches of 1st abdominal segment absent; fore and middle femora pinkish; sometimes hind femora with upper and lower medial areas black and whitish, respectively; knees of hind femora laterally black, dorsally reddish; hind tibia frequently black. Diagnosis. Externally this species sometimes resembles S. purpurascens and S. infernalis sp.n. However, S. borrei differs from these species principally by its notably elongated head (conical in both sexes) and dorsal ovipositor valves, as well as its pinkish colourations in the ventral and posterior margins of pronotum when present. At the level of male genitalia, S. borrei more closely resembles other species with a conspicuous sclerotized hollow in the sheath (all species described below). Nevertheless, S. borrei differs from other species in the genus by the following combination of male genitalia characters: lateral borders of ramus convergent, basal emargination of cingulum notably developed, sclerotized hollow of the sheath conspicuous and closed, inflections of supraramus notably developed anterolaterally, valves of cingulum triangular and stout, posteriorly developed, and aedeagal valves very small. Distribution. This species is distributed in elevations ranging from 304 to 2225 m a.s.l. and is principally restricted to the western portion of the Mexican Volcanic Belt in Colima, Guanajuato, Jalisco, Michoacan, Nayarit and Zacatecas, Mexico (Fig. 7 A). Material examined. S. borrei: lectotype m (Fig. 14 D) and paralectotype f (Fig. 14 E), from Mexico: Guanajuato (E. Duges); designation: Kevan (1960, unpublished results); location: Royal Institute of Natural Sciences of Belgium (RISCNB), Brusseles, Belgium. S. bruneri: lectotype m (Fig. 14 F) and paralectotype f (Fig. 14 G), from Mexico (Coneradt); designation: Kevan (1960, unpublished results); location: Spanish Institute of Entomology (SIE), Madrid, Spain. We could only examine the external morphology of all these type specimens. Additional material: 214 m, 178 f from 65 localities (Appendix Table 5). Taxonomic discussion. Bolivar (1884) originally described this species from one male and female syntypes from Guanajuato, Mexico. He recognized this species based on the morphology of head, antenna and ovipositor valves, as well as coloration patterns. For posterior taxonomic studies this species remain valid until M��rquez (1962) synonymised it as S. purpurascens despite he did not examined specimens of this species. Later, Boyle (1974) re-established the status of this species and this status remain stable for posterior molecular analysis (Pedraza-lara et al. 2015; Sanabria-Urb��n et al. 2015). According to our results, S. borrei is the most differentiated species in the genus in both morphological and genetic characters. Moreover, its well-supported monophyly and geographic restrictiveness support his recognition as a valid species. Sphenarium bruneri was only briefly described based on unstated number of specimens (Bolivar 1909), probably one male and two females from an unspecified Mexican locality. Originally, Bolivar (1909) stated a close relationship between this species and S. rugosum. However, he mainly considered highly variable external traits in differentiating S. brunerri from its congeners. The validity of S. bruneri is questionable and its taxonomic position has remained uncertain. This species has been considered as a synonym of S. histrio (Boyle 1974) or S. purpurascens (Kevan 1977). Nevertheless, considering the close resemblance in the morphology of head, tegmina and ovipositor valves between S. borrei and S. bruneri, it is probable that S. bruneri represents a junior synonym of S. borrei., Published as part of Sanabria-Urb��n, Salom��n, Song, Hojun, Oyama, Ken, Gonz��lez-Rodr��guez, Antonio & Castillo, Ra��l Cueva Del, 2017, Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae), pp. 1-86 in Zootaxa 4274 (1) on pages 45-46, DOI: 10.5281/zenodo.804182, {"references":["Bolivar, I. (1884) Monografia de los Pyrgomorfinos. Anales de la Sociedad Espanola de Historia Natural, 13, 1 - 500.","Bolivar, I. (1909) Orthoptera Fam. Acridiidae Subfam. Pyrgomorphinae. Genera insectorum, 90, 1 - 59.","Marquez, C. (1962) Estudios de las especies del genero Sphenarium basado en sus genitalia (Acrididae; Orthoptera), con la descripcion de una nueva especie. Anales de Instituto de Biologia UNAM Serie Zoologia, 33, 247 - 258.","Boyle, W. K. (1974) A Revision of the Genus Sphenarium (Orthoptera: Pyrgomorphidae). McGill University, McGill, 214 pp.","Sanabria-Urban, S., Song, H., Oyama, K., Gonzalez-Rodriguez, A., Serrano-Meneses, M. A. & Cueva del Castillo, R. (2015) Body size adaptations to altitudinal climatic variation in neotropical grasshoppers of the genus Sphenarium (Orthoptera: Pyrgomorphidae). Plos ONE, 10 (12), 1 - 24.","Kevan, D. K. M. (1977) The American Pyrgomorphidae (Orthoptera). Revista de la Sociedad Entomologica Argentina, 36 (1 - 4), 3 - 28."]}

Key to Sphenarium species 1. Tegmina spatula-like, wider in the apex than in the base (Fig. 4 G)............................................... 2 - Tegmina strap-like, as wide in the apex as in the base (Fig. 4 H)................................................ 12 2. Ectophallus in posterior view without sclerotized hollow in the inner-central portion of the sheath (e.g. Fig. 10 B, arrow). Endophallus in lateral view with short to very long aedeagal valves and elongated pseudoarch loosely joined to the valves of cingulum (e.g. Fig. 10 A-III, arrow)....................................................................... 3 - Ectophallus in posterior with a conspicuous sclerotized hollow in the sheath (e.g. Fig. 16 E, arrow). Endophallus in lateral view always with short aedeagal valves and short pseudoarch tightly joined to the valves of cingulum (e.g. Fig. 16 D-III, arrow)..................................................................................................... 11 3. Aedeagal valves with an apical spine in lateral view of endophallus (e.g. Fig. 10 C, arrow)............................ 4 - Aedeagal valves without apical spine in lateral view of endophallus (e.g. Fig. 12 F, arrow)........................... 6 4. Ectophallus in dorsal view with lateral borders of ramus strongly concave (e.g. Fig. 10 A-II, arrow). Apical spine of aedeagus slightly longer or shorter than the base of aedeagal sclerites. Widely distributed from the southern Altiplano to the Sierra Madre del Sur in central and southern Mexico (Fig. 7 A).......................................... S. purpurascens - Ectophallus in dorsal view with lateral borders of ramus convergent slightly rounded (e.g. Fig. 10 J-II, arrow)............. 5 5. Ectophallus in posterior view with inflections of supraramus reduced or not developed laterally and valves cingulum tonguelike (Fig. 10 K). Apical spine of aedeagus as long or shorter than the base endophallic apodemes (Fig. 10 J-III). Restricted to outer slope of the Sierra Madre del Sur in southern Mexico (Fig. 7 C)............................... S. zapotecum sp.n. - Ectophallus in posterior view with inflections of supraramus moderately developed laterally; valves of cingulum triangular to, Published as part of Sanabria-Urb��n, Salom��n, Song, Hojun, Oyama, Ken, Gonz��lez-Rodr��guez, Antonio & Castillo, Ra��l Cueva Del, 2017, Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae), pp. 1-86 in Zootaxa 4274 (1) on page 67, DOI: 10.5281/zenodo.804182

Sphenarium infernalis Sanabria-Urb��n, Song & Cueva del Castillo sp.n. (http://lsid.speciesfile.org/urn:lsid: Orthoptera.speciesfile.org:TaxonName:495096) Description. External morphology (Fig. 9 Q, R): body size ranging from 25.74 to 38.48mm in females and from 25.94 to 38.13mm in males; antennae filiform, slightly shorter in females or longer than head and pronotum together in males; head subtriangular-elongated slightly longer than wide in females or conical notably longer than wide in males, with oval eyes in both sexes; fastigium moderately elongated, nearly half the length of interocular space in females or notably elongated, nearly as long as the interocular space in males; tegmina spatula-like in both sexes; subgenital plate of males rounded, moderately developed posteriorly; dorsal ovipositor valves rounded towards the apex. Male genitalia: bridge of epiphallus moderately longer or as long as the length of lateral plates (Fig. 13 A-I). Ectophallus in dorsal view (Fig. 13 A-II) broad at the base, with lateral borders of ramus convergent; dorsal borders of ramus with lateral projections notably developed interiorly, closing the central membrane of ectophallus somewhat far from the sheath (Fig. 13 A-II, arrow); basal emargination of cingulum moderately developed; interspace between apodemal plates of cingulum moderately open to notably close. Ectophallus in posterior view (Fig. 13 B) without a conspicuous sclerotized hollow in the sheath; inflections of supraramus moderately developed with distal margins laterally or dorsally directed; valves of cingulum with distinct form (Fig. 13 B), small and slightly developed posteriorly (Fig. 13 C). Endophallus in lateral view (Fig. 13 A-III) with elongated pseudoarch, loosely joined to the valves of cingulum; aedeagal valves somewhat broad, medium-sized, with smooth ventral margins and moderately rounded in the apex (Fig. 13 B) without apical spine (Fig. 13 C); aedeagal valves and sclerites together about the same length than dorsal inflections of endophallic apodemes (Fig. 13 A-III). Colouration. Ground colours vary from green or brown. Body uniformly coloured (Fig. 9 R) or with the following colour traits (Fig. 9 Q): antennae mostly black; fastigium frequently reddish; lateral postocular bands frequently present, wide, whitish to yellow; dorsomedial line mostly absent, if present very narrow and yellowish; dorsal shades often absent, if present reduced and restricted to the apex of abdomen, brown to black; lateral shades often present, narrow, black or brown; lateral bands of blotches not evident; ventral bands of pronotum very narrow and yellow; mesonotum is partially or entirely black; whitish lateral blotches of 1st abdominal segment frequently present; tegmina green, black to magenta; generally hind femora uniformly coloured with knees laterally black and dorsally reddish; hind tibia usually black. Diagnosis. Externally this species more closely resembles S. macrophallicum, and S. crypticum sp.n. In most cases, females of S. infernalis sp.n. differ from females of other species by their rounded dorsal ovipositor valves. Nonetheless, the male genitalia of S. infernalis sp.n. clearly differ from other Sphenarium species by the following combination of traits: lateral borders of ramus convergent, dorsal borders of ramus with lateral projections developed towards the central membrane that are not evident in other species in the genus, inflections of supraramus moderately developed, and aedeagal valves somewhat broad, medium-sized, moderately rounded in the apex. Distribution. This species is apparently restricted to the western portion of the Balsas River Basin in elevations ranging from 335 to 1450 m a.s.l. Colima, Jalisco and Michoacan, Mexico (Fig. 7 A). Material examined. Holotype m (Fig. 9 Q) from Mexico: Jalisco, 5mi. NE. Tecalitlan, 19.540895��N, - 103.3147204��W, 4250ft, XI-25-1958 (T. J. Cohn #313); measurements: BS = 28.01 mm, FL = 1.03 mm, PL = 5.99 mm, HF = 13.30 mm. Paratypes from Mexico: Jalisco: 1 m, 1 f, same data as holotype; 3 m, 2 f, 1mi S Pihuamo, 869m a.s.l., XI-26-1958 (T.J. Conh #314); 2 m, 2 f, 6mi NE Tecalitlan, 4250ft, XI-30-1958 (T.J. Cohn #331); 1 m, 1 f, Tecalitlan, 19.52045714��N, - 103.3007948��W, 1213 m a.s.l., X-2-2013 (Sanabria-Urban #P78); 2 m, 4 f, Pihuamo, 19.24430234��N, - 103.4002227��W, 800 m a.s.l., X-2-2013 (Sanabria-Urban S. & Rivera-Ortiz F. #P79 [L58 MS1]). Colima: 1 m, 1 f, 9 mi E Colima, 1620ft, XI-22-1958 (T.J. Cohn #328); 1 m, 6 mi E Colima, 1250ft, XI-29-1958 (T.J. Cohn #327); 1 m, 1 f, Entrada a Colima, 19.18950356��N, - 103.6827591��W, 468 m a.s.l., X-3- 2013 (Sanabria-Urban S. & Rivera-Ortiz F. #M58). Michoacan: 1 m, 1 f, 15 mi W Apatzingan (2 mi E. Santa Ana), 1600ft, XII-2-1958 (T.J. Cohn #340); 1 m, 23rd mi WSW Ario de Rosales, 4200ft, XII-4-1958 (T.J. Cohn #348); 2 m, 1 f, 16.7mi W Apatzingan, 1100ft, IX-26-1959 (Cantral & Cohn #179); 1 m, 11mi S Uruapan on 37, IX-8-1981 (Otte, Azuma, Newlin #57); 1 m, 26rd mi NE Arteaga on Hwy. 37, 3 rd. mi. SW Rancho Nuevo, 2060ft, XI-3-1974 (T.J. & J. W. Cohn #121); 10 m, 8 f, Las Majadas Carr 120, 19.134653��N, - 102.463237��W, 301 m a.s.l., IX-23-2012 (Sanabria-Urban S. #P43 [L57 MS1]); 1 m, 1 f, Periban, 19.55953625��N, - 102.4384873��W, 1450 m a.s.l., X-1-2013 (Sanabria-Urban S. & Rivera-Ortiz F. #P75). The holotype was deposited at UMMZ and the paratypes were deposited at the IBUNAM, TAMUIC and UMMZ. Taxonomic discussion. This species is closely related morphologically to S. rugosum. In deed material of this new species was identified as S. rugosum in previous studies (Boyle 1974; Kevan 1977). Nevertheless, S. infernalis sp.n. shows a unique combination of morphologic traits, in both external and male genitalia structures, a wellsupported monophyly (Fig. 2 b), relatively high levels of interspecific genetic differentiation (Table 3). All these lines of evidence support the recognition of S. infernalis sp.n. as a valid species. Previously, we identified specimens of S. infernalis sp.n. as Sphenarium sp.n. 8 (Sanabria-Urb��n et al. 2015). For other studies in the genus this new species was unknown. Etymology. The specific name ��� infernalis ��� is derived from Latin and means ���belonging to the lower regions���. It refers to the distribution of this species in the lowest portion of the Balsas River Basin, which is also one of the warmest regions in Mexico., Published as part of Sanabria-Urb��n, Salom��n, Song, Hojun, Oyama, Ken, Gonz��lez-Rodr��guez, Antonio & Castillo, Ra��l Cueva Del, 2017, Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae), pp. 1-86 in Zootaxa 4274 (1) on pages 37-39, DOI: 10.5281/zenodo.804182, {"references":["Boyle, W. K. (1974) A Revision of the Genus Sphenarium (Orthoptera: Pyrgomorphidae). McGill University, McGill, 214 pp.","Kevan, D. K. M. (1977) The American Pyrgomorphidae (Orthoptera). Revista de la Sociedad Entomologica Argentina, 36 (1 - 4), 3 - 28.","Sanabria-Urban, S., Song, H., Oyama, K., Gonzalez-Rodriguez, A., Serrano-Meneses, M. A. & Cueva del Castillo, R. (2015) Body size adaptations to altitudinal climatic variation in neotropical grasshoppers of the genus Sphenarium (Orthoptera: Pyrgomorphidae). Plos ONE, 10 (12), 1 - 24."]}

Sphenarium histrio Gerstaecker, 1873 (http://lsid.speciesfile.org/urn:lsid: Orthoptera.speciesfile.org:TaxonName:37005) Sphenarium carinatum Bolivar, 1904 Description. External morphology (Figs. 15 Q, R, S, T; 18C, D; 20A, B, C, D, E, F, G, H, I, J): total body length ranging from 22.01 to 41.8 mm in females and from 18.57 to 38.87 mm in males. In most cases: antennae filiform, slightly shorter in females or longer than head and pronotum together in males; head subtriangular-elongated slightly longer than wide in females or conical notably longer than wide in males, with oval eyes in both sexes; fastigium moderately elongated, nearly half the length of interocular space in females or notably elongated, nearly as long as the interocular space in males; tegmina strap-like in both sexes; subgenital plate of males somewhat tapered in the apex; dorsal ovipositor valves rounded or lanceolate slightly elongated towards the apex. Male genitalia: bridge of ectophallus slightly longer than the length of lateral plates (Figs. 19 G-I, J-I; 21A-I, D-I, G-I). Ectophallus in dorsal view small with lateral borders of ramus convergent, straight (morphotypes 1, 3, and 4; Figs. 19 G-II; 21A-II, D-II, respectively) or slightly rounded (morphotypes 2 and 5; Figs. 19 J-II; 21G-II, respectively); basal emargination of cingulum notably or slightly developed; interspace between the apodemal plates notably closed. Ectophallus in posterior view with a conspicuous sclerotized hollow in the sheath closed (morphotype 4; Fig. 21 E) or moderately open (other morphotypes; Figs. 19 H, K; 21B, H); inflections of supraramus moderately developed, with distal borders ventrally (morphotypes 1, 2, and 5; Figs. 19 H, K; 21H, respectively) or laterally directed (morphotypes 3 and 4; Fig. 19 B, E, respectively); valves of cingulum principally triangular but with distinct variations, each morphotype with its own form; Figs. 19 H, K; 21B, E, H), small (morphotypes 1-4) or moderately large (morphotype 5). Ectophallus in lateral view with valves of cingulum moderately developed dorsoposteriorly (morphotype 5; Fig. 21 I) or not developed posteriorly (morphotypes 1-4; Figs. 19 I, L; 21C, F, respectively). Endophallus in lateral view (Figs. 19 G-III, J-III; 21A-III, D-III, G-III) with a short pseudoarch tightly joined to the valves of cingulum; aedeagal valves very small, tapered in the apex without apical spine; aedegala valves and sclerites as long as (morphotype 5, Fig. 21 G-III) or about �� (other morphotypes; Figs. 19 G-III, J-III; 21A-III, D-III) the length of dorsal inflections of endophallic apodemes. Colouration. Ground colours vary from olive green, yellow or brown. Body uniformly coloured (Fig. 15 R, T; 20B) or with the following colour traits (Figs. 15 Q, S; 20A, B, C, D, E, F): antennae black, gray or dark blue; fastigium blue, brown or black; lateral postocular bands frequently present, narrow and yellowish; dorsal medial line frequently present, wide, yellowish or whitish; dorsal shades frequently present black, brown to dark blue, in southern populations covering entirely the dorsal portion of the body (Fig. 20 C, D); lateral shades often present, black or dark blue; lateral bands of blotches frequently present, mostly reddish, in some populations in northern ranges of the species yellowish (Fig. 15 S); ventral bands of pronotum often present, wide, whitish or bluish; pronotum with white lateral carinas and small strips and dots in the posterior margin; mesonotum dorsally red; frequently mesonotum and metanotum laterally white or yellow; lateral blotches of 1st abdominal segment frequently present and whitish; tegmina green, red, magenta or blue; generally hind femora uniformly coloured with knees laterally black, dorsally bluish or brownish; hind tibia green, brown, reddish or blue. Diagnosis. Externally S. histrio is very similar to S. mexicanum, S. totonacum sp.n., and S. occidentalis sp.n. Nevertheless, the phallic structures of S. histrio differ from other species as follows: ectophallus small, conspicuous sclerotized hollow of the sheath moderately open or close, inflections of supraramus moderately developed with distal portions ventrally directed in most cases (except in morphotypes 3 & 4), and valves of cingulum with distinct form not developed or slightly developed dorsoposteriorly. Distribution. This species is distributed in elevations ranging approximately from 15 to 2225 m a.s.l. in Chiapas, Guerrero, Oaxaca, Tabasco, Veracuz, in southern Mexico; and in the states of Huehuetenango and Santa Rosa in Guatemala (Fig. 7 B). We observed that the five morphotypes identified within the species are apparently restricted to different geographic areas. The morphotype 1 is mainly distributed across the southern Pacific Cost and the outer slope of the Sierra Madre del Sur in Oaxaca. The morphotype 2 is distributed in the Pacific Cost south of the Isthmus of Tehuantepec, as well as in the mountain ranges of Chiapas and north-western Guatemala. The remaining morphotypes are more geographically restricted: the morphotype 3 is distributed principally in the central Valleys of Oaxaca in the Sierra Madre del Sur (Fig. 7 B, C), the morphotype 4 was observed in the southeastern portion of the Sierra Madre del Sur also in Oaxaca (Fig. 7 B, C), and morphotype 5 was observed only few localities in the Pacific Cost of Chiapas, Mexico (Fig. 7 B). Overall, here we recognised smaller distribution ranges for S. histrio than in Boyle (1974) and Kevan (1977). Within the examined material we identified two S. histrio males from Sonora (morphotype 2) and Sinaloa (morphotype 3). However, we considered these specimens as mislabelled records considering that multiple collectors, including us, have not recorded S. histrio beyond the above-specified limits. Moreover, according to the revised material, S. histrio is apparently absent from the lowlands of the Isthmus of Tehuantepec where S. mexicanum is distributed. Additional fieldwork will clarify if these two species are sympatric in this geographic region. Material examined. S. histrio: holotype m (Fig. 18 C) from Mexico; designation by monotypy; location: BZM. S. carinatum: holotype m (Fig. 18 D) from Guatemala: Santa Rosa, Testuaco; designation by monotypy; location: SIE. We could examine only the external morphology of this type material. Additional material: 398 m, 326 f, form 91 Mexican and Guatemalan localities (Appendix Table 5). Taxonomic discussion. Gerstaecker (1873) originally described S. histrio based on a single male from an unknown Mexican locality considering mainly coloration traits as diagnostic characters for this species. Bolivar (1884) re-described this species based on a male and a female, but also from an unspecified Mexican locality. The validity of this species remained constant in posterior taxonomic studies. Hebard (1932) recognized a closer relationship between S. histrio and S.mexicanum based on their morphological resemblance. Later, M��rquez (1962) and Boyle (1974) identified differences in endophallic morphology between these two species. However, Boyle (1974) and Kevan (1977) only recognized S. histrio as a subspecies of S. mexicanum (S. mexicanum histrio). Recently, Pedraza-Lara et al. (2015) and Sanabria-Urb��n et al. (2015) have proposed the recognition of S. histrio as an independent and valid species within the genus based on morphologic and genetic evidence. In our phylogenetic analysis S. histrio was recovered as a paraphyletic species despite its notable morphologic cohesiveness. Moreover, we observed notable genetic (with CO1 P-distance > 3%) and morphological differentiation in the male genitalia between S. histrio and S. mexicanum. Therefore, here we also agreed in considering S. histrio as a valid species. Moreover, during this revision we examined several specimens collected near to the reported localities of two putative new taxa recently identified, Sphenarium sp. Oax9 and Sphenarium sp. Oax2 (Pedraza-lara et al. 2015) (L11, L167, L172, and L173; Appendix Table 5). The male genitalia of these specimens were similar to those of other S. histrio individuals. Moreover, CO1 sequences of these taxa intermingled with our samples of S. histrio (Fig. 11). Therefore, we consider that these putative new taxa probably represent part of S. histrio. Sphenarium carinatum was originally described based on a single male from an uncertain Guatemalan locality, Testuaco, probably Tecuaco (Bolivar 1904). In later taxonomic studies this species was synonymised with S. m. histrio (Boyle 1974; Kevan 1977). In this study we examined multiple specimens from north-western Guatemala; which were similar to other S. histrio individuals in southern Mexico. Therefore, in the lack of additional evidence we agree in recognising S. carinatum as a synonym of S. histrio., Published as part of Sanabria-Urb��n, Salom��n, Song, Hojun, Oyama, Ken, Gonz��lez-Rodr��guez, Antonio & Castillo, Ra��l Cueva Del, 2017, Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae), pp. 1-86 in Zootaxa 4274 (1) on pages 51-52, DOI: 10.5281/zenodo.804182, {"references":["Gerstaecker, A. (1873) Acridiodea nonnulla nova insigniora descropsit. Entomologische Zeitung, 34, 185 - 197.","Bolivar, I. (1904) Notas sobre los Prygomorfidos (Pyrgomorphidae). Boletin de la Sociedad Espanola de Historia Natural, 4, 306 - 326.","Boyle, W. K. (1974) A Revision of the Genus Sphenarium (Orthoptera: Pyrgomorphidae). McGill University, McGill, 214 pp.","Kevan, D. K. M. (1977) The American Pyrgomorphidae (Orthoptera). Revista de la Sociedad Entomologica Argentina, 36 (1 - 4), 3 - 28.","Bolivar, I. (1884) Monografia de los Pyrgomorfinos. Anales de la Sociedad Espanola de Historia Natural, 13, 1 - 500.","Hebard, M. (1932) New species and records of Mexican Orthoptera. Transactions of the American Entomological Society, 58 (3), 201 - 371.","Marquez, C. (1962) Estudios de las especies del genero Sphenarium basado en sus genitalia (Acrididae; Orthoptera), con la descripcion de una nueva especie. Anales de Instituto de Biologia UNAM Serie Zoologia, 33, 247 - 258.","Pedraza-Lara, C., Barrientos-Lozano, L., Rocha-Sanchez, A. Y. & Zaldivar-Riveron, A. (2015) Montane and coastal species diversification in the economically important Mexican grasshopper genus Sphenarium (Orthoptera: Pyrgomorphidae). Molecular Phylogenetics and Evolution, 84, 220 - 231.","Sanabria-Urban, S., Song, H., Oyama, K., Gonzalez-Rodriguez, A., Serrano-Meneses, M. A. & Cueva del Castillo, R. (2015) Body size adaptations to altitudinal climatic variation in neotropical grasshoppers of the genus Sphenarium (Orthoptera: Pyrgomorphidae). Plos ONE, 10 (12), 1 - 24."]}

Sphenarium minimum Bruner, 1906 (http://lsid.speciesfile.org/urn:lsid: Orthoptera.speciesfile.org:TaxonName:36988) Sphenarium affine Bruner, 1906 Description. External morphology (Figs. 8 G, H; 9O, P): total body length ranging from 23.67 to 28.57 mm in females and 18.05 to 24.89 mm in males; antennae filiform, slightly shorter in females or notably longer than head and pronotum together in males; head subtriangular-elongated slightly longer than wide in females or conical notably longer than wide in males, with oval eyes in both sexes; fastigium moderately elongated, nearly half the length of interocular space in both sexes; tegmina spatula-like in both sexes; subgenital plate of males rounded moderately developed posteriorly; dorsal ovipositor valves lanceolate, notably elongated towards the apex. Male genitalia: bridge of epiphallus as long as the length of lateral plates in most cases (Fig. 12 J-I). Ectophallus in dorsal view (Fig. 12 J-II) with lateral borders of ramus strongly concave; basal emargination of cingulum moderately developed; interspace between apodemal plates of cingulum moderately open. Ectophallus in posterior view (Fig. 12 K) without a conspicuous sclerotized hollow in the sheath; inflections of supraramus moderately developed with distal margins laterally directed in most cases; valves of cingulum notably small, triangular and slightly developed posteriorly (Fig. 12 K, L). In lateral view of endophallus (Fig. 12 J-III) pseudoarch elongated loosely joined the valves of cingulum; aedeagal valves medium sized with smooth ventral margins and moderately rounded in the apex, without apical spine (Fig. 12 K, L); aedeagal valves and sclerites together about 1 �� fold the length of dorsal inflections of endophallic apodemes. Colouration. Ground colours vary from green, yellow or brown. Body uniformly coloured or with the following colour traits: antennae and fastigium frequently brown; lateral postocular bands frequently present, narrow and yellow; dorsomedial line generally absent, if present very narrow almost restricted to the abdomen, whitish, yellowish or pinkish; dorsal shades frequently absent, if present very narrow in head and thorax, wider in the dorsal portion of abdomen, light to dark brown; lateral shades often absent, if present very narrow, black or dark brown, almost restricted to head and abdomen; lateral bands of blotches not evident; ventral bands of pronotum often absent, if present very narrow and yellowish; mesonotum partially or entirely black; lateral whitish blotches of 1st abdominal segment sometimes evident; in most cases hind femora uniformly coloured, sometimes with lower medial area whitish or yellowish and brown knees; hind tibia frequently yellow or pale orange. Diagnosis. S. minimum closely resembles S. planum, S. purpurascens, S. tarascum sp.n., and S. zapotecum sp.n. in both external and genital morphology. Externally S. minimum only differs from the former two species by its more elongated head (in both sexes) and its lanceolate and more notably elongated dorsal ovipositor valves. Nevertheless, more conspicuous differences exist between male genital structures of all these species. Sphenarium minimum differs from these species by the following combination of male genital traits: ectophallus with lateral borders of ramus notably concave, valves of cingulum triangular and notably small, medium-sized aedeagal valves and moderately rounded in the apex without apical spine, and aedeagal valves and sclerites about 1 �� fold the length of dorsal inflections of endophallic. Distribution. This species is distributed in elevations ranging from approximately 1000 to 1596 m a.s.l. in the outer slope of southern portion of the Sierra Madre Oriental in Veracruz, Mexico (Fig. 7 A). Material examined. S. minimum: lectotype m (Fig. 8 G) from Mexico: Veracruz, Jalapa, XII (O.W. Barrett); S. affine: lectotype m (Fig. 8 H) from Mexico: Veracruz, Orizaba, XI-1887 (L. Bruner). Designation: Rehn and Hebard (1912) and location: ANSP for both type specimens. We examined only the external morphology of this type material. Additional material: 7 m, 5 f, from Jalapa (L76); 2 m from Orizaba (L292), 35 m, 26 f, from 12 adjacent localities in Veracruz (Appendix Table 5). Taxonomic discussion. Bruner (1906) originally described this species apparently based on a single male from Jalapa, Veracruz. Bolivar (1909) and Hebard (1932) recognized S. minimum as a valid species. Particularly, the later author proposed a closer relationship between S. minimum, S. histrio and S. mexicanum mainly based on their head morphology. Boyle (1974) also identified differences in head morphology and colouration patterns between S. minimum and S. purpurascens. However, this author only recognized the former taxon as a subspecies of the later, S. purpurascens minimum, principally based on the similarity between their epiphallic and ectophallic structures. Recently, Pedraza-Lara et al. (2015) and Sanabria-Urb��n et al. (2015) proposed again the re-establishment of S. minimum as valid species based principally on genetic evidence. In this study we found that despite the notable similarity in the male genitalia between S. minimum and S. purpurascens the former taxon shows a unique combination of external and male genitalia characters, a well-supported monophyly (Fig. 2), and a well-separated geographic distribution from S. purpurascens. All these lines of evidence support the recognition of S. minimum as a valid species. Sphenarium affine was only briefly described by Bruner (1906) apparently based on one female and male from Orizaba, Veracruz. Originally, Bruner (1906) recognized a close relationship between S. affine and S. minimum, only differentiating the former from the later in highly variable traits of head morphology. Later S. affine was synonymised within S. marginatum (Hebard 1932), despite the notable external differentiation between both taxa. Posteriorly, Boyle (1974) synonymized S. affine within S. p. minimum. In this study we examined several specimens collected in type localities of S. minimum and S. affine; which were practically similar among them in their male genital morphology. Therefore, we also recognise S. affine as a synonym of S. minimum., Published as part of Sanabria-Urb��n, Salom��n, Song, Hojun, Oyama, Ken, Gonz��lez-Rodr��guez, Antonio & Castillo, Ra��l Cueva Del, 2017, Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae), pp. 1-86 in Zootaxa 4274 (1) on pages 36-37, DOI: 10.5281/zenodo.804182, {"references":["Bruner, L. (1906) Insecta. Orthoptera. Vol. II. In: Godman, F. D. & Salvin, O. (Eds.), Biologia Centrali-Americana. R. H. Porter, London, pp. 199 - 207.","Rehn, J. A. G. & Hebard, M. (1912) Fixation of the single type (Lectotype) specimens of species of American Orthoptera. Proceedings of the Academy of Natural Sciences of Philadelphia, 64 (1), 60 - 183.","Bolivar, I. (1909) Orthoptera Fam. Acridiidae Subfam. Pyrgomorphinae. Genera insectorum, 90, 1 - 59.","Hebard, M. (1932) New species and records of Mexican Orthoptera. Transactions of the American Entomological Society, 58 (3), 201 - 371.","Boyle, W. K. (1974) A Revision of the Genus Sphenarium (Orthoptera: Pyrgomorphidae). McGill University, McGill, 214 pp.","Pedraza-Lara, C., Barrientos-Lozano, L., Rocha-Sanchez, A. Y. & Zaldivar-Riveron, A. (2015) Montane and coastal species diversification in the economically important Mexican grasshopper genus Sphenarium (Orthoptera: Pyrgomorphidae). Molecular Phylogenetics and Evolution, 84, 220 - 231.","Sanabria-Urban, S., Song, H., Oyama, K., Gonzalez-Rodriguez, A., Serrano-Meneses, M. A. & Cueva del Castillo, R. (2015) Body size adaptations to altitudinal climatic variation in neotropical grasshoppers of the genus Sphenarium (Orthoptera: Pyrgomorphidae). Plos ONE, 10 (12), 1 - 24."]}

Sphenarium totonacum Sanabria-Urb��n, Song & Cueva del Castillo sp.n. (http://lsid.speciesfile.org/urn:lsid: Orthoptera.speciesfile.org:TaxonName:495099) Description. External morphology (Fig. 20 O, P): total body length ranging from 32.77 to 40.47 mm in females and from 30.04 to 38.75 mm in males; antennae weakly ensiform, slightly shorter in females or longer than head and pronotum together in males; head conical notably longer than wide with oval eyes in both sexes; fastigium notably elongated, nearly as long as the interocular space in both sexes; tegmina strap-like in both sexes; subgenital plate of males somewhat tapered in the apex; dorsal ovipositor valves lanceolate, elongated towards the apex. Male genitalia: bridge of epiphallus slightly longer than the length of lateral plates (Fig. 22 A-I). Ectophallus in dorsal view (Fig. 22 A-II) small with lateral borders of ramus convergent, somewhat straight; basal emargination of cingulum notably developed; interspace between the apodemal plates notably closed. Ectophallus in posterior (Fig. 22 K) view with a conspicuous sclerotized hollow in the sheath closed; valves of cingulum drop-like (Fig. 22 B, arrow); inflections of supraramus notably developed whit distal borders ventrolateral directed. Ectophallus in lateral view (Fig. 22 C) with valves of cingulum not developed posteriorly. Endophallus in lateral view (Fig. 22 A- III) with a short pseudoarch tightly joined to the valves of cingulum; aedeagal valves small, sharply pointed apically, without apical spine; aedeagal valves and sclerites about half the length of dorsal inflections of endophallic apodemes. Colouration. Ground colours vary from green or brown. Body uniformly coloured or with the following colour traits (Fig. 20 O, P): antennae black, gray or dark brown; lateral postocular bands frequently present, narrow and yellowish; dorsomedial line frequently absent, if present very narrow, yellowish or whitish; dorsal shades absent; lateral shades often present and black; lateral bands of blotches frequently present and reddish; ventral bands of pronotum often present, wide and whitish; pronotum with white small stripes and dots in the posterior margin; dorsal portion of pronotum, metanotum and 1st abdominal segment with darker green colouration; mesonotum red; lateral blotches of 1st abdominal segment if present whitish; generally hind femora uniformly coloured with knees laterally black, dorsally brownish; hind tibia reddish. Distribution. This species is apparently restricted to the outer slope of the Sierra Madre Oriental in elevations ranging from 440 to 1145 m a.s.l. in Veracruz and Puebla, Mexico (Fig. 7 A). Diagnosis. Externally this species closely resembles S. mexicanum, whereas it is more similar to S. histrio in their male genitalia. Nevertheless, S. totonacum sp.n. differs from other Sphenarium species by the following male genitalia characters: sheath of ectophallus with a conspicuous sclerotized and closed hollow, inflexion of supraramus notably developed and ventrolateral directed, and valves of cingulum with distinct drop-like form. Material examined. Holotype m (Fig. 20 O) from Mexico: Veracruz, Plan de Hayas, 19.74138��N, - 96.64531��W, 1145 m a.s.l., XI-3-2012 (Sanabria-Urb��n S. & J��menez-Arcos V. H. # P72 [L19 MS1]); measurements: BS = 34.63 mm, FL = 1.38 mm, PL = 7.27 mm, HF = 15,22 mm. Paratypes from Mexico: Veracruz: 5 M, 5 F same data as holotype; 3 m, 3 f, Km 21 Carr. 131, ca. 7km SO de Tlapacoyan, 19.90252398��N, - 97.22993698��W, 751 m a.s.l., IX-19-2015 (Sanabria-Urb��n S. # M010-L52); 1 f, Tlapacoyan Eytepequez, 11-9- 1995 (Delgadillo J.). The holotype was deposited at IBUNAM and the paratypes were deposited at the IBUNAM and TAMUIC. Additional material: 13 m, 16 f, from seven localities (Appendix Table 5). Taxonomic discussion. This species is also closely related morphologically to S. histrio. Indeed, specimens of this new species were identified as an isolated population of S. m. histrio in the cost of the Golf of Mexico (Boyle 1974; Kevan 1977). Previously we identified specimens of this new species as Sphenarium sp.n. 4 (Sanabria-Urb��n et al. 2015). For other studies in the genus this species was unknown. In this study we found that S. totonacum sp.n. shows a unique combination and notably different male genitalia structures. Moreover, this new species has a wellsupported monophyly, shows relatively high levels of interspecific genetic differentiation (Table 3), and it is considerably separated geographically from S. histrio. All these lines of evidence support the recognition of S. totonacum sp.n. as a valid species. Etymology. Named in honour of the Totonacos, an ancient Native American people still living in the area where this species was found., Published as part of Sanabria-Urb��n, Salom��n, Song, Hojun, Oyama, Ken, Gonz��lez-Rodr��guez, Antonio & Castillo, Ra��l Cueva Del, 2017, Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae), pp. 1-86 in Zootaxa 4274 (1) on pages 54-58, DOI: 10.5281/zenodo.804182, {"references":["Boyle, W. K. (1974) A Revision of the Genus Sphenarium (Orthoptera: Pyrgomorphidae). McGill University, McGill, 214 pp.","Kevan, D. K. M. (1977) The American Pyrgomorphidae (Orthoptera). Revista de la Sociedad Entomologica Argentina, 36 (1 - 4), 3 - 28.","Sanabria-Urban, S., Song, H., Oyama, K., Gonzalez-Rodriguez, A., Serrano-Meneses, M. A. & Cueva del Castillo, R. (2015) Body size adaptations to altitudinal climatic variation in neotropical grasshoppers of the genus Sphenarium (Orthoptera: Pyrgomorphidae). Plos ONE, 10 (12), 1 - 24."]}