Sphenarium crypticum Sanabria-Urbán, Song & Cueva

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), pp. 1-86 : 41-45

Sphenarium crypticum Sanabria-Urbán, Song & Cueva del Castillo sp.n.

(http://lsid.speciesfile.org/urn:lsid: Orthoptera .speciesfile.org:TaxonName:495097)

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 1¾ 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 1st abdominal segment frequently absent; generally hind femora uniformly coloured with knees laterally black, dorsally reddish; hind tibia often black.

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.

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.

Material examined. Holotype m ( Fig. 15 E; 16A, 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. #P66 L47-MS1); 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. #P67); 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. Gro3, in Pedraza-Lara et al. (2015). During this study we examined specimens collected approximately 7 km apart from the Sphenarium sp. Gro3 population (L88; Appendix Table 5) that were undoubtedly S. crypticum sp.n. Moreover, CO1 sequences of Sphenarium sp. Gro3 clustered with our S. macrophallicum and S. crypticum sp.n. samples (PP Ż 0.95) ( Fig. 11 ). Although Sphenarium sp. Gro3 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.

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

Figures

FIGURE 2. Bayesian phylogeny of Sphenarium based on a concatenated analysis of 2527 nucleotide positions from five loci and 145 terminals (129 ingroup and 16 outgroup terminals). Higher-level phylogenetic relationships are shown on left-top box and ingroup relationships are magnified outside. Voucher and locality identifier numbers of the analysed specimens are indicated in bold characters in terminals names, except for those cases in which genetic information was retrieved from the GenBank. Different branch colours highlight the three mayor clades within the genus (brown, Clade 1; red, Clade 2; and purple, Clade 3). Different colours of the Sphenarium terminals represent the 17 taxa identified during the morphologic analysis. Black circles behind the nodes indicate PP values ± 95 %. For some important nodes we also showed the PP values in bold numbers. Black bottom bars in all cases are equal to 0.03 substitutions per site.

FIGURE 7. Geographic distribution of Sphenarium species. Numbers within parenthesis in front of taxa names indicate the number of identified morphotype within the species. White surrounded areas and upper case abbreviations denote the Mexican biogeographic provinces. AC, Altos de Chiapas; AL, Altiplano Sur; BRB, Balsas River Basin; GMC, Gulf of Mexico Coast; MVB, Mexican Volcanic Belt; PC, Pacific Coast; SMOC, Sierra Madre Occidental; SMOR, Sierra Madre Oriental; SMS, Sierra Madre Sur; SO, Soconusco; and SOX, Sierra de Oaxaca.

FIGURE 11. Bayesian phylogeny of Sphenarium species based on CO 1 sequences of Pedraza-Lara et al. (2015) and this study. The analysis was conducted using the substitution model estimated previously and under the same conditions above specified for the concatenated phylogenetic analysis in methods section. Terminals of different colours represent the 17 recognized species in this study. Black dots behind the nodes indicate PP values greater than 95 %. Black bottom bars are equal to 0.04 substitutions per site.

FIGURE 15. External morphology of S. rugosum: morphotype 2 m (A) and f (B), morphotype 3 m (C) and f (D); S. crypticum sp. n. holotype m (E) and paratype f # 45 (F); S. borrei m (G) and f (H); S. variabile: morphotype 1 paratype m # 134 (I) and f (J), and morphotype 2 m (K) and f (L); S. mexicanum: morphotype 1 m (M) and f (N), and morphotype 2 m (O) and f (P); S. histrio: morphotype 1 m (Q and S) and f (R and T) (Scale bars = 1 cm).

FIGURE 16. Male genital structures of S. crypticum sp. n. holotype (A – C); S. borrei (D – F); S. variabile: morphotype 1 paratype # 234 (G – I) and morphotype 2 (J – l). For all taxa left squares show epiphallus (I) and ectophallus (II) in dorsal view, and endophallus in lateral view (III); meddle squares show ectophallus in posterior view; and right squares show a close up of ectophallus in lateral view (Scale bars = 1 mm).