Calligrapha elegantula Jacoby, 1877

Gómez-Zurita, Jesús, 2015, Systematic revision of the genus Calligrapha Chevrolat (Coleoptera: Chrysomelidae, Chrysomelinae) in Central America: The group of Calligrapha argus Stål, Zootaxa 3922 (1), pp. 1-71 : 46-50

Calligrapha elegantula Jacoby, 1877

Jacoby, M. 1877: 519.

( Figs 7 , 12 b, 13i, 13j)

Calligrapha elegantula: Jacoby, 1882 , Biol. Centr. Am., vol. 6, pt. 1, p. 208. Polyspila elegantula: Weise, 1916 , Col. Cat., pars 68, 12, p. 39.

Calligrapha elegantula: Blackwelder, 1946 , Checklist Col., Pt. 4, p. 674. Calligrapha elegantula: Wilcox, 1975 , Checklist, Biol. Res. Inst. Amer., p. 66. Calligrapha elegantula: Flowers, 2004 , Rev. Biol. Trop., p. 80.

Calligrapha elegantula: Gómez-Zurita et al., 2006 , Evolution, 60, p. 332. Calligrapha elegantula: Montelongo & Gómez-Zurita, 2014 , Zool. Scr. 43, p. 607.

In the original description for this species, there is no indication about the number of specimens that M. Jacoby used to describe it, but it is possible that it was a single specimen (e.g., beetle length is given as a single figure, not a range), and the only useful information links the type material to his own collection. This collection is mostly split between MCZ, MfN and NHM and there are specimens labelled as types in at least two of the Institutions. Most specimens, including these putative types, are labelled as originating from the Godman-Salvin expedition to Central America, or include collecting data compatible with this expedition (e.g., Ch. van Patten or H. Rogers as collectors, or the localities that the latter visited: Cache, Irazu). Based on M. Jacoby's introduction to his Phytophaga volumes in Biologia Centrali Americana, dated May 1892, he had been working on this material for twelve years, in any case with posteriority to having described C. elegantula Jac. Therefore , the identity of the type specimen has been lost, and a new type needs to be designated. This narrowly distributed species is also extraordinarily constant in patterns and colouration therefore any of the specimens is a good representation for the taxon. However, the one in particular labelled as type at MCZ deviates from the original description in presenting narrow areas connecting arcuate band and subsutural stripe (described as completely free) and most additional spots appear fused together in different degrees, while they are described as "sometimes connected with each other" (Jacoby 1877). I arbitrarily select here as neotype one specimen at NHM, representative of the variability found in the species and fully fitting Jacoby's description.

Neotype: R. Susio, Costa Rica. H. Rogers / C. elegantula Jac. [pale blue] / Godman-Salvin Coll., Biol. Centr.- Amer. (NHM). Specimen lacks right metatarsus.

Habitus ( Fig. 12 b). Length: 8.05 mm, width: 4.98 mm. Body elongated oval. Head, mandibles, disc of labrum, antennae, mouth pieces, pronotum, scutellum, elytral markings, epipleura, ventral surfaces and legs black, shiny, with a hint of greenish metallic reflection; margins of labrum, narrow apex of antennal and maxillary segments, apex of mandibles, knees and claws dark rufous; background of elytra dark yellow, creamy yellow around markings.

Head large, deeply inserted in pronotum; surface finely microreticulate, moderately punctured, sparsely on frons; supraocular furrow impressed, straight, from antennal calli to shortly behind upper eye margin; frontal suture fine, deeper in depressed area of epistome, joining broadly V-shaped clypeal suture. Clypeus finely microreticulate, moderately strongly punctured with fine, long setae curved towards middle. Antennae slender but short, reaching humeri; antennomeres 2–6 elongated, thin, relatively smooth and almost glabrous; second antennomere half as long as first, third slightly shorter than first; antennomeres 3–5 shortening progressively, sixth longer than second; antennomeres lengthening gradually beyond fifth segment; antennomeres 7–11 thicker, widening from base to apex, rugose and pubescent, eleventh about as long as first, tappering towards blunt apex and slightly emarginate dorsally; eighth antennomere 0.86x as wide as long. Labrum relatively long, smooth, emarginate anteriorly, with very long fine setae convergent medially. Mandibles large, sturdy, densely punctured except at apical molar area, with very long fine curved setae directed to middle; sides weakly concave before strong, regularly curved preapical curvature. Maxillary palpi long, pubescent with long fine setae; last palpomere subtrapezoidal, broader apically than basally, sides subparallel before basal curvature; previous palpomere twice as broad apically as basally, elbowed. Pronotum transverse (W/L=1.96), sides straight, narrowly convergent from base, regularly smoothly curved at apical third towards moderately produced anterior angles; anterior border concave, posterior border convex; surface microreticulate, sparsely double punctured on disc, with fine and sparser minute punctures, stronger, deeper, sometimes confluent at sides; some round punctures near and at basal margin at sides; sides and apex with fine margin, invisible from above at sides; anterior angles with long setae strongly bent dorsolaterally backwards, and posterior angles with long setae bent laterally forwards. Hypomeral suture deep in basal 2/3, less impressed apically, strongly divergent from margin; hypomera shagreened, convex medially, unpunctured, transversally wrinkled basally. Prosternum concave and finely margined apically, microreticulate, punctured near coxae and laterally on process, with long fine setae; prosternal process narrow between coxae, sloping and weakly expanded at straight-cut apex. Mesanepisterna shagreened, with few irregularly impressed punctures at posterior angle. Metepisterna with weakly raised margins, densely punctured, almost rugose on shagreened background. Metaventrite leathery at sides, punctured mainly at anterior angles with fine long hairs on punctures, smooth and shiny medially. Scutellum small, lancet-like and relatively elongate (W/L=0.88), very finely microsculptured, unpunctured. Elytra elongated, broader than pronotum basally, with faintly marked shoulders, smoothly curved at sides before regular apical curvature; minutely, very sparsely punctured on pale areas, except by very regular premarginal line and at apical declivity, with scutellar row present; punctures around and within dark markings stronger. Markings: (i) sutural and subsutural stripes forming a continuous sutural marking, broadest at level with apex of scutellum, quickly narrowing at base almost reaching border of scutellum basally, and gradually narrowing posteriorly to apical angle of elytra, slightly widening again at apical declivity; (ii) arcuate band free on disc, gently curved, expanded apically as large subsquare spot; (iii) humeral spot and humeral lunule broadly confluent as large humeral marking, elongated on shoulder, with apex of lunule curved towards suture, slightly surpassing basal end of arcuate band; (iv) spot enclosed by humeral lunule large, elongated oval; (v) spot of apical declivity large, subtriangular, longitudinally elongated, free; (vi) apical spot round, medium-sized, free; (vii) eight additional medium-sized to large markings on disc; largest irregular in shape between apical enlargement of arcuate band and elytral margin; one medium-sized round spot at concavity of arcuate band; two medium-sized round spots transversally arranged at apical declivity of elytra; five large spots in lateral declivity arranged in two rows parallel to margin, externally with three spots (the apicalmost largest, at level with apical enlargement of arcuate band), internally with two. Legs slender; femora shiny, finely, relatively densely punctured, with fine long and relatively dense hairs, mainly ventrally. Tibiae very hairy ventrally, with dense and long golden setae apically on rugose background; weakly carinated internally, shortly furrowed externally. Abdominal ventrites shiny, smooth, finely, densely punctured at basal half, with fine hairs; last abdominal ventrite punctured and hairy on all surface. Penis figured in Figs 13 i, 13j.

Distribution ( Fig. 7 ). The species, which is not rare in collections, was until now considered endemic from Costa Rica, most frequent in the northern half of the country. However, in the course of this study I have found specimens collected in coastal localities along the Bahia of Coronada in the Pacific south of Costa Rica, and one specimen seemingly collected in a Panamian locality, nearby the Costarican border, expanding its known range to the south. In any case, it is a Caribbean Mesoamerican endemic species. The Colombian and Mexican records at NMNH most likely represent mislabelled specimens.

Material examined (291 specimens).

COLOMBIA

NMNH: (1) one specimen: Colombie, gift of F.C. Bowditch, gift ex MCZ Dupl. Series.

COSTA RICA

EGRC: (1) one specimen: Costa Rica, Punt. Pr., Monteverde area, 4–6.vi.1980, J.E. Wappes, Calligrapha elegantula Jac. Daccordi ’81; (2) twenty-six specimens: Costa Rica, Puntarenas, Monteverde, 26.v–3.vi.1984, E. Riley, D. Rieder & D. LeDoux; (3) three specimens: Costa Rica, Puntarenas Prov., Monteverde 1300–1600 m, 6.i.1974, C. Barfield, taken from vegetation; (4) two specimens: Costa Rica, Guan., La Pacífica nr. Cañas, 22–26.v.1984, E. Riley, D. Rieder & D. LeDoux. FSCA: (1) one specimen: Costa Rica, Puntarenas, Monteverde area, 4–6.vi.1980, J.E. Wappes, Calligrapha elegantula Jac. Det. E.G. Riley ’82; (2) two specimens: Costa Rica, Puntarenas, Monteverde, 3.vi.1984, E. Riley, D. Rider & D. LeDoux; (3) one specimen: Costa Rica, Ala. Province, 6–8 km W Atenas, 1–2.vi.1980, J.E. Wappes, Calligrapha elegantula Jac., Det. E.G. Riley ’86; (4) one specimen: Costa Rica, Puntarenas, San Luis (Monteverde) 3900’, 12–13.v.1996, E. Giesbert; (5) one specimen: Costa Rica, San Jose, Monteverde, 21.vi.1976, at blacklight, J.K. Balciunas & P.P. Habeck; (6) one specimen: Costa Rica, Puntarenas, Monteverde, 27.ii.1987, E. Giesbert; (7) two specimens: Costa Rica, Puntarenas, 4 km NE San Luis de Guacimal, 22.ii.1987, E. Giesbert. IBE-JGZ: (1) three specimens: [IBE-JGZ-0073, IBE-JGZ-0074, IBE-JGZ- 0075] Costa Rica, Zarcero, Alfaro Ruíz, 4/vi/2000, A. Solís leg., Calligrapha elegantula Jacoby, 1877 J. Gómez- Zurita det. 2003. MCZ: (1) one specimen4: Cache, Costa Rica, H. Rogers, 1st Jacoby Coll., Type 17438 [red label], 4. Specimen labelled as type prior to this study.

Jan. Jul.2004. MCZ Image Database; (2) one specimen: 473, Costa Rica, 473 Calligrapha elegantula Jac. [male; female] […]; (3) one specimen: C. Rica, elegantula ; (4) one specimen: Costa Rica; (5) one specimen: Cartago; (6) one specimen: Costa Rica, Jacoby 2nd Coll.; (7) two specimens: Costa Rica, Van Patten [one with: C. elegantula Jac. ]; (8) one specimen: C. Rica, Calligrapha elegantula Jac. ; (9) one specimen: Collection Schild-Burgdorf, Costa Rica, Zarzero, 7, F.A. Eddy Collection; (10) one specimen: Collection Schild-Burgdorf, Costa Rica, San Jose; (11) five specimens: Hamburg Farm, March, Costa Rica, C.P. Dodge; (12) seven specimens: C. Rica [one with: Cal. elegantula Jac. ]; (13) three specimens: Cache, Costa Rica, H. Rogers, 1st Jacoby Coll.; (14) one specimen: Irazu 6–7000 ft, H. Rogers, 1st Jacoby Coll.; (15) four specimens: Costa Rica, Jacoby 2nd Coll.; (16) one specimen: Bebedero, Costa Rica (Underwood, 94); (17) one specimen: San José, Costa Rica (Underwood); (18) seven specimens: C. Rica; (19) one specimen: Escazu, Costa Rica, Calligrapha elegantula Jacoby J. Gómez-Zurita det. 2010; (20) one specimen: Cariblanco, Costa Rica, Calligrapha elegantula Jacoby J. Gómez-Zurita det. 2010. MfN: (1) four specimens: Costa Rica, Van Patten, 96374 [one with: C. elegantula Jac. ]; (2) one specimen: Irazu, 6–7000ft., H. Rogers, Calligrapha elegantula Jac. ; (3) one specimen: R. Susio, Costa Rica, H. Rogers; (4) three specimens: Carthago, S. José, Costa Rica, C. Wercklé G.; (5) five specimens: 39600, Costa Rica, Fratz. [four specimens without data]; (6) three specimens: Collection Schild-Burgdorf, Costa Rica, San Jose [one with: elegantula Jac. ]; (7) two specimens: Collection Schild-Burgdorf, Costa Rica, Zarzero. MTJM: (1) two specimens: Costa Rica, Monteverde, 10º29’N 84º50’W, 1400 m, R.D. Akre, Calligrapha elegantula Jacoby J. Gómez-Zurita det. 2009 [one with: Feb. 5, 1963; one with: Feb. 6, 1963]. NHM: (1) twelve specimens: Costa Rica, Sharp Coll. 1905-313; (2) four specimens: Costa Rica, Rogers, Sharp Coll. 1905-313; (3) two specimens: Costa Rica, 92-48 [one with: Calligrapha elegantula Jac. ]; (4) two specimens: Costa Rica, Van Patten, Godman-Salvin Coll., Biol. Centr.-Amer. [one with: Calligrapha elegantula ]; (5) seven specimens: R. Susio, Costa Rica, H. Rogers, Godman- Salvin Coll., Biol. Centr.-Amer. [one with: C. elegantula Jac. ]; (6) three specimens: Irazu, 6–7000ft, H. Rogers, Godman-Salvin Coll., Biol. Centr.-Amer. NMB: (1) one specimen: Collection Schild-Burgdorf, Costa Rica, Atenas, Calligrapha elegantula J. Bechyné det. 1951; (2) one specimen: Collection Schild-Burgdorf, Costa Rica, Zarcero; (3) two specimens: Carpintera, Costa Rica, Reimoser. NMCZ: (1) one specimen: Costa Rica, 475, Coll. Achard Mus. Pragense; (2) six specimens: Costa Rica, Coll. Achard Mus. Pragense. NMNH: (1) two specimens: Quepos, Puntarenas C.R., 19-5-55, on Sida sp. [one with: Calligrapha elegantula Jacoby J. Gómez-Zurita det. 2011]; (2) one specimen: La Carpentera, Costa Rica, F. Monrós Collection 1959, Calligrapha elegantula Jac., F. Monrós det. 1954; (3) two specimens: San Jose, Costa Rica, F. Monrós Collection 1959; (4) two specimens: C. Rica, F. Monrós Collection 1959; (5) three specimens: Costa Rica, 30-iii-1948, F. Monrós Collection 1959; (6) twelve specimens: Costa Rica, El Alto de Ochomogo, 28.vi. 934, C.H. Ballon s/ maiz escobilla, F. Monrós Collection 1959; (7) one specimen: Costa Rica, San Ramón de Tres Ríos, ix.1991, J.M. Maes, Calligrapha elegantula Jacoby 1991 Det. C.L. Staines ; (8) one specimen: Irazu, Costa Rica, Coll. Schild & Burgdorf; (9) four specimens: Zarzero, Costa Rica, Coll. Schild & Burgdorf; (10) one specimen: Costa Rica, Van Patten; (11) two specimens: San Jose, C.R., J.F. Tristan coll. [one with: Apr. 28]; (12) three specimens: C. Rica, F. Monrós Collection 1959; (13) one specimen: La Palma, C.R., 1500 m alt., C.H. Lankester collector, Calligrapha elegantula Jac. F. Monrós det. 1953; (14) one specimen: San Jose, C.R., 1160 m, M. Valerio coll., Calligrapha elegantula Jac. F. Monrós det. 1953; (15) eight specimens: El Alto de Ochomogo, C.R., 6.28.34, C.H. Ballou, on corn escobilla, C.R. No 1368 [one with: Calligrapha elegantula Jac. HSB]; (16) twelve specimens (within a capsule): El Alto de Ochomago, C.R., 6-28-34, C.H. Ballou, on corn+escobilla, C.R. No. 1368, Calligrapha elegantula Jacoby J. Gómez-Zurita det. 2003; (17) one specimen: Irazu, 6–7000 ft, H. Rogers; (18) one specimen: Costa Rica, V. Patten, Calligramma elegantula Jac. ; (19) one specimen: La Carpentera, Costa Rica, April 1924, W.M. Mann; (20) one specimen: San Jose, C.R., 1160 m, M. Valerio coll.; (21) one specimen: Costa Rica, J.F. Tristan coll.; (22) one specimen: 11990, Turrialba C.R., 27-2-71, J.V. Mankins collector, Calligrapha elegantula Det. 1977 , J.V. Mankins; (23) two specimens: Costa Rica, Heredia, Finca La Selva, 21–30 July 1976, J.C. Solomon Coll.; (24) six specimens: Costa Rica, F. Nevermann, iii.33, Guayabillos SW Abhang Irazu 2200 m [one with: iii.33]; (25) one specimen: Costa Rica, F. Nevermann, 25.viii.28, San Jose 1000–1200 m, an grasnarbe; (26) three specimens: Costa Rica; (27) three specimens: Costa Rica, Oct.; (28) two specimens: Coronado, Costa Rica, July 1923, T. Assman, Nevermann Collection 1940; (29) two specimens: San Jose, C.R., March 1921, Nevermann Collection 1940; (30) one specimen: Costa Rica, Monteverde, November 1981, N.J. Besansky; (31) ten specimens: Costa Rica, Puntarenas, Monteverde area, 6–14 June 1973, 1400–1700 m, Erwin & Hevel Central American Expedition 1973; (32) two specimens: Costa Rica, Punt. Prov., Monteverde 1430 mts., 23 February 1978, E.R. Snyder Hodges; (33) four specimens: Costa Rica, Heredia, 1800 m, Monte de la Cruz, 1 July 2001, C.L. Staines; (34) one specimen: on bromeliad plants, San Jose, Costa Rica, iv.5.61, A.S. Mills Coll, Miami 16264, 10t61-11378, Calligrapha sp. det. D.M. Weisman; (35) one specimen: Irazu, 6–7000 ft, H. Rogers; (36) one specimen: Costa Rica, San Jose 10 km N, 9 Aug 1972; (37) one specimen: Costa Rica, Volcan Irazu, 15 May 1951, O.L. Cartwright; (38) one specimen: NW Costa Rica, Oct. 22–31, 1992, Coll. R. Pienkowski; (39) one specimen: Costa Rica, San Jose 1100–1200 m, August 1980, N.L.H. Krauss; (40) one specimen: Costa Rica, Vista de Mar, xi.21.1965, N.L.H. Krauss; (41) two specimens: Costa Rica, Heredia, Hwy32 at San Jose border, 8 January 1995, C.L. & S.L. Staines. NRM: (1) one specimen: Costa Rica, Van Patten, Calligr. elegantula Jac. OUMNH: (1) one specimen: Cache, Costa Rica, H. Rogers, B.C.A. duplicates, presented 1909 by F.D. Godman, Cat. No. 84, Calligrapha diversa Stål , Calligrapha elegantula Jacoby J. Gómez-Zurita det. 2011. TAMUIC: (1) two specimens: [X0537769 and X0540719], Costa Rica, Alajuela, 10 km N San Pedro, 26.vii.1990, William F. Chamberlain; (2) one specimen: X0540763, Costa Rica, Cartago, Juan Viñas, 5.viii.1990, William F. Chamberlain; (3) one specimen: X0540842, Costa Rica, Puntarenas, Monteverde, 2.viii.1990, G.M. Chamberlain; (4) one specimen: X0541102, Costa Rica, Puntarenas, Monteverde, 20–24.vi.1986, W. Hanson & G. Bohart; (5) two specimens: [X0541496 and X0547150], Costa Rica, San José, San José, 19–20.xii.1987, F.D. Parker; (6) two specimens: [X0547858 and X0549945], Costa Rica, San José, San José, 27–29.xii.1987, F.D. Parker; (7) five specimens: [X0542347, X0546293, X0546589, X0547506 and X0550458], Costa Rica, Puntarenas, Monte Verde, 6.i.1974, taken from vegetation, Carl Barfield; (8) one specimen: Costa Rica, San José, Escazu, 5–6.vii.1988, F.D. Parker.

MEXICO

NMNH: (1) one specimen: Mexico, R.F. Pearsall, Brooklyn Museum Colln 1929.

PANAMA

FSCA: (1) one specimen: Panama, Chiriquí, Hartmann’s Finca, 4–6.vii.1997, J. Huether.

UNKNOWN SOURCE

MCZ: (1) one specimen: elegantula ; (2) one specimen: F.A. Eddy Collection; (3) one specimen: 38, Jacoby 2nd Coll. MfN: (1) one specimen: [no data]. OUMNH: (1) one specimen: E. Coll. Laferté, bor, Calligrapha persimilis Reiche, Amer. bor., From the Brit. Mus. duplicates Pres. 1899, Calligrapha elegantula Jacoby J. Gómez- Zurita det. 2011.

Variation. Species very constant in appearance, showing slight differences in elytral maculation, for instance different confluence patterns among additional spots on disc, these with apex of arcuate band, or with apical spot or the spot on apical declivity; the basalmost spot can also be narrowly confluent with humeral marking externally. The basal end of the subsutural stripe is commonly confluent with sutural stripe, but sometimes it is slightly separated from it for a short distance. I have seen one specimen with the spot enclosed by humeral lunule confluent with the latter and another with the basal end of the arcuate band. The spot of apical declivity appeared very small in one specimen. Finally, in a single specimen at NMCZ, the apex of arcuate band presents two consecutive lobes.

Figures

FIGURE 7. Geographic distribution of Calligrapha elegantula Jacoby in Costa Rica and Panama (black circles) and the single Panamanian locality where C. fulvitarsis Jacoby (white circle) has been found.

FIGURE 12. Dorsal and lateral habitus of the Lectotype of Calligrapha aeneopicta Stål (a), Neotype of C. elegantula Jacoby (b), Lectotype of C. sylvia Stål (c), Lectotype of C. diversa Stål (d), and Holotype of C. simillima Stål (e). Scale bar = 5.0 mm.

FIGURE 13. Penis in ventral, lateral and dorsal view of apex of the species Calligrapha simillima Stål (a, d), C. sylvia Stål (b, e), C. diversa Stål (c, f), C. aeneopicta Stål (g, h), and C. elegantula Jacoby (i, j). Scale bar = 0.5 mm.