Sphenarium adelinae 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 : 58-60

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

(http://lsid.speciesfile.org /urn:lsid: Orthoptera .speciesfile.org:TaxonName:495100)

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 1st 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.

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.

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.

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. # 13OC [L16 MS1]) ; 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. # 15TC); 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. # 14ZC [L17 MS1]); 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. Gro7, 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. Gro7 locality (L86, L100, and L103; Appendix Table 5), which were invariably S. adelinae sp.n. Nevertheless, the CO1 sequences of Sphenarium sp. Gro7 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. Gro7 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 CO1 sequences of Sphenarium sp. Gro7 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 CO1 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).

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.

Figures

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 20. External morphology of S. histrio: morphotype 2 m (A and C) and f (B and D), morphotype 3 m (E) and f (F), morphotype 4 m (G) and f (H), and morphotype 5 m (I) and f (J); S. occidentalis sp. n. holotype m (K) and paratypes m # 2 (M), f # 3 (L) and f # 4 (N); S. totonacum sp. n. holotype m (O) and paratype f # 1 (P); S. adelinae sp. n. holotype m (Q) and paratype f # 1 (R); S. miztecum sp. n. holotype m (S) and paratype f # 1 (T) (Scale bars = 1 cm).

FIGURE 22. Male genital structures of S. totonacum sp. n. holotype (A – C), S. adelinae sp. n. holotype (D – F); and S. miztecum sp. n. holotype (G – I). 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).