taxonID	type	description	language	source
A12C6E25AC103B702CBED4E5FE0FA85C.taxon	description	Description. External morphology (Figs. 8 A, B; 9 A, 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; 10 A-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 ½ 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 1 st 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.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC103B702CBED4E5FE0FA85C.taxon	diagnosis	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.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC103B702CBED4E5FE0FA85C.taxon	distribution	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.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC103B702CBED4E5FE0FA85C.taxon	materials_examined	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-Urbán 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.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC1B3B762CBED2FBFDA3A910.taxon	description	Description. External morphology (Fig. 9 G, H): total body length ranging from 22.13 to 26.73 mm in females and from 19.29 to 24.26 mm 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 lanceolate, moderately elongated towards the apex. Male genitalia: bridge of epiphallus as long or slightly longer than the length of lateral plates in most cases (Fig. 10 J – I). Ectophallus in dorsal view (Fig. 10 J-II) with lateral borders convergent slightly rounded; basal emargination of cingulum moderately developed; interspace between apodemal plates of cingulum moderately open. Ectophallus in posterior view (Fig. 10 K) without a conspicuous sclerotized hollow in the sheath; inflections of supraramus reduced or not developed laterally (Fig. 10 K, arrow); valves of cingulum with unique tongue-like form, notably developed posteriorly (Fig. 10 L). Endophallus in lateral view (Fig. 10 J-III) with an elongated pseudoarch loosely joined to the valves of cingulum; aedeagal valves long with smooth ventral borders and an apical spine slightly longer than the width of the base of aedeagal sclerites; aedeagal valves and sclerites together about twice the length of dorsal inflections of endophallic apodemes. Colouration. Ground colours varying from green to brown. Body uniformly coloured with ground colours (Fig. 9 H) or with the following colour traits (Fig. 9 G): antennae generally light brown; fastigium brown to black; lateral postocular bands whitish; dorsomedial line narrow and pinkish in eastern populations or wide and yellowish in western populations; dorsal shades black, grey or brown, frequently covering partially the dorsal portion of the body; lateral black shades generally absent or restricted to head; lateral bands of blotches not evident; ventral bands of pronotum generally absent, instead ventral borders of pronotum lightly-coloured; mesonotum partially or entirely black; lateral blotches of 1 st abdominal segment if present white; hind femora uniformly coloured with knees laterally black, dorsally brownish; hind tibia orange.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC1B3B762CBED2FBFDA3A910.taxon	diagnosis	Diagnosis. Externally this species closely resembles S. purpurascens and S. variabile. However, S. zapotecum sp. n. generally differs from these two species by its more elongated head in both sexes. At the male genitalia level S. zapotecum sp. n. more closely resembles S. purpurascens and S. tarascum sp. n., which also show an apical spine in the aedeagus and lack the sclerotized hollow in the sheath of ectophallus. Nevertheless, S. zapotecum sp. n., differs from these latter species by its reduced or undeveloped inflections of supraramus, as well as the unique form of its valves of cingulum notably projected posteriorly.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC1B3B762CBED2FBFDA3A910.taxon	distribution	Distribution. This species is apparently restricted to the outer southern slope of the Sierra Madre del Sur in Oaxaca, Mexico, occurring in elevations ranging from 1016 to 1457 m a. s. l. (Fig. 7 B, C).	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC1B3B762CBED2FBFDA3A910.taxon	materials_examined	Material examined. Holotype m (Figs. 9 G; 10 J, K, L) from Mexico: Oaxaca, Pluma Hidalgo (1), 15.93987876 ° N, - 96.42996051 ° W, 1153 m a. s. l., XII- 11 - 2013 (Sanabria-Urbán S., Fontana P. & Mariño-Pérez R. # L 27); measurements: BS = 23.74 mm, FL = 1.29 mm, PL = 4.68 mm, HF = 11.77 mm. Paratypes from Mexico: Oaxaca: 4 m, 3 f, same data as holotype; 2 m, 2 f, Carr. 175 Km. 172, XII- 12 - 2013, 16.01864845 ° N, - 96.5303105 ° W, 1457 m a. s. l. (Sanabria-Urbán S., Fontana P. & Mariño-Pérez R. # L 31); 1 m, Oaxaca road ca. 85 km N Pto. Angel, IX- 1 - 1981 (Otte, Azuma & Newlin # 43); 2 m, 1 f, 24 - 25 mi. N Pto. Escondido rd. to Oaxaca, IX- 2 - 1981 (Otte, Azuma & Newlin # 45). The holotype was deposited at IBUNAM and the paratypes were deposited at the ANSP, IBUNAM and TAMUIC. Additional material: 5 m, 4 f, from three additional localities (Appendix Table 5). Taxonomic discussion. We observed that S. zapotecum sp. n. is mainly related morphologically and genetically to S. purpurascens. Nevertheless, its unique combination of morphologic traits (both in external and male genitalia structures), as well as its geographic isolation supports its recognition as an independent species within the genus. Sphenarium zapotecum sp. n. was not included in previous studies on the genus except for Pedraza-Lara et al. (2015), who identified specimens from a locality within the ranges of this new species as Sphenarium sp. Oax 6. Using the CO 1 sequences of Pedraza-Lara et al. (2015) and the present study, the only locus represented in both studies, we conducted a combined Bayesian phylogenetic analysis, in which all Sphenarium sp. Oax 6 samples strongly clustered (PP> 0.95) with S. zapotecum sp. n. samples (Fig. 11). Considering their geographical and phylogenetic proximity it is probable that Sphenarium sp. Oax 6 and S. zapotecum sp. n. represent the same species.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC1B3B762CBED2FBFDA3A910.taxon	etymology	Etymology. Named in honour of the Zapotecos, an ancient Native American people still living in the area where this species is distributed.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC193B772CBED403FE3AA95F.taxon	description	Description. External morphology (Fig. 9 I, J): total body length ranging from 21.41 to 30.07 mm in females and from 19.44 to 25.6 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 females or notably elongated, nearly as long as the interocular space in males; tegmina are 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 or slightly longer than the length of lateral plates in most cases (Fig. 12 A-I). Ectophallus in dorsal view (Fig. 12 A-II) with lateral borders of ramus convergent slightly rounded; basal emargination of cingulum moderately developed; interspace between apodemal plates of cingulum moderately open. Ectophallus in posterior view (Fig. 12 B) without a conspicuous sclerotized hollow in the sheath; inflections of supraramus moderately developed with distal margins dorsally directed; valves of cingulum triangular to slightly quadrangular, moderately developed posteriorly (Fig. 12 C). Endophallus in lateral view (Fig. 12 A-III) with elongated pseudoarch, loosely joined to the valves of cingulum; aedeagal valves long with smooth ventral borders and an apical spine slender, slightly bent, and notably longer than the width of the base of aedeagal sclerites; aedeagal valves and sclerites together about 1 ½ fold the length of dorsal inflections of endophallic apodemes. Colouration. Ground colours varying from green to brown; body uniformly coloured (Fig. 9 J) or with the following colour traits (Fig. 9 I): antennae brown to magenta; fastigium often reddish or brownish; lateral postocular bands yellowish; dorsomedial line frequently absent, if present very narrow, whitish, almost entirely restricted to the abdomen; dorsal shades brown, grey or magenta covering almost entirely the dorsal portion of thorax and abdomen, generally less evident in pronotum; lateral shades generally absent or restricted to head; lateral bands of blotches not evident; ventral bans of pronotum generally absent, instead ventral borders of pronotum lightly-coloured; mesonotum partially or entirely black; lateral blotches of 1 st abdominal segment if present, whitish or yellowish; in most cases hind femora uniformly coloured with knees laterally brown, dorsally reddish; hind tibia sometimes brown or yellowish.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC193B772CBED403FE3AA95F.taxon	diagnosis	Diagnosis. Externally this species resembles other species with adjacent distribution, such as S. purpurascens, S. rugosum, and S. infernalis sp. n., as well as other more geographically distant species such as S. minimum and S. zapotecum sp. n. Generally, S. tarascum sp. n. differs from the first three species by its longer antennae, more elongated head (in both sexes) and dorsal ovipositor valves. The male genitalia of S. tarascum sp. n. more closely resemble those of S. purpurascens and S. zapotecum sp. n., which also have an apical spine in the aedeagus. However, S. tarascum sp. n., differs from these species by the following combination of male genital characters: lateral borders of ramus convergent slightly rounded, inflections of supraramus moderately developed and apical spine of aedeagus slender, slightly bent and longer than in any other species in the genus.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC193B772CBED403FE3AA95F.taxon	distribution	Distribution. This species is restricted north of the central portion of the Balsas River Basin in Michoacan, Mexico (Fig. 7 A), in elevations ranging from 1394 to 1759 m a. s. l.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC193B772CBED403FE3AA95F.taxon	materials_examined	Material examined. Holotype m (Fig. 9 I) from Mexico: Michoacan, Las Cañas sobre Carr. 120, 19.08443296 ° N, - 101.768692 ° W, 1394 m a. s. l., X- 8 - 2015 (Sanabria-Urbán S. # M 011 - 11 ARO); measurements: BS = 22.19 mm, FL = 1.04 mm, PL = 4.31 mm, HF = 10.60 mm. Paratypes from Mexico: Michoacan: 9 m, 10 f same data as holotype; 3 m, 1 f, Rancho el Mirador, 19.20727001 ° N, - 101.735125 ° W, 1759 m a. s. l., X- 8 - 2015 (Sanabria- Urbán S. # M 012). The holotype was deposited at IBUNAM and the paratypes were deposited at the IBUNAM and TAMUIC. Additional material: 5 m, 2 f, from the same locality as holotype but different date; and 3 m, 1 f, from two additional localities (see Appendix Table 5). Taxonomic discussion. This species is closely related morphologically to S. purpurascens and genetically to S. rugosum. Nevertheless, its unique combination of morphologic traits (both in external and male genitalia structures), monophyly and geographic isolation supports its recognition as a separate species within the genus. Previously, we identified specimens of S. tarascum sp. n. as, Sphenarium sp. n. 5 (Sanabria-Urbán et al. 2015). For other studies in the group this species was unknown.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC193B772CBED403FE3AA95F.taxon	etymology	Etymology. Named in honour of the Tarascos, an ancient Native American people still living in the area where this species was found.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC183B482CBED4CAFE1BA80B.taxon	description	Description. External morphology (Figs. 8 C, D; 9 K, 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 (12 D-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 ½ 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 1 st 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.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC183B482CBED4CAFE1BA80B.taxon	diagnosis	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.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC183B482CBED4CAFE1BA80B.taxon	distribution	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.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC183B482CBED4CAFE1BA80B.taxon	materials_examined	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-Urbán 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.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC273B492CBED536FB1BA85E.taxon	description	Description. External morphology (Figs. 8 E, F; 9 M, N): total body length total body length ranging from 23.47 to 38.88 mm in females and 23.12 to 36.15 mm in males; antennae filiform; notably shorter in females or slightly longer than head and pronotum together in males; head subtriangular-elongated nearly as wide as long in females or moderately longer in males wide, 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 notably developed posteriorly; dorsal ovipositor valves rounded or lanceolate moderately elongated towards the apex. Male genitalia: bridge of epiphallus shorter or as long as the length of lateral plates (Fig. 12 G-I). Ectophallus in dorsal view (Fig. 12 G-II) large with lateral borders of ramus parallel; basal emargination of cingulum notably reduced; interspace between apodemal plates notably open. Ectophallus in posterior view (Fig. 12 H) without a conspicuous sclerotized hollow in the sheath; inflections of supraramus reduced; ramus of cingulum distinctly developed ventrally; valves of cingulum small with unique form, centred at the middle of the sheath. In lateral view of ectophallus valves of cingulum slightly developed posteriorly (Fig. 12 I). Endophallus in lateral view (Fig. 12 G- III) with elongated pseudoarch, loosely joined to the valves of cingulum; aedeagal valves very slender and long, with smooth ventral margins and moderately rounded in the apex without apical spine; aedeagal valves and sclerites together about 2 ½ to 3 fold the length of dorsal inflections of endophallic apodemes. Colouration. Ground colours vary from green to brown. Body uniformly coloured or with the following colour traits: antennae generally black or dark brown; fastigium frequently reddish; lateral postocular bands frequently present, wide and yellow; dorsomedial line frequently absent, if present very narrow and yellowish; dorsal shades generally absent; lateral shades frequently absent, if present narrow and black, principally evident in head, metanotum, 1 st abdominal segment and apex of 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 1 st abdominal segment frequently absent; hind femora mostly uniformly coloured, sometimes with lower medial area yellowish and knees laterally black, dorsally reddish; hind tibia generally black.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC273B492CBED536FB1BA85E.taxon	diagnosis	Diagnosis. Externally this species closely resembles S. crypticum sp. n. and S. infernalis sp. n., which are distributed contiguously. Generally, males of S. macrophallicum differ from males of other species by its rounded subgenital plate more notably developed posteriorly. Moreover, the male genitalia of S. macrophallicum are unique among its congeners and differ by the following combination of characters: ectophallus notably large with lateral borders of ramus of ectophallus parallel, inflections of supraramus reduced, ramus of cingulum distinctly developed ventrally, valves of cingulum small with distinct form and centred at the middle of the sheath, aedeagal valves very long and slender with smooth ventral margins and moderately rounded in the apex, and aedeagal valves and sclerites together longer than in any other species ranging from 2 ¾ to 3 fold the length of dorsal inflections of endophallic apodemes.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC273B492CBED536FB1BA85E.taxon	distribution	Distribution. This species is distributed northwest of the middle portion of the Balsas River Basin in elevations ranging from 207 to 1607 m a. s. l. in Michoacan and Guerrero, Mexico (Fig. 7 A).	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC273B492CBED536FB1BA85E.taxon	materials_examined	Material examined. Holotype m (Fig. 8 E) from Mexico: Guerrero, 11 rd. mi. NE. of Arcelia, XII- 8 - 1958, 3000 ft ± (T. J. Cohn # 360). Allotype f (Fig. 8 F) from Mexico: Guerrero, Cd. Altamirano, X- 20 - 1957. Paratypes from Mexico: Guerrero: 1 m same locality as holotype (Figs. 7 M; 10 G, H, I), 1 m, 1 f, Temisco, XI- 10 - 1928 (T. W: Bouchelle). Designation: Kevan and Boyle (1974) and location: UMMZ for all these specimens. We were able to examine both external and genital morphology of this type material. Additional material: 45 m, 43 f, from six localities. Locality information and depositories of these additional specimens examined is provided in Appendix Table 5. Taxonomic discussion. This species was partially described from eight specimens collected from three different localities in Guerrero, Mexico (Kevan 1977). After the original description, the validity of this species has remained unchanged, even in the most recent studies in the genus (Pedraza-Lara et al. 2015; Sanabria-Urbán et al. 2015). Our phylogenetic analysis recovered S. macrophallicum as a paraphyletic species and we observed relatively low levels of genetic differentiation between S. macrophallicum and other adjacent (S. crypticum sp. n., S. tarascum sp. n., S. rugosum, S. purpurascens) and distant species (S. variabile and S. zapotecum sp. n.) (Table 3). Nevertheless, the decidedly different and unique combination of male genital traits, and restricted geographic distribution of this species supports its recognition as an independent species within the genus.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC243B4C2CBED2FBFB4EAD14.taxon	description	Description. External morphology (Figs. 8 G, H; 9 O, 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 ¼ 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 1 st 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.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC243B4C2CBED2FBFB4EAD14.taxon	diagnosis	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 ¼ fold the length of dorsal inflections of endophallic.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC243B4C2CBED2FBFB4EAD14.taxon	distribution	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).	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC243B4C2CBED2FBFB4EAD14.taxon	materials_examined	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 (L 76); 2 m from Orizaba (L 292), 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-Urbán 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.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC233B4E2CBED000FE70AB2B.taxon	description	Description. External morphology (Fig. 9 Q, R): body size ranging from 25.74 to 38.48 mm in females and from 25.94 to 38.13 mm 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 1 st 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.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC233B4E2CBED000FE70AB2B.taxon	diagnosis	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.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC233B4E2CBED000FE70AB2B.taxon	distribution	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).	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC233B4E2CBED000FE70AB2B.taxon	materials_examined	Material examined. Holotype m (Fig. 9 Q) from Mexico: Jalisco, 5 mi. NE. Tecalitlan, 19.540895 ° N, - 103.3147204 ° W, 4250 ft, 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, 1 mi S Pihuamo, 869 m a. s. l., XI- 26 - 1958 (T. J. Conh # 314); 2 m, 2 f, 6 mi NE Tecalitlan, 4250 ft, 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 # P 78); 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. # P 79 [L 58 MS 1]). Colima: 1 m, 1 f, 9 mi E Colima, 1620 ft, XI- 22 - 1958 (T. J. Cohn # 328); 1 m, 6 mi E Colima, 1250 ft, 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. # M 58). Michoacan: 1 m, 1 f, 15 mi W Apatzingan (2 mi E. Santa Ana), 1600 ft, XII- 2 - 1958 (T. J. Cohn # 340); 1 m, 23 rd mi WSW Ario de Rosales, 4200 ft, XII- 4 - 1958 (T. J. Cohn # 348); 2 m, 1 f, 16.7 mi W Apatzingan, 1100 ft, IX- 26 - 1959 (Cantral & Cohn # 179); 1 m, 11 mi S Uruapan on 37, IX- 8 - 1981 (Otte, Azuma, Newlin # 57); 1 m, 26 rd mi NE Arteaga on Hwy. 37, 3 rd. mi. SW Rancho Nuevo, 2060 ft, 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. # P 43 [L 57 MS 1]); 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. # P 75). 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-Urbán et al. 2015). For other studies in the genus this new species was unknown.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC233B4E2CBED000FE70AB2B.taxon	etymology	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.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC213B402CBED603FEA0AFDF.taxon	description	Description. External morphology (Figs. 14 A, B, C; 9 S, T; 15 A, 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 ¾ (morphotype 2; Fig. 13 G-III) to 1 ½ (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; 15 B, 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 1 st 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.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC213B402CBED603FEA0AFDF.taxon	diagnosis	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.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC213B402CBED603FEA0AFDF.taxon	distribution	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).	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC213B402CBED603FEA0AFDF.taxon	materials_examined	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. Márquez (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-Urbán 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 (L 414, L 435, L 440, and L 449; 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 (CO 1 P-distances <3 %; Table 3) and close phylogenetic relationships (Fig. 11) between S. rugosum and the putative new taxa Sphenarium sp. Gro 1 + Gro 8 and Sphenarium sp. n. 6 recognized in Pedraza-Lara et al. (2015) and Sanabria-Urbán et al. (2015) respectively. Therefore, we consider that these taxa represent part of the same species, S. rugosum (morphotype 2 and 3, respectively).	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC2F3B442CBED377FAD1AD14.taxon	description	Description. External morphology (Fig. 15 E, F): total body length ranging from 32.22 to 35.92 mm in females and from 19.04 to 34.35 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 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 or lanceolate, moderately elongated towards the apex. Male genitalia: bridge of epiphallus as long or slightly longer than the length of lateral plates (Fig. 16 A-I). Ectophallus in dorsal view (Fig. 16 A-II) with lateral borders of ramus convergent; basal emargination of cingulum moderately developed; interspace between apodemal plates moderately open. Ectophallus in posterior view (Fig. 16 B) 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, slightly developed posteriorly (Fig. 16 C). Endophallus in lateral view (Fig. 16 A-III) with elongated pseudoarch, 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 ranging from 1 ¾ to 2 fold the length of dorsal inflections of endophallic apodemes. Colouration. Ground colours vary from green to brown. Body uniformly coloured or with the following colour traits: antennae frequently black or dark brown; fastigium frequently reddish; lateral postocular bands frequently present, wide and yellow; dorsomedial line frequently absent, if present very narrow and yellowish; dorsal black shades frequently absent, if present restricted principally to apex of the abdomen; lateral shades often present, narrow, black or dark brown; 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 1 st abdominal segment frequently absent; generally hind femora uniformly coloured with knees laterally black, dorsally reddish; hind tibia often black.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC2F3B442CBED377FAD1AD14.taxon	diagnosis	Diagnosis. Externally this species closely resembles S. macrophallicum, S. rugosum, and S. infernalis sp. n. In most cases, males of S. crypticum sp. n. differ from males of S. macrophallicum by their rounded, moderately developed posteriorly subgenital plate. Sphenarium crypticum sp. n. differs from S. rugosum by lacking the dorsomedial line and the yellow lateral bands of blotches; whereas the former species is almost indistinguishable from S. infernalis sp. n. externally. Nevertheless, S. crypticum sp. n. conspicuously differs from all Sphenarium species by the following combination of male genitalia traits: lateral borders of ramus of ectophallus convergent, inflections of supraramus moderately developed, and aedeagal valves and sclerites long, moderately rounded in the apex without apical spine.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC2F3B442CBED377FAD1AD14.taxon	distribution	Distribution. This species is apparently restricted to the western portion of the inner slope of the Sierra Madre del Sur in Guerrero, Mexico (Fig. 7 A), in elevations ranging approximately from 395 to 1662 m a. s. l.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC2F3B442CBED377FAD1AD14.taxon	materials_examined	Material examined. Holotype m (Fig. 15 E; 16 A, B, C) from Mexico: Guerrero, El Pinzan Morado Carr. 134, 18.114954 ° N, - 100.945988 ° W, 730 m a. s. l., X- 13 - 2012 (Sanabria-Urbán S. # P 66 L 47 - MS 1); measurements: BS = 34.04 mm, FL = 1.32 mm, PL = 7.25 mm, HF = 15.66 mm). Paratypes from Mexico: Guerrero: 2 m, 5 f, same locality as holotype; 1 m, 1 f, Vallecitos Carr. 134, 18.039971 ° N, - 101.046337 ° W, 916 m a. s. l., X- 13 - 2012 (Sanabria-Urban S. # P 67); 2 m, Coyuca de Catalan Microondas El Nopal, 18.30111 ° N, - 100.798333 ° W, 395 m, XI- 17 - 2006 (G. Ortega, L. Cervantes & C. Mayora). The holotype was deposited at IBUNAM and the paratypes were deposited at IBUNAM and TAMUIC. Additional material: 8 m, 5 f, same locality as holotype; 6 m, 1 f, same localities as paratypes (Appendix Table 5). Taxonomic discussion. S. crypticum sp. n. is mainly related genetically to S. macrophallicum. Nevertheless, these two species clearly differ each other in their male genital structures. Moreover, S. crypticum sp. n. shows a unique combination of morphologic traits, in both external and male genitalia structures, a well-supported monophyly (Fig. 2 b) and separated geographic distribution. All these lines of evidence support the recognition of S. crypticum sp. n. as a valid species. Material of S. crypticum sp. n. was identified as Sphenarium sp. n. 7 in Sanabria-Urbán et al. (2015), whereas the specimens collected within the ranges of S. crypticum sp. n., were considered as an unambiguous new species, Sphenarium sp. Gro 3, in Pedraza-Lara et al. (2015). During this study we examined specimens collected approximately 7 km apart from the Sphenarium sp. Gro 3 population (L 88; Appendix Table 5) that were undoubtedly S. crypticum sp. n. Moreover, CO 1 sequences of Sphenarium sp. Gro 3 clustered with our S. macrophallicum and S. crypticum sp. n. samples (PP Ż 0.95) (Fig. 11). Although Sphenarium sp. Gro 3 and S. crypticum sp. n. did not form a monophyletic group, we consider that these taxa should represent the same species, considering their phylogenetic and geographic proximity. For other taxonomic studies in the genus S. crypticum sp. n. was unknown.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC2F3B442CBED377FAD1AD14.taxon	etymology	Etymology. The specific name “ crypticum ” is derived from Latin and means “ concealed ”. It refers to the condition in which this species remained undiscovered and also the close similarity with other adjacent species	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC2B3B452CBED000FF5FACE1.taxon	description	Description. External morphology (Figs 14 D, E, F, G; 15 G, 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 ¾ 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 1 st 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.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC2B3B452CBED000FF5FACE1.taxon	diagnosis	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.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC2B3B452CBED000FF5FACE1.taxon	distribution	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).	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC2B3B452CBED000FF5FACE1.taxon	materials_examined	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 Márquez (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-Urbán 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.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC2A3B462CBED653FB46AA03.taxon	description	Description. External morphology (Figs. 14 H; 17 A; 15 I, J, K, L): total body length ranging from 21.38 to 25.57 mm in females and from 16.29 to 28.22 mm in males; antennae filiform, slightly shorter in females or notably 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 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 moderately longer or as long as the length of lateral plates (Fig. 16 G-I, J-I). Ectophallus in dorsal view (Fig. 16 G-II, J-II) with lateral borders of ramus convergent, slightly sinuous; basal emargination of cingulum notably developed reducing completely the interspace between the apodemal plates. Ectophallus in posterior view (Fig. 16 H, K) with a conspicuous sclerotized hollow in the sheath moderately closed or open; inflections of supraramus moderately developed laterally (morphotype 1; Fig. 16 H, arrow) or anteriorly (morphotype 2; Fig. 16 K, arrow); valves of cingulum small whit distinct finger-like form, slender (morphotype 1; Fig. 16 H) or somewhat broad (morphotype 2; Fig. 16 K); notably developed posteriorly (morphotype 1; Fig. 16 I) or not (morphotype 2; Fig. 16 L) in lateral view of ectophallus. Endophallus in lateral view (Fig. 16 G-III, 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 about ¾ or the same length as the dorsal inflections of endophallic apodemes. Colouration. Ground colours vary from green to brown. Body uniformly coloured (Fig. 15 J, L) or with the following colour traits (Fig. 15 I, K): antennae black, brown or pale orange; fastigium often brownish; lateral postocular bands frequently present, narrow, yellowish or whitish; dorsomedial line often present, narrow and whitish; dorsal shades frequently present, black or brown, covering partially the dorsal part of the body, frequently well delimited in pronotum by the lateral carinas (Fig. 15 I); lateral shades frequently present, black or dark brown, restricted to head and pronotum; lateral bands of blotches not evident; ventral bands of pronotum often present, wide and whitish; mesonotum partially or entirely black; lateral blotches of 1 st abdominal segment whitish frequently present; hind femora with lower medial area sometimes whitish with knees laterally black, dorsally brownish; hind tibia black, green or orange.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC2A3B462CBED653FB46AA03.taxon	diagnosis	Diagnosis. Externally this species is very similar to S. purpurascens. Sometimes, principally in sympatry, these species differ from each other in their colouration patterns. For instance, in S. variabile males the dark dorsal shades are more conspicuous and notably delimited by the lateral carinas of pronotum (e. g. Figs. 14 H; 15 I) than in S. purpurascens males (Fig. 9 A, C). Moreover, in S. variabile the dorsomedial line of body frequently lack the pinkish colourations that are commonly found in S. purpurascens. Nonetheless, S. variabile clearly differs from other species of Sphenarium by its unique lateral borders of ectophallus convergent, slightly sinuous and its fingerlike valves of cingulum.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC2A3B462CBED653FB46AA03.taxon	distribution	Distribution. This species is restricted to inner valleys and highlands of the Sierra Madre del Sur in Oaxaca, Mexico, in elevations ranging from 838 to 2279 m a. s. l. (Fig. 7 A, C). According to our results, the two morphotypes of this species are distributed separately northeast (morphotype 1) and southwest (morphotype 2) of the southern portion of the Sierra Madre del Sur.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC2A3B462CBED653FB46AA03.taxon	materials_examined	Material examined. Holotype m (Fig. 14 H) from Mexico: Oaxaca, 18 mi. NW. La Reforma (Km 695 onHwy. 190; 32 mi WNW. Tequisistlan), 2750 ft, IX- 14 - 1959 (I. J. Cantrall & T. J. Cohn # 116). Allotype f (Fig. 17 A) same data as holotype. Paratypes from Mexico: Oaxaca: 1 f, same locality as holotype; 1 m, 45 mi. NW. Tequisistlan (1 mi. S. El Camaron), 2000 ft, XII- 21 - 1958 (T. J. Cohn # 384) (Fig. 12 I; 13 G, H, F); 1 m, 13 mi. SE. Tlacolula, XII- 21 - 1958 (T. J. Cohn # 383); 1 m, 1.0 mi. NW. Tamazulapan at Km 388, IX- 9 - 1958 (T. J. Hubell, I. J. Cantrall & T. J. Cohn # 87); 1 m, 2 f, 2 mi. SE. Tlacolula on Hwy. 190, 5700 ft, IX- 11 - 1961 (T. J. Hubell, I. J. Cantrall & T. J. Cohn # 91); 1 m, 3 f, 7 mi. SE. El Camaron (37 mi. NW. Tequesistlan on Hwy. 190), 3500 ft, IX- 14 - 1959 (I. J. Cantrall & T. J. Cohn # 117); 1 m, Mitla, 5000 ft, VIII- 11 - 1938 (H. R. Roberts). Designation: Kevan and Boyle (1974); location: UMMZ for all these type specimens. We examined both external and genital morphology of the type material. Additional material: 89 m, 38 f, from 15 new localities (Appendix Table 5). Taxonomic discussion. This species was briefly described based on several specimens from seven different localities from Oaxaca, Mexico (Kevan 1977). In this original description the holotype, allotype and 68 paratypes of were designated. Since its original description the validity of S. variabile has remained unchanged for later studies (Pedraza-lara et al. 2015; Sanabria-Urbán et al. 2015). According to our results, S. variabile is genetically close and paraphyletic with respect to S. purpurascens and S. zapotecum sp. n. lineages. However, the differentiation between their male genital structures is greater than that observed between other congeners with greater genetic differentiation. Therefore, we also agree in considering S. variabile a valid species.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC293B5A2CBED70EFEBDAE33.taxon	description	Description. External morphology (Figs. 17 B, C, D, E, F, G, H; 18 A, B; 15 M, N, O, P): total body length ranging from 33.23 to 46.66 mm in females and from 30.99 to 41.58 mm in males. In most cases: antennae filiform to weakly ensiform, 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 rounded, moderately developed posteriorly; dorsal ovipositor valves rounded or lanceolate moderately elongated towards the apex. Male genitalia: bridge of epiphallus slightly longer than the length of lateral plates (Fig. 19 A-I, D-I). Ectophallus in dorsal view (Fig. 19 A-II, D-II) large with lateral borders of ramus convergent; basal emargination of cingulum notably or slightly developed; interspace between the apodemal plates close. Ectophallus in posterior view (Fig. 19 B, E) with a conspicuous sclerotized hollow in the sheath notably open; inflections of supraramus notably reduced; valves of cingulum claw-like rather stout (Fig. 19 A-III, D-III). Ectophallus in lateral view (Fig. 19 C, F) with valves of cingulum notably (morphotype 1, Fig. 19 C) or moderately (morphotype 2, Fig. 19 F) developed posteriorly, always evident. Endophallus in lateral view (Fig. 19 A-III, 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 as long as (morphotype 1, Fig. 19 A-III) or about ¾ (morphotype 2, Fig. 19 D-III) the length of dorsal inflections of endophallic apodemes. Colouration. Ground colours vary from olive green to brown. Body uniformly coloured (Fig. 15 N) or with the following colour traits (Fig. 15 M, O, P): antennae black, gray or dark blue; fastigium often blue or black; lateral postocular bands frequently present, narrow and yellowish; dorsomedial line frequently present, wide and yellowish; dorsal shades if present black to dark blue, generally absent in most of the range of the species except in populations in the Isthmus of Tehuantepec (Fig. 15 O, P); lateral shades often present, black or dark blue; lateral bands of blotches frequently present and reddish; ventral bands of pronotum often present, wide, whitish, rarely bluish; frequently pronotum with white lateral carinas and small stripes and dots in the posterior dorsal margin; mesonotum partially or entirely red; lateral blotches of 1 st abdominal segment frequently present and whitish; generally hind femora uniformly coloured with knees laterally black, dorsally bluish; hind tibia light brown to reddish.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC293B5A2CBED70EFEBDAE33.taxon	diagnosis	Diagnosis. This species is similar to S. histrio and S. totonacum sp. n. in both male genitalia and external morphology. Nevertheless, S. mexicanum clearly differs from other Sphenarium species by the following combinations of male genital characters: ectophallus relatively large, notably conspicuous sclerotized hollow in the sheath, inflections of supraramus notably reduced, and valves of cingulum claw-like, rather stout, notably or moderately developed posteriorly.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC293B5A2CBED70EFEBDAE33.taxon	distribution	Distribution. This species is distributed in elevations ranging approximately from 6 to 1280 m a. s. l. in the southern portion of the Golf of Mexico Cost, the outer slope of the Sierra de Oaxaca and the Isthmus of Tehuantepec in Oaxaca, Veracruz and Tabasco, Mexico (Fig. 7 B). The morphotype 1 of this species is widely spread across most of the range of the species; whereas morphotype 2 was only observed in a single population near Tehuantepec (L 14) (Fig. 7 B).	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC293B5A2CBED70EFEBDAE33.taxon	materials_examined	Material examined. S. mexicanum: lectotype f (Fig. 17 B) and paralectotype m (Fig. 17 C) from Mexico: Litoral du Mexique (M. H. Saussure); designation: Kevan (1960, unpublished results); located: Natural History Museum Geneva (NHMG), Geneva, Sweden. O. crassipes: holotype m (Fig. 17 D) from Mexico: Veracruz; designation by monotypy; location: British Museum of Natural History (BMNH), London, England. S. ictericum: lectotype m (Fig. 17 E) and paralectotype f (Fig. 17 F) from Mexico; designation: Kevan (1962, unpublished results); location: BZM. S. bolivari: lectotype m (Fig. 17 G) and paralectotype m (Fig. 17 H) from Mexico: Oaxaca, Salina Cruz, XII- 1898 (C. C. Deem); designation: (Rehn & Hebard 1912); location: ANSP. S. marginatum: lectotype m (Fig. 18 A) from Mexico: Veracruz, Orizaba, XI and paralectotype m (Fig. 18 B) from Mexico: Veracruz, Medellin (T. Heyde); designation: (Rehn & Hebard 1912); location: ANSP. S. magnum: holotype m, allotype f and paratypes (2 m, 2 f), from Mexico: Oaxaca, Santo Domingo, XII- 16 - 1 953 (L. Vázquez); designation: Márquez (1962); location: IBUNAM. For S. bolivari we were able to examine the external and male genital morphology of the paratype; whereas for the remainder taxa we could examine only their external morphology. Additional material: 212 m, 169 f, form 50 Mexican localities (Appendix Table 5). Taxonomic discussion. Saussure (1859) briefly described S. mexicanum based on an unstated number of male and female syntypes from “ Mexico Calida ”, probably Veracruz, Mexico (Saussure 1859). The validity of S. mexicanum remained unchanged until Boyle (1974) ans Kevan (1977) considered this taxon as the subspecies S. mexicanum mexicanum (Boyle 1974; Kevan 1977). Recently, Pedraza-lara et al. (2015) and Sanabria-Urbán et al. (2015) have proposed the elevation of this taxon at the species level. According to our results, this species is closely related and / or paraphyletic with respect to S. histrio and S. occidentalis sp. n. lineages. However, S. mexicanum clearly differs from these species genetically (CO 1 P-distance> 3 %) and in male genital structures. Therefore, we also agree in considering S. mexicanum a valid species.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC353B5B2CBED3E5FCC8A90B.taxon	description	Description. External morphology (Figs. 15 Q, R, S, T; 18 C, D; 20 A, 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; 21 A-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; 21 A-II, D-II, respectively) or slightly rounded (morphotypes 2 and 5; Figs. 19 J-II; 21 G-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; 21 B, H); inflections of supraramus moderately developed, with distal borders ventrally (morphotypes 1, 2, and 5; Figs. 19 H, K; 21 H, 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; 21 B, 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; 21 C, F, respectively). Endophallus in lateral view (Figs. 19 G-III, J-III; 21 A-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 ¾ (other morphotypes; Figs. 19 G-III, J-III; 21 A-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; 20 B) or with the following colour traits (Figs. 15 Q, S; 20 A, 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 1 st 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.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC353B5B2CBED3E5FCC8A90B.taxon	diagnosis	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.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC353B5B2CBED3E5FCC8A90B.taxon	distribution	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.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC353B5B2CBED3E5FCC8A90B.taxon	materials_examined	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, Márquez (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-Urbán 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 CO 1 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. Oax 9 and Sphenarium sp. Oax 2 (Pedraza-lara et al. 2015) (L 11, L 167, L 172, and L 173; Appendix Table 5). The male genitalia of these specimens were similar to those of other S. histrio individuals. Moreover, CO 1 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.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC343B5D2CBED43DFB1EAE4C.taxon	description	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. 21 K) 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 ¾ 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 1 st abdominal segment frequently present and whitish; hind femora uniformly coloured with knees laterally black laterally, dorsally bluish or brownish.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC343B5D2CBED43DFB1EAE4C.taxon	diagnosis	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.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC343B5D2CBED43DFB1EAE4C.taxon	distribution	Distribution. This species is distributed in elevations ranging approximately from 30 to 750 m 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).	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC343B5D2CBED43DFB1EAE4C.taxon	materials_examined	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-Urbán S. # P 39); 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, 31 mi 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, 3 rd mi SW Arteaga church (0.1 mi 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, 15 mi W Caleta de Campos (W playa Azul), X- 9 - 1981 (Otte # 62); 1 m, 1 f, 12 mi W Caleta de Campos (W playa Azul), X- 9 - 1981 (Otte # 63); 1 m, 1 f, 26 mi n La Mira, VIII- 9 - 1981 (Otte # 59); 1 m, 1 f, 16 - 20 km NE Rt 200 Ixtapa-Altamirano Rd., VIII- 9 - 1981 (Otte # 60); 6 M, 6 F, Las Peñitas, 17.99 0621 ° N, - 102.026924 ° W, 36 m a. s. l., IX- 23 - 2012 (Sanabria-Urbán S. # P 40 [L 7 MS 1]); 1 M, 1 f, Zicuirán Carr. 120 Km 144, 18.881308 ° N, - 101.967336 ° W, 239 m a. s. l., IX- 22 - 2012 (Sanabria-Urbán S. # P 38 [L 8 MS 1]); 1 m, 1 f, Carr. 37 D Km 236, 18.3868 ° N, - 101.89475 ° W, 197 m a. s. l., IX- 23 - 2012 (Sanabria-Urbán S. # M 055). Guerrero: 1 m, 5 mi N Acapulco, IX- 15 - 1940 (C. Bolivar & H. R. Roberts); 2 m, 1 f, 13 mi 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-Urbán S. # 16 ACA [L 9 MS 1]). 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-Urbán 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. Gro 6 and Sphenarium sp. Gro 9, 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. Gro 6 and Gro 9 (e. g. L 09, L 309, and L 307, L 315; Appendix Table 5) that were undoubtedly S. occidentalis sp. n. Moreover, CO 1 sequences of Sphenarium sp. Gro 6 of S. occidentalis sp. n. formed a well supported (PP Ż 0.95) monophyletic group (Fig. 11). Therefore, we consider that Sphenarium sp. Gro 6 and Sphenarium sp. Gro 9 probably represent additional populations of S. occidentalis sp. n.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC343B5D2CBED43DFB1EAE4C.taxon	etymology	Etymology. This species is named after its distribution in the occidental region of Mexico.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC323B512CBED3E8FE45A988.taxon	description	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 1 st abdominal segment with darker green colouration; mesonotum red; lateral blotches of 1 st abdominal segment if present whitish; generally hind femora uniformly coloured with knees laterally black, dorsally brownish; hind tibia reddish.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC323B512CBED3E8FE45A988.taxon	distribution	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).	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC323B512CBED3E8FE45A988.taxon	diagnosis	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.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC323B512CBED3E8FE45A988.taxon	materials_examined	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-Urbán S. & Jímenez-Arcos V. H. # P 72 [L 19 MS 1]); 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. 7 km SO de Tlapacoyan, 19.90252398 ° N, - 97.22993698 ° W, 751 m a. s. l., IX- 19 - 2015 (Sanabria-Urbán S. # M 010 - L 52); 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-Urbán 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.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC323B512CBED3E8FE45A988.taxon	etymology	Etymology. Named in honour of the Totonacos, an ancient Native American people still living in the area where this species was found.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC3E3B532CBED4BBFACCAEF8.taxon	description	Description. External morphology (Fig. 20 Q, R): total body length ranging from 27.91 to 35.25 mm in females and from 26.47 to 33.1 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 slightly elongated towards the apex. Male genitalia: bridge of epiphallus slightly longer than the length of lateral plates in most cases (Fig. 22 D-I). Ectophallus in dorsal view (Fig. 22 D-II) small with lateral borders of ramus convergent, straight or slightly rounded; basal emargination of cingulum notably developed closing the interspace between the apodemal plates. Ectophallus in posterior (Fig. 22 E) view with a conspicuous sclerotized hollow in the sheath notably closed; valves of cingulum small with distinct form; inflections of supraramus notably developed laterally and dorsally forming a dorsal fold; dorsal borders of inflections of supraramus very close but not fused (Fig. 22 E, arrow). Ectophallus in lateral view with valves of cingulum barely evident slightly developed posteriorly (Fig. 22 F). 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 light brown. Body uniformly coloured or with the following colour traits: antennae dark to light brown; fastigium brownish; lateral postocular bands frequently present, wide and yellowish, whitish or cyan; dorsomedial line frequently present, wide and yellowish; dorsal shades brown to dark magenta, frequently covering entirely the dorsal portion of the abdomen; lateral shades often present, dark to light brown; lateral bands of blotches absent; ventral bands of pronotum often present, wide, yellowish or cyan; pronotum sometimes with small dots in the dorsal posterior margin; mesonotum partially or entirely brownish; lateral blotches of 1 st abdominal segment frequently present and yellowish; hind femora with medial area uniformly coloured and lower marginal area with distinct blue colouration, specially in the apex of femur (Fig. 20 Q, arrow); knees of hind femora without dark colouration; hind tibia intense yellow.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC3E3B532CBED4BBFACCAEF8.taxon	diagnosis	Diagnosis. Externally this species closely resembles S. histrio and S. miztecum sp. n. In most cases, S. adelinae sp. n. differs from S. histrio by its weakly ensiform antennae and the blue colourations in the lower marginal area of the hind femora; whereas it differs from S. miztecum principally by lacking the cyan lateral band of blotches. Nevertheless, more conspicuous differences exist among the male genital structures of these species. Sphenarium adelinae sp. n. differs from these species by the following combinations of male genital traits: sclerotized hollow in the sheath notably closed, valves of cingulum small with distinct form, and inflections of supraramus notably developed laterally and dorsally with dorsal borders almost fused above the valves of cingulum.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC3E3B532CBED4BBFACCAEF8.taxon	distribution	Distribution. This species is only known from a small region in the outer slope of the Sierra Madre del Sur in Guerrero, Mexico (Fig. 7 B). The altitudinal distribution of this species ranges approximately from 420 to 1210 m a. s. l.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC3E3B532CBED4BBFACCAEF8.taxon	materials_examined	Material examined. Holotype m (Fig. 20 Q) from Mexico: Guerrero, 14 mi. S. Chilpancingo, IX- 4 - 1981 (Otte, Azuma & Newlin # 52); measurements: BS = 32.94 mm, FL = 1.38 mm, PL = 6.28 mm, HF = 15.51 mm. Paratypes from Mexico: Guerrero: 12 m, 9 f, Acahuizotla ca. de Ocotito, Carr. 95, 17.355494 ° N, - 99.480226 ° W, 1018 m a. s. l., X- 4 - 2011 (Sanabria-Urbán S. # 13 OC [L 16 MS 1]); 5 m, 4 f, Tierra Colourada Carr. 95, 17.204383 ° N, - 99.508199 ° W, 534 m a. s. l., X- 4 - 2011 (Sanabria-Urbán S. & Días de la Vega A. # 15 TC); 5 m, 5 f, Palo Blanco Carr. 95, 17.409805 ° N, - 99.466767 ° W, 1210 m a. s. l., X- 4 - 2011 (Sanabria-Urbán S. # 14 ZC [L 17 MS 1]); 1 m, Tierra Colourada, 1800 ft, IX- 16 - 1940 (C. Bolivar & H. R. Roberts); 2 m, 16 rd. mi S Chilpancingo (Km 298 on Hwy. 95), 3800 ft (I. J. Cantrall & T. J. Cohn # 147); 1 m, 2 mi S Tierra Colourada (Km 335 on Hwy. 95), 1400 ft, IX- 19 - 1959 (I. J. Cantrall & T. J. Cohn # 148). The holotype was deposited at ANSP and the paratypes were deposited at IBUNAM and TAMUIC. Additional material: 22 m, 24 f, form the same first three paratypes localities (Appendix Table 5). Taxonomic discussion. This species is closely related morphologically to S. histrio. Specimens of this new species were identified S. m. histrio in previous studies (Boyle 1974; Kevan 1977). In this study we found that S. adelinae sp. n. shows a unique combination of morphologic traits, both external and on male genitalia, differencing this new species from S. histrio. Moreover, this new species has a well-supported monophyly, shows relatively high levels of interspecific genetic differentiation (Table 3). We consider that all these lines of provide evidence for the recognition of S. adelinae sp. n. as a valid species. Previously, we identified specimens of this new species as Sphenarium sp. n. 2 (Sanabria-Urbán et al., 2015). Pedraza-Lara et al. (2015) recognised a putative new species, Sphenarium sp. Gro 7, externally similar and geographically close to S. adelinae sp. n. Indeed, we examined several specimens collected less than 9 km apart from the Sphenarium sp. Gro 7 locality (L 86, L 100, and L 103; Appendix Table 5), which were invariably S. adelinae sp. n. Nevertheless, the CO 1 sequences of Sphenarium sp. Gro 7 are different but closely and strongly related (PP Ż 0.95) to the S. adelinae sp. n. / S. miztecum sp. n group (Fig. 11). Considering their geographical and phylogenetic proximity, both Sphenarium sp. Gro 7 and S. adelinae sp. n. probably represent the same species. The paraphyletic relationships between these taxa suggest a greater genetic diversity within S. adelinae sp. n. than the one we recognised during our genetic analysis. Another possibility is that the genetic differentiation observed between CO 1 sequences of Sphenarium sp. Gro 7 and S. adelinae sp. n. is due to the different methodology used to obtain the sequences. We obtain mitochondrial sequences following a long-PCR amplification protocol in order to avoid coamplification nuclear mitochondrial pseudogenes; whereas Pedraza-Lara et al. (2015) followed standard CO 1 barcoding protocols, which usually results in coamplification of multiple paralogous numts haplotypes of different divergences, leading to the overestimation of genetic diversity (Song et al., 2008).	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC3E3B532CBED4BBFACCAEF8.taxon	etymology	Etymology. This species is dedicated to the memory of Adelina Cedillo Franco, who always supported and encouraged the senior author in his biological studies. The specific name is a female noun in the genitive case.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC3C3B542CBED02BFE45AE84.taxon	description	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 1 st 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.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC3C3B542CBED02BFE45AE84.taxon	diagnosis	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.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC3C3B542CBED02BFE45AE84.taxon	distribution	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.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC3C3B542CBED02BFE45AE84.taxon	materials_examined	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-Urbán S. # 18 CPI [L 18 MS 1]). 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- Urbán et al. 2015). For other studies in the genus, this new species was unknown.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
A12C6E25AC3C3B542CBED02BFE45AE84.taxon	etymology	Etymology. Named in honour of the Mixtecos, an ancient Native American people still living in the area where this species was found.	en	Sanabria-Urbán, Salomón, Song, Hojun, Oyama, Ken, González-Rodríguez, Antonio, Castillo, Raúl Cueva Del (2017): Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae). Zootaxa 4274 (1): 1-86, DOI: 10.5281/zenodo.804182
