Scaria Bolívar, 1887

Cadena-Castañeda, Oscar J., Mendes, Diego Matheus De Mello, Silva, Daniela Santos Martins, Granda, Juan Manuel Cardona, García, Alexander García & Tumbrinck, Josef, 2019, Systematics and biogeography of the genus Scaria Bolívar, 1887 (Orthoptera: Tetrigidae: Batrachideinae), Zootaxa 4675 (1), pp. 1-65 : 13-15

Genus Scaria Bolívar, 1887

Type species: Scaria hamata Bolívar, 1887 , by subsequent designation ( Kirby, 1910).

Diagnosis. Medium-sized and slender ( Figs. 8A , 19A , 20A , 24A , 26A ). Eyes strongly globose, frontal costa sulcate ( Figs. 8B , 19B , 20B , 24B , 26B ), antennae elongated and filiform. Pronotum with the dorsum flat and with the anterior edge of pronotum curved forward, sharp and hook-shaped, always somewhat ascendant, usually markedly so, not extending beyond the fastigium of vertex ( Figs. 12C , 19C , 24C , 27C ). No brachypronotal form is known. Humeral angles very obtuse, known species fully winged, elytra black, oblong, with a pale mark or without it ( Figs. 12C , 19C , 24C , 27C ). Mid femora usually with an internal, dorso-apical spine. Female subgenital plate variable emarginated, with or without a median lobe ( Figs. 9E , 11F , 18F , 21E , 25F , 28E ). Penultimate male sternite projecting ventro-apically in various degrees of prominence ( Figs. 26F , 27F ).

Description. Body slightly granulate, slim and elongated ( Figs. 8A , 19A , 20A , 24A , 26A ). Head: frontal costa bifurcation located at the top of the vertex, between the compound eyes; scutellum narrow; lower margin of the antennal grooves located between of the lower margin of the compound eyes; eyes with a rounded dorsal base, globose or subglobose and exerted above the pronotum ( Figs. 8B , 19B , 20B , 24B , 26B ); antennae 20–22 segmented, filiform with elongate articles; vertex variable, usually flat, sometimes very weakly domed or concave, median carina absent. Facial outline essentially flat, occasionally indented bellow the frontal costa. Fastigium of vertex not reaching the most anterior margin of the eyes, usually well behind this point, truncate from above; fastigio-fascial angle usually rectangular-rounded, sometimes obtuse or subobtuse-angulate; palpi elongated and flattened, always whitish. Pronotum: slender, surpassing the tip of hind femora (macropronotal), pronotal disc flat; anterior margin produced in a delicate to stout, acute, and elongate spine which is ascendant behind the head and curves over the vertex termination behind the fastigium; humeral angles rounded; lateral lobe sub-triangulated; stripe of the upper half of the lateral margin of pronotum strip of upper half of pronotum usually brownish, covering the lateral lobes of the pronotum ( Figs. 8D , 10B , 12D , 15B , 17B , 19B , 20D , 24D ). Wings: tegmina black, spotless, or with spots of various shapes, usually with a circular or oblong spot, that can be accompanied with additional strips, or not. ( Figs. 8C , 12C , 19C , 24C , 27C ); hind wings surpassing the abdomen tip. Legs: fore and mid legs elongate, fore femur obsolete to weakly sulcate above, internal dorso-apical spine on fore femur well developed; mid femora with a moderate to well-developed internal, dorso-apical spine, conspicuously sulcate above for the entire length; antegenicular tooth conspicuous; chevrons ridges in the external surface well visible. Terminalia: penultimate male sternite ventro-apically usually emarginated or with a small to prominent medio apical process; male subgenital plate inflated or slender with different angles curvature of its apex ( Figs. 3 A–G). Ovipositor valves normal ( Figs. 21F , 25G , 28F ), only in S. laeta , short and broad ( Fig. 9F ); female subgenital plate emarginated along ventro apical edge ( Figs. 21E , 25F , 28E ).

Comments: The genus Scaria was erected by Bolívar in 1877, for two species S. hamata ( De Geer, 1773) and S. lineata Bolívar, 1887 . Later, Hancock (1907 a, b, 1909) adds more species to the genus ( S. brevis , S. fasciata , S. ferruginea and S. producta ). De Geer (1773) Giglio-Tos (1898), Bruner (1920) and Günther (1940) described a species each ( S. hamata , S. maculata , S. boliviana and S. laeta respectively); recently Cadena-Castañeda & Cardona (2015) synonymized S. laeta under S. ferruginea .

Comparative notes. This genus can be easily distinguished from the Neotropical genera Rehnidium and some macropronotal species of Batrachidea . It can be separated from Rehnidium on account of body size ( Rehnidium is of a smaller size) ( Fig. 1B ), frontal costa is also more pronounced in Rehnidium than in Scaria . Batrachidea its more robust ( Figs. 1A ) than Scaria , and with a weakly decurved pronotal spine. Superficially, Scaria is, in body appearance, similar to Palaioscaria ; they can be separated by the lance-shaped tegmina, longer legs and subgenital plate of Palaioscaria ( Grant, 1966) .

As additional characters to separate these three closely related genera, it can be observed that for the species of Scaria , no brachipronotal population has even been found, (except for S. brevis which is placed below in Batrachidea ), all are macropronotals ( Figs. 1D ), and the few known individuals of Rehnidium , are all brachypronotals ( Fig. 1B ). Likewise, in Batrachidea only two species have been found to have macropronotal individuals: B. flavonotata (seemingly atypical individuals, represented by a small percentage of the specimens examined in several locations in Colombia (Cadena-Castañeda in prep.)) and B. mucronata , with close to half the specimens being macropronotals according to the material reviewed by Grant (1956a).

Besides, tegmina in Scaria are black, accompanied by stripes or spots that separate the species ( Figs. 2 J–M), whereas in Batrachidea and Rehnidium tegmina is brown, like in the rest of the genera of the subfamily. The body is slim and elongated in Scaria , contrasting with the other two genera, which are from stout to squat even in the macropronotal forms of Batrachidea . Hind femora in Batrachidea and Rehnidium is not sulcate, unlike in Scaria . In any case, male terminalia in Scaria are quite diverse, but they differ from the other genera in which the penultimate sternite is prolonged from slightly to noticeably ( Figs. 21F , 25G , 28F ), while the other genera lack this character, the penultimate sternite being rounded, with any protrusion whatsoever in the remaining Batrachideinae .

Notes on literature and comments on some collections: Harold J. Grant Jr. (1921–1966) was a orthopterologist who made remarkable contributions to various genera of the order Orthoptera , including members of Tetrigidae (on this family he focused his efforts), Eumastacidae , Romaleidae , Acrididae and Tettigoniidae . Unfortunately, he suffered an untimely death, on 27 February, 1966, resulting from a swimming accident off the coast of Trinidad ( Phillips, 1966).

Many of Grant’s contributions are focused on the subfamily Batrachideinae , covering revisions of several genera of this subfamily ( Grant, 1955a, b, 1956a, b, 1966; Rehn & Grant, 1958, 1961). His doctoral thesis was a revision of the subfamily Batrachideinae (1962) , where his previous works on this group are condensed and developed in which would eventually become his future contributions on this subfamily of pygmy grasshoppers. By what can be inferred from what arguably was his magnum opus, several revisions stemming from his doctoral thesis were left unpublished.

In Scaria ’s case, in Grant’s thesis (1962) no new species were reported, but a revision of the type specimens was performed. He established the synonym of S. maculata under S. hamata and S. ferruginea under S. lineata . He also redescribed all the species and provided a key for their identification. In this paper, the reference base was Grant’s thesis, and we only used the data as far as we could agree with it. We had, however, several conclusions diverging from Grant’s, which we could otherwise corroborate, in particular distributional data about the species, which are duly recorded in this paper.

Regrettably, many of the specimens of the family Tetrigidae and others groups described by Lawrence Bruner and deposited in the Carnegie Museum of Natural History, Pittsburgh, Pennsylvania, USA (CMNH), were exchanged with the Academy of Natural Sciences, Philadelphia (ANSP) and some type specimens were lost along the way ( Cadena-Castañeda & Cortés-Torres, 2013). So part of the data published by this author could not be tracked or confirmed to be included in our distribution maps.

Distribution: This genus is distributed in Amazonia, and only S. fasciata is found elsewhere, namely along the great forests of the Pacific region in northwest South America, in what is now called the Tumbes-Chocó-Magdalena ecoregion (Mittermeier et. al. 2005) (formerly known as the Chocó-Darién-Western Ecuador Hotspot). This species ranges all the way north to southern Nicaragua ( Map 1 ).

Figures

FIGURE 1. Characters used in phylogeny. A. Batrachidea flavonotata Bolívar, 1887. B. Rehnidium omnivagum Grant, 1956. C. Tettigidea parradae Cadena-Castañeda & Cardona, 2015. D. Scaria rafaeli sp. nov. Habitus in lateral view to all specimens. Numbers indicate characters and states used in the cladistics analysis.

MAP 1. Distributional records of Scaria species.

FIGURE 2. Characters used in phylogeny. A. Scaria ferruginea Hancock, 1909 (habitus in dorsal view). B. Apex of pronotum in dorsal view of B. flavonotata. C–F. Head and pronotum in lateral view: C. S. ferruginea. D. T. parradae. E. S. laeta. F. S. producta. G and H. Fore and mid legs respectively of S. rafaeli. I. Hind femur of S. producta. J–M. Tegmina: J. S. laeta. K. S. granti sp. nov. L. S. jonasi sp. nov. M. S. ferruginea. Numbers indicate characters and states used in the cladistics analysis.

FIGURE 3. Characters used in phylogeny. A–G, J. Male terminalia and H, I, K, L. Female terminalia: A, D, H. S. laeta. B, E. S. granti sp. nov. C, F, I. S. ferruginea. G, J, K. S. hamata. L. S. jonasi sp. nov. Numbers indicate characters and states used in the cladistics analysis.

FIGURE 8. Scaria laeta Günther, 1940 stat. ressurr. (male). A. Habitus in lateral view. B. Frons. C. Head and pronotum in lateral view. D. Habitus in dorsal view. E–G. Male terminalia. E. Terminalia in latero-dorsal view. F. ventral view and G. lateral view respectively.

FIGURE 9. Scaria laeta Günther, 1940 stat. ressurr. (female). A. Habitus in lateral view. B. Frons. C. Head and pronotum in lateral view. D. Habitus in dorsal view. E. Terminalia in ventral view. F. Terminalia in lateral view.

FIGURE 10. Scaria maculata Giglio-Tos, 1898 (male). A. Habitus in lateral view. B. Habitus in dorsal view C. Frons. D. Head and pronotum in dorsal view. E. Terminalia in lateral view. (Photos: Filippo Maria Buzzetti).

FIGURE 11. Scaria producta Hancock, 1907 (female) A. Habitus in lateral view. B. Frons. C. Head and pronotum in lateral view. D. Habitus in dorsal view. E. Terminalia in lateral view. F. Terminalia in ventral view. (Photos: Gustavo Tavares).

FIGURE 12. Scaria producta Hancock, 1907 (male). A. Habitus in lateral view. B. Frons. C. Head and pronotum in lateral view. D. Habitus in dorsal view. E. Terminalia in lateral view. F. Terminalia in ventral view. (Photos: Gustavo Tavares).

FIGURE 15. Scaria lineata Bolívar, 1887 (male). A. Habitus in lateral view and. B. dorsal view respectively. (Photos: Mercedes París, MNCN).

FIGURE 17. Scaria verutum (Grant, 1956) comb. nov. A. Habitus in lateral view and. B. dorsal view respectively.

FIGURE 18. Scaria ferruginea Hancock, 1909 (female). A. Habitus in lateral view. B. Frons. C. Head and pronotum in lateral view. D. Habitus in dorsal view. E. Terminalia in dorso-lateral view. F. Terminalia in ventral view. G. Terminalia in lateral view.

FIGURE 19. Scaria ferruginea Hancock, 1909 (male). A. Habitus in lateral view. B. Frons. C. Head and pronotum in lateral view. D. Habitus in dorsal view. E–G. Male terminalia. E. Terminalia in dorsal view. F. ventral view and G. lateral view respectively.

FIGURE 20. Scaria hamata (De Geer, 1773) (male). A. Habitus in lateral view. B. Frons. C. Head and pronotum in lateral view. D. Habitus in dorsal view. E–G. Male terminalia. E. Terminalia in latero-dorsal view. F. ventral view and G. lateral view respectively.

FIGURE 21. Scaria hamata (De Geer, 1773) (female). A. Habitus in lateral view. B. Frons. C. Head and pronotum in lateral view. D. Habitus in dorsal view. E. Terminalia in ventral view. F. Terminalia in lateral view.

FIGURE 24. Scaria rafaeli sp. nov. (male). A. Habitus in lateral view. B. Frons. C. Head and pronotum in lateral view. D. Habitus in dorsal view. E–G. Male terminalia. E. Terminalia in dorsal view. F. ventral view and G. lateral view respectively.

FIGURE 25. Scaria rafaeli sp. nov. (female). A. Habitus in lateral view. B. Frons. C. Head and pronotum in lateral view. D. Habitus in dorsal view. E. Terminalia in dorsal view. F. Terminalia in ventral view. G. Terminalia in lateral view.

FIGURE 26. Scaria granti sp. nov. (male). A. Habitus in lateral view. B. Frons. C. Head and pronotum in lateral view. D. Habitus in dorsal view. E–G. Male terminalia. E. Terminalia in dorsal view. F. ventral view and G. lateral view respectively.

FIGURE 27. Scaria jonasi sp. nov. (male). A. Habitus in lateral view. B. Frons. C. Head and pronotum in lateral view. D. Habitus in dorsal view. E–G. Male terminalia. E. Terminalia in latero-dorsal view. F. ventral view and G. lateral view respectively.

FIGURE 28. Scaria jonasi sp. nov. (female). A. Habitus in lateral view. B. Frons. C. Head and pronotum in lateral view. D. Habitus in dorsal view. E. Terminalia in ventral view. F. Terminalia in lateral view.