Dellia multicolor Carl, 1916

Yong, Sheyla, 2018, Revision of the genus Dellia Stål, 1878 (Orthoptera: Acrididae). Part 1. Two redescriptions and seven new species from Cuba, Ecologica Montenegrina 16, pp. 58-107 : 63-71

Dellia multicolor Carl, 1916 status resurrected

Figs. 4b – 6 , 16– 24 , 25e–f , 26f . Table I

Dellia multicolor Carl, 1916: 512–513 . Rehn & Hebard, 1938: 224.

Dellia insulana : Amédégnato et al., 1995: 694 (misidentification).

Types (corrected). CUBA : somewhere between Havana City (Havana Province) and Limonar Town (Matanzas Province); January 29 th through February 22 th, 1855; H. de Saussure; 1♀ lectotype, 1♀ paralectotype ( MHNG, dry pinned). Notes. The lectotype bears three labels handwritten in black ink that textually read: 1) " Cuba Mr H de S." [aged white card]; 2) " Dellia multicolor Carl " [green card]; 3) " Dellia multicolor Carl. Hololectotypus ♀ C S Carbonell - 1966" [red card, handwritten by Carlos S. Carbonell]. The precise data given above were not stated in the original description nor labels, but gathered from a reliable supplementary source (see detailed discussion below, in Remarks section) .

Additional material examined. CUBA : Mayabeque Province: Santa Cruz del Norte Municipality: Boca de Canasí (23°08'36"N - 81°46'22"W, 20 m a.s.l.); transition from sea-grape to semideciduous forest, on Zanthoxylum fagara bush; 26/June/2014; S. Yong, I. Rodríguez, L. J. Forcelledo, L. Valdés, E. Fonseca; 2♂♂ ( SY, dry pinned).

Diagnosis (emended). Size medium to moderately large for the genus (18–23 mm). 1) Coloration predominantly light olivaceous green, with two irregular yellow lateral stripes on each side of the body, dorsum yellowish with a broad green median longitudinal stripe, epimera I–II each with a conspicuous bright yellow spot, tergites without conspicuous lateral spots, sternites red, hind femur pale olivaceous green, conspicuously darker on transversal ridges and upper and lower margins, hind knees with upper half blackish and lower half dark bluish, tarsomeres whitish. 2) Tegmina very short and digitiform, reaching anterior margin of abdominal segment I. 3) Supra-anal plate relatively large, subpentagonal, wider than long, and sparsely setose; anterior margin shallowly convex and medially moderately lobed, lateral margins straight and essentially parallel, posterior margin narrowly paraboloid; median sulcus complete, moderately deep and narrow, flanked by a pair of low, coarse, medially convergent ridges. 4) Furcula short, strongly sclerotized, subtriangular with the apex curved inwards.

Description (adult male from Boca de Canasí). Size medium for the genus (total length 18.2 mm).

Coloration. Head basically vivid olivaceous green, except as follows: labrum reddish, vertex with a pair of thin, irregular, longitudinal yellow stripes just above eyes, genae with a thick, irregular, longitudinal yellow stripe just below each eye; antennae blackish with whitish tips, scapus, pedicel and basal segments of flagellum pale bluish; eyes medium gray, with a bluish to greenish shade (depending on the view angle). Pronotum pale olivaceous green, with two irregular pale lateral stripes that are continuous with those on the head and extend continuously through epimera I–II and leg III coxae, these pale stripes are mostly yellow but alternate with bright white spots on anterior and posterior corners of pronotum and basal segments of abdomen, and are interrupted by the green transversal grooves. Epimera I and II each with a conspicuous bright yellow spot. Tegmina with a conspicuously bicolored, longitudinally striped pattern: three thin, equidistant black stripes (anterior, median and posterior), separated by two thick, white stripes. Fore and mid legs of same coloration: coxa and trochanter olivaceous green, femur light reddish, tibia and tarsomeres pale bluish with irregular infuscation; hind legs with coxa and trochanter olivaceous green, femur pale olivaceous green, conspicuously darker on transversal ridges and upper and lower margins, hind knees with upper half blackish and lower half dark bluish, tibia bright blue with spines blackish, tarsomeres whitish. Abdomen with complex longitudinally striped pattern on tergites: single dorsomedian (thin, dark bluish green), paired submedians (thick, pale reddish-cream), paired dorsolaterals (thin, dark bluish green) and paired laterals (thick, vivid red), lateral margins of tergites conspicuously infuscate; sternites vivid red, essentially immaculate. See figure 5, 25e –f and table I.

Head (figs. 16–18). Moderately large, wider than long (ratio = 1.3). Tegument shiny, but sparsely and coarsely punctate, with thin and short setae scattered all over. Vertex convex in lateral view, subtriangular in dorsal view; fastigium prominent, broad in dorsal view, semicircular in lateral view and strongly compressed in frontal view. Eyes very large, rounded and prominent; ocelli minute. Costa frontalis very well-marked and keyhole-shaped, with lateral branches almost parallel and shallowly sinuose, i.e., upper portion wider and inverted U-shaped, lower portion narrower and almost circular. Genae almost straight in frontal view, but not flat. Frons coarsely and sparsely punctate. Antennae standard for Acrididae in size and shape, with 18/18 flagellomeres; scapus subcylindrical, wider than long (ratio = 1.1), oval in cross-section, mostly glabrous; pedicel clearly of more than half the length of scapus.

Thorax (figs. 19–20). Tegument shiny, but densely and coarsely punctate and with thin, short setae scattered all over. Pronotum subrectangular in dorsal view, longer than wide (ratio = 1.1); anterior and posterior margins clearly bilobed, the former shallowly notched medially, the latter moderately notched medially, lateral margins shallowly sinuose; median keel strong, lateral keels irregularly crenulate and deeply incised by the three transverse grooves. Metanotum wider than long (ratio = 2.3) and shorter than abdominal segment I (ratio = 0.7), trapezoidal in dorsal view with tegument coarsely punctate and shiny. Tegmina very short (reaching anterior margin of abdominal segment I), narrow and digitiform, costal margin basically straight, anal margin slightly sinuose and tip rounded.

Legs (fig. 5). Covered all over by thin and short setae. Profemur slender, cylindrical and unarmed; protibia slightly shorter than profemur (ratio = 0.9), very slender and straight, ventral surface with 6:5 / 4:5 subapical spines and two small apical spurs. Mid legs very similar to forelegs, but mesotibia slightly longer than mesofemur and with 3:8 / 8:3 subapical spines. Metafemur robust, longer than metatibia (ratio = 1.1), oval in cross-section; metatibia with 6:8 / 8:5 subapical spines and five inward-curved apical spurs: two small-sized laterodorsals, two medium-sized laterals, and one large ventral.

Abdomen. Large and slender (conspicuously narrower than pronotum), subcylindrical and evenly tapering posteriorly. Tegument shiny but coarsely and densely punctate, with long setae scattered mainly in the ventral surface and abdominal segments with median carina very well marked. Tympanic organ large, rounded and located laterally on segment I. Furcula strongly sclerotized, short, curved inwards and with a sharp tip. Supra-anal plate (fig. 21) relatively large, subpentagonal, wider than long (ratio = 1.1), and sparsely setose; anterior margin shallowly convex and medially moderately lobed, lateral margins straight and essentially parallel, posterior margin narrowly paraboloid; median sulcus complete, moderately deep and narrow, flanked by a pair of low, coarse, medially convergent ridges. Cerci very short, conical and densely covered by thin setae, reaching posterior margin of Supra-anal plate but not surpassing abdomen tip. Subgenital plate (fig. 23) suboval, slightly wider than long (ratio = 1.1) and sparsely setose; anterior margin shallowly concave, lateral margins shallowly convex, posterior margin strongly convex. See figures 5, 21–23 and table I.

Female lectotype: Very similar to male, except as follows: slightly larger (23.0 mm), more robust, with tegmina slightly narrower. See original description by Carl (1916) and figs. 6, 16–23 herein.

Comparisons. D. multicolor stat. resurr. is easy to distinguish from the other 17 members of the genus by its unique color pattern, i.e., head and pronotum with four solid alternate stripes of vivid green and yellow, combined with lower half of abdomen uniformly bright red (figs. 5–6).

Distribution (fig. 24). This species is confirmed to occur only in northwestern Cuba , in northern Havana-Matanzas Coastal Plain.

Ecological notes. As usual for ordinary members of the genus, D. multicolor stat. resurr. seems to be rare and/or highly localized. The Havana-Matanzas Coastal Plain is undoubtedly the longest and most intensively sampled region in Cuba , but only four specimens have been collected and a single precise locality is confirmed. There at Boca de Canasí, the same situation holds true: despite multiple and intensive day-andnight searches, the two available specimens were found together at the same spot.

These specimens from Boca de Canasí were found at dusk, standing in close proximity on the same twig of the bush Zanthoxylum fagara (L.) Sarg., 1890 ( Rutaceae ), less than 1.0 m above the ground, in a sea grape forest. Repeated searches at the same spot and surroundings (including the adjacent semicaducifolious forest and xerophytic scrub), were totally unsuccessful.

One of the males from Boca de Canasí had an unidentified egg-like organism attached to right epimeron II. The other was taken as subadult and molted to adult at night, a few hours after collection.

Remarks. The comparison of the name-bearing types of D. insulana (holotype) and D. multicolor stat. resurr. (lectotype), plus two additional, fresh adult males of the latter, revealed that both taxa are clearly not conspecific. Such synonymy was introduced by Amédégnato et al. (1995) as follows: " The synonymy of both specific names is unquestionable from the study of the types of both species and the comparison of their photographs. The differences annotated by Carl (1916) do not exist in the specimens with respect to the ovipositor. Its distinct coloration corresponds to that specified by Zayas (1976) and we believe it is due to intraspecific variability " ( Amédégnato et al., 1995: 694; originally in Spanish, italics and English translation added herein). Nevertheless, such arguments were actually flawed: 1. Amédégnato et al. (1995) wrongly regarded the holotype and the false type of D. insulana (same two specimens examined herein) as conspecific and mistook the evident differences as intraspecific variation, apparently by regarding them as sexual dimorphism. 2. Because the false type of D. insulana is very similar in coloration to the lectotype of D. multicolor stat. resurr. designated by them (see figs. 3 and 6 herein), Amédégnato et al. (1995) also wrongly concluded that both taxa were conspecific, missing the crucial point that the identity of the former is indivisibly linked to its name-bearing type, i.e., the holotype male. 3. Amédégnato et al. (1995) trusted the D. insulana concept of Zayas (1976), who nevertheless did not study its holotype nor the (then) syntypes of D. multicolor stat. resurr. and even did not mention the latter. Strangely, they overlooked an interesting comment given on the same page: " It looks like there are many species members of this genus... " ( Zayas, 1976: 100; originally in Spanish, italics and

English translation added herein).

The present study of multiple samples where both sexes of the same species are included (e.g., D. atrostriata n. sp., D. megalapida n. sp. and D. multicolor stat. resurr., plus the available literature on Hispaniolan and Jamaican taxa) revealed that adult intraspecific variation is virtually nonexistent and sexual dimorphism is largely restricted to genitalia and does not affect the color pattern (i.e., males are slightly smaller and more slender than females and their coloration is only subtly brighter, but keeps exactly the same pattern).

On the basis of the argumentation given above, D. multicolor Carl, 1916 is herein restored as valid species and the following differences support its accurate distinction from D. insulana : 1. General coloration. D. multicolor stat. resurr.: sharply contrasting, head and pronotum with four solid alternating stripes of vivid green and yellow, abdomen with lower half uniformly bright red. D.

insulana: plain, basically olivaceous green, only with a blurred pair of dark brown lateral stripes. See figs. 2, 5–6. 2. Shape of pronotum. D. multicolor stat. resurr.: anterior margin shallowly notched medially, lateral margins shallowly sinuose, posterior margin moderately notched medially. D. insulana : anterior margin moderately notched medially, lateral margins convex, posterior margin deeply notched medially. See figs. 19–20. 3. Shape and length of tegmina. D. multicolor stat. resurr.: digitiform and very short, not reaching posterior margin of metanotum. D. insulana : lanceolate and short, but reaching anterior margin of abdominal segment II. See figs. 19–20. 4. Coloration of epimera I–II. D. multicolor stat. resurr.: each with conspicuous bright yellow spot.

D. insulana : each with conspicuous bright red spot. See figs. 2, 5–6.

5. Furcula. D. multicolor stat. resurr.: strongly sclerotized, short, falcate. D. insulana : normally sclerotized, minute, triangular. See fig. 21.

6. Supra-anal plate. D. multicolor stat. resurr.: wider than long, subrectangular, sparsely setose, median sulcus broad. D. insulana : as long as wide, subpentagonal, moderately setose, median sulcus narrow. See fig. 21.

The exact locality and collection date of the types of D. multicolor stat. resurr. were not stated in the original description, nor included in their labels (fig. 4c); nevertheless, there is a supplementary literature source that clarifies both data. Hollier & Hollier (2012: 238) reported that Saussure traveled to Cuba from January 29 th through February 22 th, 1855, and visited the following localities: Havana, Guanabacoa, Santa María del Rosario, Matanzas and Limonar.

All these sites are well-known cities and towns scattered across 100 km of the northern part of Havana-Matanzas Coastal Plain, in the northwestern region of Cuba (fig. 24). The types of D. multicolor stat. resurr. were most likely collected somewhere along this route, and following Article 76.2 and Recommendation 76A.1 of the Code (ICZN, 1999: 71), their originally vague type data are herein corrected accordingly (see above, in the Type data section). It is worth mentioning here that the single locality where specimens of this species have been found again (Boca de Canasí) is not only placed within Saussure's route, but roughly in its center (fig. 24).

Figures

Fig. 4. Original labels of: a) adult male holotype of Dellia insulana; b) adult female of Dellia aff. multicolor stat. resurr., wrongly labeled as paratype of Dellia insulana (see detailed discussion in the text); c) adult female lectotype of Dellia multicolor stat. resurr. Photos courtesy G. Lindberg (a–b) and J. Hollier (c).

Fig. 6. Adult female lectotype of Dellia multicolor stat. resurr., full-body views: a) dorsal; b) lateral; c) ventral. Scale bar in millimeters. Photos courtesy J. Hollier.

Fig. 16. Dorsal close-up of head and anterior margin of pronotum of all Cuban Dellia recognized in the present revision.

Fig. 17. Lateral close-up of head and anterior margin of pronotum of all Cuban Dellia recognized in the present revision.

Fig. 18. Frontal close-up of head of all Cuban Dellia recognized in the present revision.

Fig. 19. Dorsal close-up of thorax and abdominal segment I of all Cuban Dellia recognized in the present revision.

Fig. 20. Lateral close-up of thorax and abdominal segment I of all Cuban Dellia recognized in the present revision.

Fig. 21. Dorsal close-up of abdominal apex of all Cuban Dellia recognized in the present revision.

Fig. 22. Lateral close-up of abdominal apex of all Cuban Dellia recognized in the present revision.

Fig. 23. Ventral close-up of abdominal apex of all Cuban Dellia recognized in the present revision.

Fig. 24. Geographical distribution of all Cuban Dellia recognized in the present revision. Note three nearby localities for D. atrostriata n. sp. and precise occurrence still wanting for D. insulana. The upper left inset (frame equals to 150 x 75 km) depicts the single precise locality known for D. multicolor stat. resurr. (red symbol), superimposed to all sites visited by Saussure in 1855 (yellow symbols), from left to right: Havana, Guanabacoa, Santa María del Rosario, Matanzas and Limonar.

Fig. 25. Live habitus of: a) Dellia albida n. sp., adult male holotype, in captivity; b–c) Dellia atrostriata n. sp., adult male from El Cojo, in nature; d) Dellia brauni n. sp., adult male holotype, in nature; e–f) Dellia multicolor stat. resurr., adult male topotype, in nature; g–i) Dellia megalapida n. sp., adult male holotype (g), adult male and adult female paratypes while mating (h), subadult female paratype (i), all in nature; j–k) Dellia melici n. sp., adult female holotype, in nature; l) Dellia zephyra n. sp., adult female holotype, in nature. Photos courtesy Rayner Núñez (b, c), Rubén Marrero (d), and Tomás M. Rodríguez (j, k, l).

Fig. 26. Habitat at the exact collecting site of: a) Dellia albida n. sp; b) Dellia atrostriata n. sp.; c) Dellia brauni n. sp.; d) Dellia megalapida n. sp., with the author and her collaborator Rolando Teruel while photographing some of the type specimens; e) Dellia melici n. sp.; f) Dellia multicolor stat. resurr.; g) Dellia siboney n. sp.; h) Dellia zephyra n. sp., note the vegetated pits (micro-dolines or large skylights) all along Las Perlas cave, the holotype was found in the dark passage connecting two of these pits. Photos courtesy Rolando Teruel (b, f, h), Tomás M. Rodríguez (c, e), and Luis A. Casanella (d).