Calligrapha argus Stål, 1859

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 : 39-45

Calligrapha argus Stål, 1859

Stål, C. 1859: 324.

( Figs 2 d, 6b, 9i, 9j, 10)

Chrysomela argus: Stål, 1865 , Mon. Chrys. Amer., 3, p. 277.

Calligrapha argus: Gemminger & Harold, 1874 , Cat. Col., XI, p. 3432.

Calligrapha argus: Steinheil, 1877 , Mittheil. Münch. Ent. Ver. 1, p. 32

Calligrapha argus: Jacoby, 1882 , Biol. Centr. Am., vol. 6, pt. 1, p. 201.

Calligrapha famularis: Jacoby, 1882 , Biol. Centr. Am., vol. 6, pt. 1, p. 201. Calligrapha argus: Jacoby, 1892 , Biol. Centr. Am., vol. 6, pt. 1, suppl., p. 246. Chrysomela argus: Dugés, 1901 , Cat. Col. Coleópt. Mex., p. 97.

Polyspila argus: Weise, 1916 , Col. Cat., pars 68, 12, p. 38.

Calligrapha argus: Blackwelder, 1946 , Checklist Col., Pt. 4, p. 674.

Calligrapha ramulifera var. argus: Bechyné, 1952 , Entom. Arb. Mus. Frey 3, p. 4. Calligrapha ramulifera s. argus : Blackwelder, 1957, Checklist Col., Pt. 6, p. 1436. Calligrapha argus: Bechyné & Springlová de Bechyné, 1965, Rev. Fac. Agron. Maracay 3, p. 48. Calligrapha argus: Wilcox, 1975 , Checklist, Biol. Res. Inst. Amer., p. 66. Calligrapha famularis: Wilcox, 1975 , Checklist, Biol. Res. Inst. Amer., p. 66. Calligrapha argus: Burgos-Solorio & Anaya-Rosales, 2004 , Acta Zool. Mex. 20(3), p. 45. Calligrapha argus: Flowers, 2004 , Rev. Biol. Trop., p. 80.

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

Jacoby (1882) expressed doubts about the conspecificity of this taxon with C. ramulifera Stål , and this prompted some nomenclature instability by subsequent authors possibly without revising relevant type material, which would have shown at once the remarkable differences between both taxa (e.g., Gómez-Zurita et al. 2006). Bechyné (1952), for instance, proposed that C. argus was but a mere colour variety of C. ramulifera with larger spots on elytra, a stance followed shortly by Blackwelder (1957) in his corrigenda to the Checklist of American beetles, but proposing a subspecific status for the former taxon. Years later, J. Bechyné actually withdraw his earlier opinion explicitly describing several differences found between both species (Bechyné & Springlová de Bechyné 1965), a viewpoint which I completely endorse here.

I found two specimens from Mexico, the locality mentioned in the original description, labelled as types, one in the collection in Stockholm (NRM) and one in London (NHM). The original description of C. argus is, like most others, very succinct but it is possible to deduce that it was based on a single specimen, since the author gave a discrete size for the specimen. The specimen at NRM fits these dimensions as well as the other details in the description. The one at NHM was very likely seen by C. Stål because it is labelled "Campeche", an unusual locality mentioned precisely for this species in his later monograph. However, this origin is not given in the orignal description and the size of the specimen has poorer fit with the measures given by C. Stål, therefore, I designate here as lectotype the specimen in the NRM collection.

Lectotype by present designation: Mexico / Chevrol. / Type / Typus [red] (NRM).

Paralectotype: Campeche, Pilate / Type Stål Coll: Deyrolle / Baly Coll. / argus Stål Mexico [underneath: Type Stål Col. Deyrolle] (NHM).

Habitus (Fig. 6b). Length: 9.39 mm, width: 5.57 mm. Body oval oblong, moderately convex. Head, pronotum, scutellum, epipleura, apendages, ventral surfaces and elytral markings reddish brown; antennae and palpi paler. Apex of mandibles and narrow anterior and lateral margins of pronotum blackish. Elytra creamy yellow, brighter around margin and markings.

Head broad, deeply inserted in pronotum; surface microreticulate, moderately strongly punctured, sparser on vertex and around antennal insertions; longitudinal frontal suture perpendicularly joining obtuse V-shaped clypeal suture; supraocular sulci parallel to frontal suture, running close to dorsal margin of eye; eyes elongated dorsoventrally, finely faceted. Antennae relatively short, reaching humeri; first antennomere long, thick nearly straight posteriorly and convex at anterior border; antennomeres 2–5 elongated, thin, relatively smooth and nearly glabrous; second antennomere half as long as first, third slightly shorter than first; antennomeres 3–6 shortening progressively, sixth subequal to second; antennomeres gradually lengthening beyond sixth antennomere; antennomeres 7–11 thicker, darker, widening from base to apex, rugose and pubescent beyond eighth antennomere; eleventh about as long as first, tappering towards blunt apex and slightly emarginate dorsally. Labrum relatively small, sides regularly curved, anterior border emarginate, laterally on disc with nearly straight, long pale traslucent setae directed forwards. Mandibles large, strong, largely protruding, about 4x beyond apex of labrum; sides concave before strong preapical curvature; surface covered by very strong punctures except at smooth molar area, each puncture with a very long whitish seta. Last maxillary palpomere broad, parallel-sided at apical half, slightly obliquely truncated at apex. Pronotum 1.88 times broader at base than long medially; surface microsculptured with irregularly sparse punctation, stronger (2– 3 x) and denser towards sides; apical border bisinuated, finely margined behind head, with markedly protruding anterior angles, with thicker margin; posterior border broadly convex, unmargined; sides margined, nearly straight, subparallel at basal 1/3, gently and regularly curved towards anterior angles. Hypomeral suture ( Fig. 1 d) deep, parallel to pronotal margin, curved basally, close but not reaching pronotal base, and regularly curved apically, following basal contour of anterior pronotal angle; hypomera finely shagreened, unpunctured, finely transversely wrinkled at base. Prosternum punctured near coxae; prosternal process narrow between coxae, gradually expanding apically, reaching slightly beyond coxae, cut almost straight at apex. Metepisterna strongly punctured; punctures elongated, dense, confluent at apical 1/3. Metaventrite regularly punctured at sides, with fine sparse pubescence. Scutellum narrowly triangular (W/L=0.7) with sides gently curved; feebly convex, shiny, unpunctured. Elytra slightly broader than pronotum; surface finely and sparsely punctured with brownish minute spots, punctures irregularly scattered except at regular premarginal line; space between premarginal line and margin of elytra smooth, unpunctured; dark markings surrounded by stronger punctures; scutellar row of punctures present. Markings: (i) uninterrupted sutural stripe basally surrounding scutellum and reaching elytral apex suddenly and obliquely narrowed; (ii) subsutural stripe divergent from sutural stripe basally, shortly for 5–6 punctures; slightly widened preapically; (iii) arcuate band continuous and confluent for most of its length with subsutural stripe, except briefly at basal end and apically, shaped as big round spot; (iv) humeral lunule well defined, also by surrounding punctures; base detached from basal margin of elytron, apex at level with basal end of arcuate band; confluent basally by half of its length with (v) elongated humeral spot, basally free from basal margin of elytron, slightly broader than humeral lunule, with punctures inside marking and imperfectly surrounded by punctures; (vi) spot enclosed by humeral lunule large, roundish with small basal emargination but punctures defining U-shape; (vii) spot of apical declivity of elytra large and laterally confluent with preapical enlargement of subsutural stripe; (viii) apical spot round, free; (ix) midlateral spot large (seven punctures of premarginal line), subrectangular, laterally confluent with brownish elytral margin; (x) nine additional discal spots loosely arranged in a 3-1-2-1-2 oblique pattern. Legs mostly unpunctured, very finely and sparsely pubescent except at tibial apices, with dense golden pubescence; tibiae broadening apically, externally furrowed at apical third, furrow broadening towards tarsal insertion. Penis as in Figs 9 i, 9j.

Distribution. This is one of the species in the group with a wider distribution, ranging from the southern half of Mexico (particularly in the Caribbean domain) to Venezuela, through Guatemala, Belize, Honduras, El Salvador, Nicaragua, Costa Rica, Panama and Colombia ( Fig. 10 ). It is one of the three species in this evolutionary lineage that possibly reached the Caribbean NW South American region from its main distribution in the Caribbean Mesoamerican domain in recent times (Montelongo & Gómez-Zurita, 2014).

Material examined (350 specimens).

BELIZE

NMB: (1) one specimen: Br. Honduras, Punta Gorda, 1915, Col. R. Vitalis, Calligrapha argus Stål J. Bechyné det. 1954. NMNH: (1) one specimen: Belize, Toledo dist., Blue Creek Village, 18 June 1981, W.E. Steiner, Earthwatch Belize Expedition 1981, D.H. Messersmith, W.E. Steiner et al., Calligrapha argus Stål J. Gómez-Zurita det. 2011.

COLOMBIA

MCZ: (1) one specimen: Sevilla, Mgd. Colombia, v.6.28, Darlington, Calligrapha argus Stål J. Gómez-Zurita det. 2010. NHM: (1) one specimen: Calligrapha pacta Dej. Columbia , [illegible], Chrysomela argus Stål, Baly Coll. ; (2) one specimen: E. Coll. Laferté, 426, Calligrapha pacta Dej. Colombia, argus Stål Stål , 67-56; (3) one specimen: E. Coll. Laferté, 479, Granada, 67-56. NRM: (1) one specimen: Columbia, Stål.

COSTA RICA

EGRC: (1) one specimen: Costa Rica, Monte Verde, Cordillera de Tilarán, 10.iii.1991, M.E. Rice, Calligrapha argus Stål J. Gómez-Zurita det. 2011. FSCA: (1) one specimen: Costa Rica, Guanacaste, Finca Taboga Res. Farm, 9.vii.1966, G.R. Buckingham; (2) two specimens: Costa Rica, Ala. Pr., 8 km S S. Ramon, 31.v.1980, J.E. Wappes, Calligrapha argus St. Det. E.G. Riley ’80; (3) one specimen: Costa Rica, Guanacaste, La Pacífica, nr. Cañas, 8.vi.1983, J.E. Wappes; (4) two specimens: Costa Rica, Pun., 13 mi NW Esparza, 250’, 19.vi.1974, C.W. & L.B. O’Brien & G.B. Marshall, Calligrapha argus Stål J. Gómez-Zurita det. 2011; (5) one specimen: Costa Rica, Puntarenas, Monteverde, 23–27.v.1987, E. Giesbert, Calligrapha argus Stål J. Gómez- Zurita det. 2011; (6) four specimens: Costa Rica, Puntarenas, 4 km NE San Luis de Guacimal, 22.ii.1987, E. Giesbert, Calligrapha argus Stål J. Gómez-Zurita det. 2011. IBE-JGZ: (1) one specimen: IBE-JGZ-0077, Costa Rica, Guanacaste, Area Conservación Guanacaste, Sector Santa Elena, Potrero Grande, 27/05/2000, A. Solís leg. MfN: (1) one specimen: Costa Rica, Piedras Negras, Collection Schild-Burgdorf, Calligrapha notatipennis Stål. NHM: (1) one specimen: Costa Rica, 92-18, C. suboculata Stål ; (2) one specimen: Irazu, 6–7000ft, H. Rogers, Godman-Salvin Coll., Biol. Centr.-Amer.; (3) one specimen: Surface of fallen tree, Dry tropical forest, Costa Rica, Volcán Rincón de la Vieja, 10º47'N 85º19'W 2700', 18.iii.1982, R.J. Kirby & S.A. Speight BM1982-260. NMB: (1) one specimen: Turrialba, Costa Rica, C. argus ab. famularis St. J. Bechyné det. 1951; (2) one specimen: Bebedero, Costa Rica, Reimoser; (3) one specimen: Turrialba, Costa Rica. NMCZ: (1) two specimens: Costa Rica, Coll. Achard Mus. Pragense [one with: Calligrapha argus St. ]. NMNH: (1) one specimen: Costa Rica, F. Nevermann, 17.iii.30, Turrucares 600 m, Pacifikseite, unter loser Rinde vorsteckt, Calligrapha argus Stål J. Gómez-Zurita det. 2011; (2) one specimen: Costa Rica, F. Nevermann, Guanacaste, Mai 1932, P. Assmann leg.; (3) one specimen: Costa Rica, F. Nevermann, 17.i.30, Sta Maria (Tilerán) 800 m, W.S. Thomas leg.; (4) one specimen: Costa Rica, Guanacaste, Hacienda Palo Verde, 6–15 July 1976, J.C. Solomon coll., 122, Calligrapha argus Stål J. Gómez- Zurita det. 2011; (5) one specimen: Costa Rica, Cent. Am. ix.1.1906, F. Monrós Collection 1959; (6) two specimens: Finca La Pacífica, 7 km NW Cañas, Guanacaste Prov., Costa Rica, DHJ, Jun, 17–24, 1969 [one with: Calligrapha argus Stål J. Gómez-Zurita det. 2011]; (7) one specimen: Costa Rica, Rio Corobici, Las Canas, 15 June 1967, Flint & Ortiz; (8) one specimen: Costa Rica, Guanacaste Prov., Lomas Barbudel Res., 13 July 1989, leg. David G. Furth, Calligrapha argus Stål J. Gómez-Zurita det. 2011. TAMUIC: (1) one specimen: Costa Rica, Guanacaste, 29 km WSW Cañas, Sta. OTS Paloverde, 10º21’N 85º21’W, 30 June 1976, H.A. Hespenheide, Calligrapha argus Stål J. Gómez-Zurita det. 2011.

EL SALVADOR

EGRC: (1) one specimen: El Salvador, Ahuachapan, Apaneca, 4500 ft, 7–12.ix.2002, D. Marqua, Calligrapha argus Stål J. Gómez-Zurita det. 2011. FSCA: (1) one specimen: El Salvador, Tamanique 1000 m, 5.xii.1971, S. & L. Steinhauser, Calligrapha argus Stål J. Gómez-Zurita det. 2011. NMNH: (1) one specimen: El Salvador, Vol. Conchagua, 27–29 May 1958, O.L. Cartwright, Calligrapha argus Stål J. Gómez-Zurita det. 2011; (2) two specimens: No. 444-2806, 5.vii.54, Metapan, M.S.V.; (3) two specimens: No. 444-7813, 15.vi.54, La Unión, M.S.V.

GUATEMALA

EGRC: (1) one specimen: Guatemala, Guat. City, Las Hamacas Tr. Pk., 5–8.viii.1979, Thomas & Case, Calligrapha argus St. det. Daccordi ’81. FSCA: (1) one specimen: Guatemala, Baja Veracruz, 6–9 km E Purulhá 5000’, 15–24.iv.1990, E. Giesbert, Calligrapha argus Stål J. Gómez-Zurita det. 2011. MCZ: (1) three specimens: Chacoj, Vera Paz, Champion, Ex Godman and Salvin [one with: Calligrapha argus Stål ]; (2) one specimen: Capetillo, Guatemala, C. Champion, 1st Jacoby Coll.; (3) one specimen: Chacoj, Vera Paz, Champion, 1st Jacoby Coll.; (4) one specimen: Panzos, Vera Paz, Conradt, Jacoby 2nd; (5) one specimen: Teleman, Vera Paz, Champion, Jacoby 2nd Coll.; (6) one specimen: Chacoj, Vera Paz, Champion, Jacoby 2nd Coll. MfN: (1) two specimens: Chacoj, Vera Paz, Champion, 96424 [one with: Calligrapha argus St. ]; (2) one specimen: Guatemala, [illegible]. MTJM: (1) one specimen: Guatemala, Suchitepequez Dept., Finca Los Tarrales, ca. 8 km N of Patulul, 14º32.37’N 91º08.17’W, 760 m, 4 June 2005, R.S. Zack, Calligrapha argus Stål J. Gómez-Zurita det. 2009. NHM: (1) one specimen: El Reposo, 800ft., Champion, Godman-Salvin Coll., Biol. Centr.-Amer.; (2) four specimens: Chacoj, Vera Paz, Champion, Godman-Salvin Coll., Biol. Centr.-Amer.; (3) two specimens: S. Geronimo, Guatemala, Champion, Godman-Salvin Coll., Biol. Centr.-Amer.; (4) one specimen: Cerro Zunil, 4000ft., Champion, Godman- Salvin Coll., Biol. Centr.-Amer.; (5) one specimen: Capetillo, Guatemala, Rodríguez, Biol. Centr. Amer. Collection; (6) one specimen: Checoj, Vera Paz, Champion, Biol. Centr. Amer. Collection; (7) one specimen: Cerro Zunil, 4–5000ft., Champion, Godman-Salvin Coll., Biol. Centr.-Amer. NMNH: (1) one specimen: Guatemala, Chicacao, 13.vii.1949, T.H. Farr, Calligrapha argus Stål J. Gómez-Zurita det. 2011; (2) two specimens: Chacoj, Vera Paz, Champion [one with: Calligrapha argus Stål ]; (3) one specimen: Guatemala, San Pedro, 11823, 9-13- ’60, 22083, in Orchidaceae, Calligrapha sp. DMW. NRM: (1) two specimens: Chacoj, Vera Paz, Champion; (2) one specimen: Cerro Zunil, 4–5000ft. Champion, Calligrapha argus Stål ; (3) one specimen: Cerro Zunil, 4–5000ft. Champion; (4) one specimen: Capetillo, Guatemala, G.C. Champion. OUMNH: (1) nine specimens: Chacoj, Vera Paz, Champion, B.C.A. duplicates pres. 1909 by F.D. Godman Cat. No. 84; (2) one specimen: Panajachel, 5000 ft, Champion, B.C.A. duplicates pres. 1909 by F.D. Godman Cat. No. 84, Calligrapha argus Stål J. Gómez-Zurita det. 2011; (3) one specimen: Panzos, Vera Paz, Conradt, B.C.A. duplicates pres. 1909 by F.D. Godman Cat. No. 84, Calligrapha argus Stål J. Gómez-Zurita det. 2011; (4) one specimen: Capetillo, Guatemala, G.C. Champion, B.C.A. duplicates pres. 1909 by F.D. Godman Cat. No. 84, Calligrapha argus Stål J. Gómez-Zurita det. 2011; (5) six specimens: Cerro Zunil, [four with: 4–5000 ft; two with: 4000 ft], Champion, B.C.A. duplicates pres. 1909 by F.D. Godman Cat. No. 84, Calligrapha argus Stål J. Gómez-Zurita det. 2011. ZSM: (1) sixteen specimens: Guatemala, Suchitepequez, Patului, [one with: 15.8.83; three with: 1.7.83; two with: 10.8.83; three with: 16.8.83; seven with: 10.7.83], A. Poll, Calligrapha argus Stål J. Gómez-Zurita det. 2011.

HONDURAS

EGRC: (1) one specimen: Honduras, Cortes Hill, behind San Pedro Sula, 17.ix.1984, C.W. O’Brien. FSCA: (1) two specimens: Honduras, Intibuca, 18 km W La Esperanza, 3.xii.1995, F.W. Skillman; (2) one specimen: Honduras, Choluteca, Cerro Guanacuare, 4.vi.1993, F.W. Skillman, Jr.; (3) one specimen: Honduras, Francisco Morazon Zamorano, 4.x.1993, R. Turnbow, Calligrapha argus Stal Det. E.G. Riley ’93; (4) one specimen: Honduras, Francisco, Morazon, 25.5 km SSW Talanga, 3.vi.1993, M.C. Thomas, Calligrapha argus Stål J. Gómez- Zurita det. 2011; (5) four specimens: Honduras, El Paraíso, Yuscarán, R. Turnbow, Calligrapha argus St. Det. E.G. Riley ’03 [three with: 14.vii.2001; one with: 21.vii.2001]; (6) one specimen: Honduras, El Paraíso, Yuscarán 840 m, 4–8.viii.1992, L. Stange & C. Porter, degradated wet forest, Calligrapha argus Stål J. Gómez-Zurita det. 2011. MCZ: (1) one specimen: Sn. P. Sula, Hond. NHM: (1) one specimen: Honduras, 45-123; (2) two specimens: Hond. [illegible], Baly Coll. NMNH: (1) one specimen: Hond., El P. vic Yuscaran, 18 May 1995, J.E. Wappes, Calligrapha argus (Stål) det. J.E. Wappes 2002 . TAMUIC: (1) one specimen: X0534460, Honduras, Comayagua, 30 mi E Tegucigalpa, 31.vii.1982, Robert W. Jones, Calligrapha argus Stål J. Gómez-Zurita det. 2011.

MEXICO

EGRC: (1) three specimens: Mexico, Tamaulipas, Bocatoma, 7 km SSE Gomez Farias, 27–28.v.1979, E.G. Riley, on Guazuma ulmifolia (Sterculiaceae) ; (2) two specimens: Mexico, Tamaulipas, Bocatoma, 7 km SSE Gomez Farias, [one with: 25–30.iii.1978; one with: 19–23.v.1979], E.G. Riley; (3) four specimens: Mexico, Tamaulipas, Bocatoma, 6 mi S Gomez Farias, 25–30.iii.1978 [one with: F. Breitenbach; one with: Marlin E. Rice Coll.]; (4) six specimens: Mexico, San Luis de Potosí, El Salto, at the falls, 26.v.1979 [three with: Marlin E. Rice Coll.; three with: E.G. Riley]; (5) one specimen: Mexico, Oaxaca, 37 mi NW Tehuantepec, 1500 ft, 12.viii.1974, O’Brien & Marshall; (6) one specimen: Mexico, Chiapas, 16 km W Ocozocautla, Aguacera, 16.vi, D.B. Thomas; (7) one specimen: Mexico, Chiapas, Oaxaca border, Hwy90, 28.vii.1988, D.B. and A.M. Thomas; (8) one specimen: Mexico, Chiapas, El Aguacero, 25.vi.1989, P. Lago, J. Burne & D. Thomas; (9) three specimens: Mexico, Chiapas, El Aguacero, D.B. & A.M. Thomas [one with: 14.vi.1988; two with: 29.vii.1988, Calligrapha argus Stål J. Gómez-Zurita det. 2011]; (10) one specimen: Mexico, Chiapas, Simojovel, 20.x.1988, Thomas, Calligrapha argus Stål J. Gómez-Zurita det. 2011. FSCA: (1) one specimen: Mexico, Chiapas, 16 km W Ocozocoautla, 20.vi.1987, E. Giesbert, Calligrapha argus St. Det. J. Watts 1993 ; (2) one specimen: Mexico, San Luis de Potosí, El Salto Falls, 3.vii.1990, J.K. Adams; (3) one specimen: Mexico, San Luis de Potosi, El Salto Falls, 29.vi.1968, H.V. Weems, Jr.; (4) one specimen: Mexico, Tamaulipas, 1–2 mi E N Morelos, 2.vi.1982, J.E. Wappes; (5) sixteen specimens: Mexico, Veracruz, 16.5 mi S Catemaco, Hwy180, 17–25.vi.1985, Askevold & Heffern [one with: Calligrapha argus Stal det. M. Daccordi 1986 ]; (6) two specimens: Mexico, Veracruz, 5.2 mi S Catemaco, Hwy180, 17.vi.1985, Askevold & Heffern [one with: Calligrapha argus Stal det. I. Askevold 1986 ]; (7) two specimens: Mexico, Veracruz, 1.5 mi NE Tatahuicapan, 25.vi.1985, Askevold & Heffern; (8) two specimens: Mexico, Veracruz, vic. of El Salto de Eyipantla, 15 km S San Andres Tuxtla, 15–28.vi.1985, Askevold & Heffern; (9) three specimens: Mexico, Veracruz, Lake Catemaco, 7.vii.1965, G.H. Nelson, sweeping roadside vegetation, Calligrapha argus St. det. A.J. Gilbert ’87; (10) one specimen: Mexico, Veracruz, Hwy145, 6 mi S Tinaja, 25.vii.1972, Calligrapha argus Stål J. Gómez-Zurita det. 2011; (11) one specimen: Mexico, San Luis de Potosí, 7.4 km E Tancuayalab, 24.vii.1988, R. Turbow, Calligrapha argus St. det. E. G. Riley ’93; (12) one specimen: Mexico, San Luis de Potosí, El Salto, 19.vi.1973, H.W. Weems, Calligrapha argus Stål J. Gómez-Zurita det. 2011; (13) one specimen: Mexico, Chiapas, Hwy190, Chiapas de Corsa, 3.viii.2001, W. Opitz, Calligrapha argus Stål J. Gómez- Zurita det. 2011; (14) one specimen: Mexico, Tamaulipas, Mun. de Aldana, Rancho Nuevo, Barra Coma, 11–28.vi.1979, D.F. Gicca, Calligrapha argus Stål J. Gómez-Zurita det. 2011; (15) one specimen: Mexico, Chiapas, 9 km N Arriaga, 15.x.1988, E. Giesbert, Calligrapha argus Stål J. Gómez-Zurita det. 2011. MCZ: (1) one specimen: Mex.; (2) one specimen: Mex., A.P. Morse Coll., Calligrapha argus Stål J. Gómez-Zurita det. 2010; (3) one specimen: Tuxtla, San Andrés, Mexico, Sallé Coll., 1st Jacoby Coll.; (4) one specimen: Yucatan, contigua Chev. [illegible], 1st Jacoby Coll.; (5) two specimens: Tapachula, Chiapas, Höge, Jacoby 2nd Coll. MfN: (1) one specimen: 29787, argus Stål , Mexico, Ehrenb.; (2) two specimens: Atoyac; (3) one specimen: Tapachula; (4) one specimen: Sierra de Zungolica [Zongolica], 583, scalaris Lec. ; (5) one specimen: Tupataro, 21; (6) two specimens: Mexico, Flohr. NHM: (1) one specimen: Temax, N. Yucatan, Gaumer; (2) one specimen: Tepachula, Chiapas, Höge, Godman-Salvin Coll., Biol. Centr.-Amer.; (3) one specimen: Juquila, Mexico, Sallé Coll., Calligrapha multipunctata Stål apud Sallé, Godman-Salvin Coll., Biol. Centr.-Amer.; (4) one specimen: Veracruz, Mexico, Sallé Coll., 643, Sp. figured, Godman-Salvin Coll., Biol. Centr.-Amer.; (5) one specimen: Minas Viejas, Mexico, Dr. Palmer, Godman-Salvin Coll., Biol. Centr.-Amer.; (6) one specimen: Almolonga, Mexico, Hoege, Godman- Salvin Coll., Biol. Centr.-Amer.; (7) one specimen: C. Campeche, Mexico, Sallé Coll., Calligrapha argus Stål apud Sallé, Godman-Salvin Coll., Biol. Centr.-Amer. [one with: 644]; (8) one specimen: Calligrapha , Mexico, Pascoe Coll. 9360; (9) one specimen: Chiapas, Mexico, Sallé Coll., Calligrapha famularis Stål apud Sallé, Godman-Salvin Coll., Biol. Centr.-Amer.; (10) one specimen: Tuxtla, Mexico, Sallé Coll., Calligrapha famularis Stål apud Sallé, Biol. Centr. Amer. Collection; (11) one specimen: Etla, Mexico, Sallé Coll., Calligrapha famularis Stål apud Sallé, Godman-Salvin Coll., Biol. Centr.-Amer.; (12) one specimen: Atoyac, Vera Cruz, May, H.H.S., Calligrapha argus, Godman-Salvin Coll., Biol. Centr. -Amer.; (13) one specimen: Tapachula, Chiapas, Höge, Godman-Salvin Coll., Biol. Centr.-Amer. NMCZ: (1) one specimen: Mexique, famularis Stål, Coll. Achard Mus. Pragense , Calligrapha argus Stål J. Gómez-Zurita det. 2009; (2) three specimens: Mexique, Coll. Achard Mus. Pragense. NMNH: (1) one specimen: Mexico, at Miami, 6.xi.1968, H.L. Rubin, with Tillandsia sp. plants 68-27132, Calligrapha sp. nr. labyrinthica Stal d. R.E. White, Calligrapha argus Stål J. Gómez-Zurita det. 2011; (2) one specimen: Chilpancingo, Guerrero, Mex. ii.11.46, Laredo Tex. on orchid plant 46-1591, Calligrapha sp. perhaps notatipennis Stal HSB’46, 46-1591; (3) one specimen: Sierra de Durango, gift F.C. Bowditch, Calligrapha argus Stål J. Gómez- Zurita det. 2011; (4) one specimen: Calligrapha sp. Det. S.W. Lingafelter 1997, Ex Mexico on Chamaedorea , III- 17-97, Calligrapha argus Stål J. Gómez-Zurita det. 2003; (5) one specimen: Temax, N. Yucatan, Gaumer, Calligrapha multipustulata Stål ; (6) one specimen: Tampico Mex., Lar. Tex. 60649, Mar. 13-60, 8556, bromeliad, Calligrapha sp. prob. argus Stål DMW; (7) one specimen: Mexico Dist. Federal 31.i.1949, Laredo, F. Monrós Collection 1959, Calligrapha argus Stål F. Monrós det. 1954; (8) one specimen: Mexique, gift of F.C. Bowditch; (9) one specimen: Chiapas, 19 km N Arriaga, June 23, 1987, J.E. Wappes; (10) one specimen: Tamazunchale, S.L. Potosí, Mex., vi.19.41, col. & pres. by Henry S. Dybas, F. Monrós Collection 1959, Calligrapha argus Stål J. Gómez-Zurita det. 2011; (11) one specimen: Chiapas, Mex., L. Hotzen ’19, Pacific Slope Cordilleras 800–1000 m, Calligrapha argus Stål J. Gómez-Zurita det. 2011; (12) five specimens: Mexico, Chiapas, 33.7 mi N Huixtla 6000’, ii.26 –27.1966, in bromeliads, George E. Ball & D.R. Whitehead collectors; (13) one specimen: Mexico, Veracruz, 22.0 mi E Jalapa, Rte. 140, 1000’, iii.9.66, in bromeliads, George E. Ball & D.R. Whitehead collectors; (14) one specimen: Mexico, Ver., Cordoba, vii.28.1964, A.B. Lau, Calligrapha argus Stål J. Gómez-Zurita det. 2011; (15) one specimen: Veracruz, Mexico, Sallé Coll.; (16) two specimens: Mexico, Vera Cruz, 30.i.67, Gonzalez et al. #Brownsville 90583, Lot.#67-4686, Tillandsia sp., Calligrapha sp. Det. R.E. White; (17) one specimen: Mexico, Vera Cruz, 30.i.67, Gonzalez et al. #Brownsville 90587, Lot.#67-4690, Bromeliads sp.; (18) two specimens: Atoyac, vii.14.41, Vera Cruz, Mex., Col. by H.S. Dybas, F. Monrós Collection 1959 [one with: Calligrapha argus Stål J. Gómez-Zurita det. 2011; (19) one specimen: Mexico, Campeche, Edizna, 150’, iv. 22.1966 in bromeliads, George E. Ball & D.R. Whitehead collectors; (20) two specimens: Mexico, at Laredo 30.xi.67, Warren & Bauman on bromeliad plnts [one with: Calligrapha sp. nr. diversa Stal d. R. White]; (21) one specimen: Mexico, Laredo 29.iii.1947, F. Monrós Collection 1959; (22) one specimen: Laredo, Tex., ex. S.L. Potosi, Mex., iii.19.56, Watt, Tillandsia plant 56-3676, Calligrapha sp. DMW56; (23) one specimen: Mexico, Huichihuayan, S.L.P., iii.27.58, Fouts colr., on Tillandsia plant, Laredo Tex. 58517, 58-6331, Calligrapha sp. det. D.M. Weisman, Calligrapha argus Stål J. Gómez-Zurita det. 2011; (24) one specimen: Mexico, SLP at Brownsville, 11.16.66, Heinrich & Burgess w. bromeliads 66-30323, Calligrapha sp. d. R. White; (25) one specimen: Tamazanchule, S.L.P., Mex., ix.23.46, E.S. Ross collector, Calligrapha argus Stål J. Gómez-Zurita det. 2011; (26) one specimen: Texas/ Mexico, 4 I 1972, D. Johnston, San Antonio 2914, Florist whse., Lot. 72-2882, Calligrapha sp. det. R. White, Calligrapha argus Stål J. Gómez-Zurita det. 2009. TAMUIC: (1) two specimens: X0533764 and X0547082, Mexico, Chiapas, [Aguaceros], 40 km W Tuxtla Gutiérrez, 21.vi.1987, at light, William F. Chamberlain, Calligrapha argus Stål J. Gómez-Zurita det. 2011; (2) one specimen: X0534994, Mexico, Chiapas, [Aguaceros], 40 km W Tuxtla Gutiérrez, 20.vi.1987, William F. Chamberlain, Calligrapha argus Stål J. Gómez-Zurita det. 2011; (3) four specimens: [X0533940, X0538585, X0540150 and X0544532], Mexico, Veracruz, Panuco, 27.viii.1962, William F. Chamberlain, Calligrapha argus Stål J. Gómez-Zurita det. 2011; (4) one specimen: X0537162, Mexico, Veracruz, San Andrés Tuxtla, 15.vii.1964, Merril H. Sweet; (5) two specimens: [X0539034 and X0541214], Mexico, Veracruz, Los Tuxtlas Range, Laguna Encantada, 18.viii.1960, D.C. Robinson; (6) two specimens: [X0542378 and X0546254], Mexico, Veracruz, Catemaco, 22.viii.1967, Horace R. Burke & J. Hafernik; (7) one specimen: X0547338, Mexico, Tabasco, 20 km W Cardenas, Campo Exp. CSAT, 22.vii.1980, Joseph C. Schaffner, Janis Weaver & Timothy P. Friedlander; (8) one specimen: X0552475, Mexico, Chiapas, 9.5 mi NW Pichucalco, 3.viii.1980, Joseph C. Schaffner, Jannis Weaver & Timothy P. Friedlander.

NICARAGUA

IBE-JGZ: (1) two specimens: [IBE-JGZ-C105, IBE-JGZ-C106], Nicaragua, Rivas, Finca Sierra Serena UTM 16P-657865-1242291, 21 September 2009, J.-M. Maes leg.; (2) one specimen: IBE-JGZ-C232, Nicaragua, Rivas, Finca Guadalupe, 10–12/07/2010, J.-M. Maes leg.; (3) one specimen: IBE-JGZ-C233, Nicaragua, Estelí, P. T. P. Miraflor-Moropotente, Finca Las Flores, 13.192742N 86.331123W 870m, Abril 2010, J.-M. Maes leg.; (4) two specimens: [IBE-JGZ-C240, IBE-JGZ-C254], Nicaragua, Rivas, Finca Isla Vista, 6–7/07/2010, J.-M. Maes leg.; (5) four specimens: [IBE-JGZ-C257, IBE-JGZ-C258, IBE-JGZ-C259, IBE-JGZ-C260], Nicaragua, Rivas, Cárdenas, Finca Sierra Serena, 14/07/2010, J.-M. Maes leg.; (6) one specimen: IBE-JGZ-2181, Nicaragua, Estelí, P. T. P. Miraflor-Moropotente, Finca Bartolo UTM 16P-0573112-1459212 993m, Junio 2010, A. del Socorro & B. Nimia leg.; (7) one specimen: IBE-JGZ-2183, Nicaragua, Montelimar, Reserva Natura, parcela 1, 8/10/2010, J.-M. Maes leg. MCZ: (1) one specimen: Chontales, Nicaragua, 1st Jacoby Coll. NHM: (1) one specimen: Chontales, Nicaragua, T. Belt, Godman-Salvin Coll., Biol. Centr.-Amer.; (2) one specimen: Janson, Nicarag., Chont.s, Fry Coll. 1905.100. NMNH: (1) one specimen: La Calera, Nic., 15/viii/55, trampa luminica, C. Espinoza R. Bradley 57-70, Calligrapha argus Stål J. Gómez-Zurita det. 2011; (2) one specimen: Nicaragua, Pachomil, vi.71, J. Maldonado C.; (3) one specimen: Consequina Slope, Nicaragua, vii.7.32, M. Willows Jr. collector, Templeton Crocker Exp. 1932, Calligrapha argus Stål J. Gómez-Zurita det. 2011.

PANAMA

EGRC: (1) one specimen: Panama, Canal Zone, La Pita Signal Station rd., 8.vi.1976, E.G. Riley; (2) three specimens: Panama, Canal Zone, Fort Kobbe, [one with: 28.vi.1970; one with: 6.vi.1976; one with: 15.vi.1976], E.G. Riley; (3) two specimens: Panama, Canal Zone, Diablo Heights, [one with: 30.iv.1971; one with: 15.vi.1970], E.G. Riley; (4) one specimen: Panama, Canal Zone, Gamboa, 11.vi.1976, E.G. Riley; (5) one specimen: Panama, Canal Zone, Achiote rd, 10 km SW Gatun, 12.vi.1976, E.G. Riley; (6) one specimen: Panama, C.Z., Cardenas Village, 22.v.1980, E.G. Riley & D. LeDoux; (7) two specimens: Panama, C.Z., K-1 rd., near Fort Kobbe, 19.v.1980, E.G. Riley & D. LeDoux; (8) one specimen: Panama, C.Z., Gamboa, 18.v.1980, E.G. Riley & D. LeDoux. FSCA: (1) one specimen: Panama, Summit Gardens, Canal Zone, 25.v.1977, R. Davis, Jr.; (2) one specimen: Panama, Colon, vic. Ft. Sherman, 24.ii.1999, J.E. Wappes, Calligrapha argus Stal Det. E.G. Riley ’01; (3) one specimen: Panama, Panama, 10 km N El Llano, 15.ii.1987, E. Giesbert, Calligrapha argus Stål J. Gómez- Zurita det. 2011; (4) two specimens: Panama, Canal Zone, Ft. Gulick, H.J. Harlan, E.J. Gerberg Collection, Calligrapha argus Stål J. Gómez-Zurita det. 2011 [one with: June 1979; one with: 17.v.1979]. IBE-JGZ: (1) three specimens: Panama, Isla Colón, ii/95, lgt. D. Buzga. MCZ: (1) four specimens: Panama, Panama [one with: Cal. argus St.]; (2) one specimen: Panama, Bella Vista, 2.vii.1924, N. Banks, Calligrapha argus Stål J. Gómez-Zurita det. 2010; (3) one specimen: V. de Chiriqui, 25–4000 ft, Champion, Jacoby 2nd Coll.; (4) four specimens: Panama, Panama. NHM: (1) one specimen: V. de Chiriqui, 25–4000ft, Champion, Godman-Salvin Coll., Biol. Centr.- Amer.; (2) one specimen: San Lorenzo, Panama, Champion, Godman-Salvin Coll., Biol. Centr.-Amer. NMNH: (1) one specimen: Panama, Chiriqui Prv., 4 km E Boqueti, 16 May 1996, Wappes, Huether & Morris; (2) one specimen: April 23, 1911, Panama Pan., E.A. Schwarz collector; (3) one specimen: Calebra, C.Z., Pan., Collection F. Knab; (4) one specimen: Canal Zone 100 m, 5.0 mi NW Gamboa 9º10’00’’N 79º45’00’’W, sample 5–7, 6 Apr 1976, Montgomery & Lubin coll., canopy fogging experiment in Luehea seemanni, Pyrethrin fog; (5) one specimen: Paraiso CZ, Pan, May 9, 11, E.A. Shwarz, Calligrapha argus Stål J. Gómez-Zurita det. 2011; (6) one specimen: Pan., Panama Prov., Cerro Campana 17.v.1999, Morris/Wappes, Calligrapha argus Stål J. Gómez-Zurita det. 2011. TAMUIC: (1) one specimen: X0535600, Panamá, Chiriquí, vic. Hornito, 14–18.v.1996, James E. Wappes, Huether & Morris, Calligrapha argus Stål J. Gómez-Zurita det. 2011; (2) one specimen: X0536682, Panama, Cerro Campana, 7.i.1994, James E. Wappes, Calligrapha argus Stål J. Gómez-Zurita det. 2011.

VENEZUELA

NMCZ: (1) one specimen: Caracas, Sallé, simillima var . Stal Caracas, famularis Stal Mex., Coll. Achard Mus. Pragense , Calligrapha argus Stål J. Gómez-Zurita det. 2009; (2) two specimens: Vénézuéla, L. Laglaize, Déc. 1896, Coll. Achard Mus. Pragense [one with: Calligrapha argus Stål J. Gómez-Zurita det. 2009]. NMNH: (1) two specimens: Venezuela, Trujillo, La Puerta 1700 mts., 1.iv.1947, H.E. Box. NRM: (1) one specimen: Venezuela, Tarnier, … Deg.

UNKNOWN SOURCE

MfN: (1) five specimens: [no data]. NHM: (1) one specimen: [illegible], Baly Coll.; (2) one specimen: C., 160, Baly Coll.; (3) two specimens: E. Coll. Laferté, 67-56; (4) one specimen: 67-56; (5) one specimen: E. Coll. Thomson, 67-56; (6) one specimen: Baly Coll. NMCZ: (1) one specimen: Coll. Achard Mus. Pragense; (2) one specimen: Calligrapha famularis St., Coll. Achard Mus. Pragense , Calligrapha argus Stål J. Gómez-Zurita det. 2009. NMNH: (1) one specimen: Amer. Mer., gift of F.C. Bowditch; (2) one specimen: Norway, 17.ix.1971, L. Beikman, aircraft Lot 72-1433, Calligrapha sp. d. R. White. NRM: (1) one specimen: 342, Calligrapha argus .

Variation. My interpretation of C. argus is perhaps more consistent with considering this taxon a species complex, given that it exhibits a certain degree of variation that would allow further subdividing it into more taxa. Nevertheless, in general most specimens conform to the type in several characters, including divergent basal ends of subsutural stripe, free humeral marking, free and round spot enclosed by humeral lunule, divergent basal end and roundish divergent apical end of arcuate band, well-delimited and orderly arranged additional spots on disc, and brick-reddish tint to dark body parts. However, some specimens tentatively placed within this group with large midlateral spot, laterally confluent with margin of elytron, deviate in one or several characters from the type and are reminiscent of C. multipustulata . These include a narrow connection of humeral marking with dark basal margin of elytron; an irregular spot enclosed by humeral lunule narrowly connected to it; an humeral lunule with its apical half separated from the rest; a broken, irregular apex of arcuate band; certain variation in the number of additional spots due to confluence/fragmentation (although they are generally ten), the innermost spot usually confluent with external concavity of arcuate band; and/or the pigmentation can be much darker, blackish bronzy, and the contrast between pale background and paler area surrounding markings can be exacerbated. In my opinion, and judging from the illustration in Biologia Centrali Americana for C. famularis, Jacoby (1882) could have interpreted this taxon as some of these atypical forms of C. argus , in particular the large darker forms similar to C. multipustulata typical from Nicaragua. This interpretation would have grounded his proposition of C. famularis as an aberration of C. argus . The specimens from Venezuela held at MNPC have somewhat larger markings, with the spot enclosed by humeral lunule large, ovoid, elongated anteriad, and the spot on the concavity of arcuate band large, slightly oblique and narrowly connected to the latter marking, and they are also reminiscent of the configuration observed in C. famularis .

Figures

FIGURE 1. Schematic representations of the typical apical ends of flagella in the penis of species in the Calligrapha argus group (a), in the C. dislocata and C. philadelphica groups (d), in the C. polyspila group (c), and in the C. labyrinthica group (d).

FIGURE 2. Distinguishing traits of major Calligrapha lineages: (a) connate claws of Calligrapha (Zygospila) mexicana Jacoby; (b) divergent claws of C. labyrinthica Stål; (c) unimpressed hypomera in C. percheroni Stål; (d) hypomera with impressed suture in C. argus Stål (white arrowhead); (e) habitus of C. californica Brown showing striped elytral pattern; (f) habitus of C. amelia Knab showing spotted elytra; (g) habitus of C. vigintimaculata (Chevrolat) showing reversed contrasting pattern of pale markings on dark background; (h) habitus of C. serpentina (Rogers) showing typical pattern of dark markings on pale background; (i) left elytron of C. sponsa Stål with a single spot enclosed by humeral lunule (white arrowhead); (j) left elytron of C. aladina Bechyné with arrangement of two spots enclosed by humeral lunule (white arrowheads); (k) left elytron of C. polyspila (Germar) with subhumeral (white arrowhead) and midlateral (black arrowhead) spots; and (l) left elytron of C. multipustulata Stål only with midlateral spot (black arrowhead).

FIGURE 9. Penis in ventral, lateral and dorsal view of apex of the species Calligrapha ancoralis Stål (a, d), C. anabelae sp. nov. (b, e), C. catarinae sp. nov. (c, f), C. ramulifera Stål (g, h), and C. argus Stål (i, j). Scale bar = 0.5 mm.

FIGURE 10. Geographic distributions of Calligrapha argus Stål (black circles) and C. ramulifera Stål (white circles).