Calligrapha discrepans (Achard, 1923), 2021

Gómez-Zurita, Jesús, 2021, Systematic revision of Calligrapha s. str. Chevrolat (Coleoptera: Chrysomelidae, Chrysomelinae), Zootaxa 4953 (1), pp. 1-111 : 53-58

Calligrapha discrepans (Achard, 1923) stat. rev.

( Figs 7c–d , 8d–e , 10 )

Calligrapha serpentina (pars): Jacoby, 1891, Biol. Centr.-Amer., 1, suppl., p. 246.

Polyspila serpentina var. discrepans Achard, 1923b , Fragm. Entom., 5, p. 69.

Calligrapha serpentina v. discrepans: Blackwelder, 1946 , Checklist Col. Amer., 4, p. 674.

Calligrapha serpentina subsp. temaxensis Bechyné, 1952 , Ent. Arb. Mus. Frey, 3, p. 4. syn. nov.

Calligrapha serpentina ab. discrepans: Scherer, 1959 , Ent. Arb. Mus. Frey, 10, p. 609.

Calligrapha serpentina: Flowers & Janzen, 1997 , Florida Entom. 80, p. 344.

Calligrapha serpentina: Flowers, 2004 , Rev. Biol. Trop., 52, p. 80.

Calligrapha serpentina var. discrepans: Montelongo & Gómez-Zurita, 2014 , Zool. Scr., 43, p. 608.

Calligrapha serpentina ssp. discrepans: Maes & Gómez-Zurita, 2016 , Rev. Nic. Ent., 76, p. 29.

Calligrapha serpentina temaxensis: Merckx et al., 2018 , Biodiv. Inf., 13, p. 45.

Julien Achard (1923b) described what he considered a variety of Calligrapha serpentina (Rogers) based on an indeterminate number of specimens from Sierra de Durango ( Mexico) with obvious similarities with that species, but presenting striking differences particularly in the maculae in the posterior half of elytra. The type material of many of the Chrysomelidae described by Achard is part of the entomology collection of the National Museum of the Czech Republic, in Prague. Indeed, this collection hosts three syntype specimens labeled by the author as types, of which the one explicitly mentioning “serpentina” in the label is selected here as the lectotype. In the course of the current study, I compiled data showing that the typical C. serpentina is parapatric with the range of specimens fitting the type of C. serpentina var. discrepans (Achard) , which present a more meridional distribution. Moreover, phylogenetic analyses of Calligrapha based on several molecular markers and including representatives of both phenotypes ( Montelongo & Gómez-Zurita 2014; Gómez-Zurita, unpubl.) resulted in a distant phylogenetic relationship between them, compatible with differences measured between other species of Calligrapha . Based on these new sources of data and the fixed and remarkable morphological differences between both taxa, I propose upgrading Achard’s taxon to species rank, as Calligrapha discrepans (Achard) stat. rev.

Lectotype ( Fig. 7c ), by present designation: Sierra de Durango [green label], Type [red label], serpentina v. discrepans m. J. Achard det. TYPE, Coll. Achard Mus. Pragense ( NMCZ).

Paralectotypes, by present designation: (1) Sierra de Durango [green label], Type [red label], discrepans n. var. J. Achard det. TYPE, Coll. Achard Mus. Pragense ( NMCZ); (2) Sierra de Durango [green label], Type [red label], v. discrepans m. J. Achard det. TYPE, Coll. Achard Mus. Pragense ( NMCZ) .

Description. Body elliptic, elongate with sides subparallel, moderately convex and slightly depressed dorsally. Head, pronotum, scutellum and apex of mandibles dark brown with bronzy metallic reflections; mandibles and ventral surfaces dark reddish brown; five apical antennomeres, labrum, dark markings on elytra and epipleura brown, with slight metallic reflections on epipleura and some elytral markings basally; basal antennomeres, mouth appendages, legs, including trochanters, and fifth abdominal ventrite orange; pale background of elytra creamy yellow. Length: 10.3 mm; width: 5.9 mm.

Head broad, relatively short and deeply inserted into prothorax; surface finely microreticulate, nearly unpunctured on frons, with free round punctures posteriorly and behind eyes, slightly elongate smaller punctures anteriorly and near fronto-clypeal suture, sometimes confluent as large, deep impressions; antennal calli unpunctured except anteriorly at sides; frontal suture finely impressed, anteriorly convergent with broadly bisinuous frontoclypeal suture; supraocular furrows deep from above antennal calli to upper eye margins. Eyes relatively small and dorso-ventrally elongate, entire and finely faceted. Clypeus transverse, slightly depressed posteriorly, finely microreticulate, with isolated punctures on disc, more densely punctured at sides, with row of medially convergent golden setae anteriorly. Labrum small, short and transverse, curved at sides and anterior angles, weakly emarginate at apex, punctured and with long medially convergent golden setae anteriorly and at sides. Mandibles large, robust, much longer than clypeus, shortly concave basally and divergent at sides before strong median curvature; surface with strong punctures and long golden setae. Maxillary palpi elongate, slender; last palpomere about as long as wide, subtrapezoidal, with sides feebly curved and apex obliquely truncate; previous palpomere narrow and cylindrical at base, elbowed and strongly enlarged apically, broader than last palpomere; basal palpomere slender, longer than second, clavate, obliquely cut at apex. Antennae short but slender, reaching humeri, clavate beyond sixth antennomere; scape about twice as long as wide, dilated anteriorly and flattened posteriorly; antennomeres 2–6 slender, clavate, glossy, with few scattered setae; pedicel about half as long as scape, slightly clavate; third antennomere more than 1.5x longer than pedicel; fourth antennomere slightly longer and more slender than pedicel; fifth antennomere about as long as pedicel and sixth antennomere shorter than pedicel; antennomeres 7–11 enlarged, with granulose surface and dense fine short pubescence; seventh antennomere as long as pedicel and antennomeres 8–10 subequal and as long as fourth antennomere; eighth antennomere slightly more than 0.8x as wide as long; last antennomere as long as scape, widest in apical 2/3 with blunt pointed apex.

Pronotum transverse, slightly over 1.7x as wide at base as long at middle, weakly convex, following body convexity in sagittal plane, and more convex than elytra transversally, particularly toward anterior angles, with abrupt vertical premarginal declivity; sides straight and parallel in basal half and smoothly curved in apical half toward strongly produced anterior angles, sides finely margined with margin thicker at anterior angles; anterior angles acute, with large, deep trichobothrium at angle; anterior border of pronotum straight behind frons and curved at sides behind eyes toward anterior angles, finely margined throughout; basal border strongly convex posteriorly, unmargined and with posterior angles obtuse; surface finely microreticulate with relatively dense small shallow punctation on disc, becoming larger, deeper and occasionally confluent fossette-like punctures at sides, particularly basally to anterior angles, aligned laterally in premarginal area and along and confluent with basal border laterally. Hypomera triangular, convex on disc and depressed at margins, surface glossy, unpunctured, wrinkled at lateral and basal depressions; hypomeral suture impressed, slightly wavy and subparallel to pronotal margin at base and divergent apically. Prosternum relatively long, with anterior transverse area slightly shorter than prosternal process; anterior border concave, raised and finely margined; transverse area before procoxae finely microgranulose, unpunctured in anterior half and with large, deep, confluent punctures posteriorly with posteriorly recumbent long fine setae; prosternal process convex, as narrow between procoxae as apical maxillary palpomere, gradually expanding posteriorly and nearly straight at apex, carinated longitudinally with strong punctures and semierect long setae at sides. Scutellum lancet-shaped, 1.1x longer than wide basally, with blunt round apex, finely alutaceous, unpunctured. Mesoventrite wider than apex of prosternal process, raised at middle and impressed at sides, glossy, with large deep round punctures posteriorly. Mesepimera finely microreticulate, with unpunctured uneven surface. Mesanepisterna microreticulate and entirely covered by large deep round punctures. Metanepisterna long, with raised anterior and external smooth broad margins, surface of disc microreticulate with dense, relatively small punctures anteriorly and rugose-punctate posteriorly. Metaventrite as long as ensemble prosternum and mesoventrite, with wider than long anterior process between mesocoxae thickly margined and slightly concave posterior border between metacoxae; disc smooth, glossy with fine transverse scratches, longitudinally impressed in posterior half medially and with sparse fine punctures with short recumbent fine yellowish setae; sides finely shagreened and more strongly and deeply punctured in anterior half.

Elytra elongate, about 0.8x as long as body, wider at base than base of pronotum, with broad humeri, sides subparallel, broadest at middle, and regularly round in apical third; surface finely microreticulate, with sparse dot-like, shallow dark punctures, and slightly bigger and deeper punctures around dark markings; punctures confused in scutellar area with scutellar row of about 15 punctures; regular premarginal line of tight punctures from subhumeral spot to sutural angle and confused in humeral area. Dark markings of elytra ( Fig. 7c ) consisting of: (i) sutural stripe thin, narrowly surrounding apical half of scutellum without reaching base of elytra, gradually tapering at apical declivity of elytron toward sutural angle; (ii) subsutural stripe narrow, with basal end at level with apex of scutellum, slightly curved and divergent from sutural stripe at base until end of scutellar row of punctures, fused with sutural stripe elsewhere, slightly broadened at apical declivity of elytron and gradually thinning apically without reaching sutural angle; (iii) arcuate band entire, weakly concave externally, long and as thin as and subparallel to subsutural stripe, separated by distance greater than width of stripes, with ends slightly widened, free in left elytron and narrowly connected to humeral marking in right elytron of type; (iv) humeral spot thin, long and weakly curved, completely fused laterally with humeral lunule, with basal end distant from base of elytron; (v) humeral lunule broadly separated from base of elytron, sinuous and about twice as long as humeral spot with convexity of apical half toward suture and reaching middle of elytron, fused externally to basal end of large irregular postmedian discal spot in lateral declivity of elytron; (vi) small spot at internal declivity of humerus, confluent with base of elytron; (vii) spot enclosed by humeral lunule elongate, half as long as humeral spot, with narrower basal end at level with basal ends of humeral spot and lunule and base of subsutural stripe, slightly closer to humeral lunule than to subsutural stripe, and longitudinally aligned with curvature of arcuate band; (viii) subhumeral spot small, shortly elliptical, free but narrowly distant from margin of elytron, not delimited by punctures and irregularly crossed by five punctures of premarginal line of punctures; (ix) midlateral spot about twice as big as subhumeral spot, elliptic, elongate, covering some eleven punctures of premarginal line of punctures, and fused externally with margin of elytron; (x) spot of apical declivity elongate, as long as midlateral spot, with narrower apex obliquely directed toward sutural angle, free and at level with apical enlargement of subsutural stripe; (xi) apical spot smaller than spot of apical declivity, elongate, placed between spot of apical declivity and margin of elytron, slightly closer to margin, with narrower apical end directed to sutural angle; (xii) pale median area external to humeral lunule at lateral declivity of elytron with two small round spots transversally arranged, and one minute spot aligned with outer one and with midlateral spot; and pale area behind large irregular spot connected to humeral marking at apicolateral declivity of elytron with two irregular spots. Epipleura slanted ventrally, sinuous and visible in lateral view, wide at base, conspicuously narrowed at middle and gradually thinning posteriorly toward sutural angle; very finely microsculptured, unpunctured and with many transverse scratches, particularly in apical half, with fringe of short erect setae apically.

Femora slender, widened medially, with smooth surface and scattered punctures with short, recumbent pale yellow setae. Pro- and mesotibiae weakly curved and metatibiae straight, slender and as long as their corresponding femora, gradually widened toward apex and externally furrowed in apical half with furrow widening toward tarsal insertion; longitudinal edges finely carinated; surface glossy, sparsely but more deeply punctured than femora, with short, semierect setae, and densely pubescent with short golden setae at apex ventrally. Tarsi slender, slightly shorter than corresponding tibia; first tarsomere longer than wide at apex, shorter than tarsomeres 2–3 together; third tarsomere wider than first, weakly bilobed; onychia with acute ventral tooth at apex, and claws long, simple, acute and weakly divergent at base. First abdominal ventrite as long as metaventrite, with broad subtrapezoidal anterior intermetacoxal process; second abdominal ventrite nearly half as long as first, fourth ventrite 1/3 as long as first, and third ventrite of intermediate length; fifth abdominal ventrite as long as second, with bisinuous and finely margined apical border and weak median convex lobe; abdominal ventrites glossy, with fine transverse scratches on disc, slightly wrinkled at sides, sparsely and finely punctured, with short fine translucent setae. Penis ( Fig. 8d ) slender, tubular, nearly as thick as wide, moderately curved ventrally, gradually tapering dorsally in apical fifth; sides almost parallel, slightly narrowed around distal end of gonopore, with apical border curved, projecting a short convex lobe medially and short acute, slightly recurved teeth laterally, with distance between tips not much wider than average width of penis; gonopore large, ovoid, bound at base dorsally by slightly bilobed flap, longitudinally membranous at middle; distal end of flagellum largely expanded, with big central orifice and membranous at periphery.

Material examined (56 specimens).

COSTA RICA

NMNH: (1) one specimen*, AF105, Costa Rica, Guanacaste Prov., D.H. Janzen et al., Calligrapha sp. det. D.R. Whitehead, Calligrapha serpentina ssp. temaxensis Bech. J. Gómez-Zurita det. 2011.

MEXICO

FSCA: (1) one specimen, Mexico, Jalisco, 5 km S Tuxcacuesco, 29.ix.1991, E. Giesbert leg., Calligrapha serpentina discrepans Ach. J. Gómez-Zurita det. 2011. MCZ: (1) one specimen, Mexico, Morelos, Tepoztlán, 14.vi.1993, leg. D. Furth, Calligrapha serpentina discrepans (Achard) J. Gómez-Zurita det. 2010; (2) one specimen, Ventanas, Mex., 2000 ft., Forrer, 1st Jacoby Coll., Calligrapha serpentina discrepans (Ach.) J. Gómez-Zurita det. 2010; (3) one specimen, Tuxtla, San Andrés, Mexico, Sallé Coll., 1st Jacoby Coll., Calligrapha serpentina discrepans (Ach.) J. Gómez-Zurita det. 2010; (4) one specimen, Iguala, Guerrero, Höge, Jacoby 2nd Coll., Calligrapha serpentina discrepans (Ach.) J. Gómez-Zurita det. 2010; (5) one specimen, Guanajuato, Mexico, Calligrapha serpentina discrepans (Ach.) J. Gómez-Zurita det. 2010; (6) two specimens, Tuxpan, Jalisco, Mex., 6.ix.1903, J.F. McClendon, Calligrapha serpentina discrepans (Ach.) J. Gómez-Zurita det. 2010; (7) one specimen, Río Balsas, Gro., Mex., Wickham, not typical, ant. club dark, no spot on thighs, Calligrapha serpentina discrepans (Ach.) J. Gómez-Zurita det. 2010; (8) four specimens*, Temax, N. Yucatán, Gaumer [three with: Jacoby 2nd Coll., Calligrapha serpentina temaxensis Bech. J. Gómez-Zurita det. 2010]; (9) two specimens*, Yucatán, Calligrapha serpentina temaxensis Bech. J. Gómez-Zurita det. 2010. MfN: (1) one specimen*, Temax, N. Yucatan, Gaumer, 96376. NHM: (1) two specimens,Acapulco, Guerrero, Höge, Godman-Salvin Coll., Biol. Centr.-Amer., Calligrapha serpentina discrepans (Achard) J. Gómez-Zurita det. 2009; (2) one specimen, E. Coll. Thomson, Mexiq.?, 149, Calligrapha serpentina discrepans (Achard) J. Gómez-Zurita det. 2009; (3) five specimens*, Temax, N. Yucatan, Gaumer, Godman-Salvin Coll.-Biol. Centr.-Amer. [one with: Calligrapha serpentina var.]; (4) one specimen*, Tuxtla, San Andrés, Mexico, Sallé Coll., 655, Godman-Salvin Coll., Biol. Centr.-Amer. NHMB: (1) two specimens [SYNTYPES of Calligrapha serpentina subsp. temaxensis Bechyné, 1952 ]*, Temax, N. Yucatan, Gaumer, Type [pink label], [one with: HOLOTYPE; one with: ALLOTYPE] C. serpentina ssp. temaxensis m. J. Bechyné det., 1951; (2) one specimen, Jalapa, Mexico, Polyspila serpentina v. discrepans J. Achard det.; (3) two specimens, Acapulco, Mex., x.1958, leg. G. Frey. NMCZ: (1) three specimens, Ventanas [Villa Corona], Durango, Höge, Coll. Achard Mus. Pragense; (2) one specimen, Acapulco, Guerrero, Höge, Coll. Achard Mus. Pragense. NMNH: (1) seven specimens, Jalisco, Mex., La Venta, Aug. 11, 1954, R. Ryckman, C. Christianson, R. Lee collrs. [one with: Calligrapha serpentina ssp. discrepans Achard J. Gómez-Zurita det. 2011]; (2) one specimen, on orchidaceae, Mex., 20.iii.1939, San Francisco, Cal., 16229; (3) one specimen, orchid (on), Panuco, Sin., Mex., 9.vi.1938, Nogales 26741; (4) one specimen*, Temax, N. Yucatan, Gaumer, Calligrapha serpentina ssp. temaxensis Bech. F. Monrós det. 1953; (5) one specimen*, Temax, N. Yucatan, Gaumer, Calligrapha serpentina ssp. temaxensis Bech. J. Gómez-Zurita det. 2011. OSAC: (1) one specimen*, Mexico, Jalisco, 60 mi W Guadalajara, 12.vi.1967, on Sida pyramidata , 16 36 - CJ A (I). RBINS: (1) one specimen*, Temax, North Yucatan, Gaumer. ZSM: (1) one specimen, Mexico, Durango, Staatssammlung München 1975 Erwerb Coll. Machatschke, Calligrapha serpentina discrepans Ach. J. Gómez-Zurita det. 2011; (2) one specimen, Mexico, Tapalpa, 25.v.1982, leg. Fittkeu, Calligrapha serpentina discrepans Ach. J. GómezZurita det. 2011; (3) one specimen*, Temax, N. Yucatán, Gaumer, Staatssammlung München 1975 Erwerb Coll. Machatschke.

NICARAGUA

JGZC: (1) two specimens, Nicaragua, Estelí, P.T.P. Miraflor-Moropotente, Finca Las Flores , 13.192570ºN 86.331465ºW, 865 m, iv.2010, J.M. Maes leg.

UNKNOWN

NMNH: (1) two specimens, Wild plant, Nogales, Ariz., Nogales 65088, 1.ix.46, 46-15366 [one with: Calligrapha serpentina ssp. discrepans Ach. J. Gómez-Zurita det. 2003]. MCZ: (1) one specimen*, Townsend M., [illegible], Calligrapha serpentina temaxensis Bech. J. Gómez-Zurita det. 2010.

Variation. In 1952, Jan Bechyné described a new subspecies of C. serpentina (Rogers) from Temax, in Yucatán ( Mexico), based on a pair of specimens that deviated from typical C. serpentina in the elytral markings being brown, instead of metallic blue or purple, and with these markings merged and showing heavier punctation on the disc of elytra, while they were free and more finely punctured in C. serpentina ( Bechyné 1952) . Jacoby (1891) had noted the existence of this variety already and from the same locality, but found no reason to separate it from C. serpentina . However, the rediscovery of this local morph by Bechyné ascribed it subspecific status under C. serpentina ssp. temaxensis Bechyné. The examination of the syntypes of this subspecies showed them to be indistinguishable from the types of C. discrepans , apart from the apparent lack of metallic reflections in dark elytral markings and the legs and antennae of a darker tinge. However, these traits, particulary the coloration of maculae on elytra, have been shown to be polymorphic in C. discrepans , where some individuals have faint or absence of metallic shine on their elytra, maybe related to the maturity of the specimens, but not necessarily, since I have seen mature specimens with markings ranging from brown to dark green throughout the range of the species. Based on these comparisons and new data, as well as the lack of differences in the male genitalia of type specimens from both taxa (compare Fig. 8d with 8e), I am compelled to propose these two taxa to be conspecific, as Polyspila serpentina var. discrepans Achard, 1923 = Calligrapha serpentina subsp. temaxensis Bechyné, 1952 syn. nov. (The specimens consistent with the “temaxensis” type are indicated with an asterisk in the list above.) Apart from slight variations in coloration and development of metallic shine as mentioned, there are other variable features in the species. For example, Höge’s specimens conserved at NHM and NMCZ are smaller than usual, two show large humeral markings laterally confluent with the spot enclosed by humeral lunule and one has the base of subsutural stripe almost entirely confluent with sutural stripe (as it occurs in a specimen intercepted in Nogales, Ariz., at NMNH); these specimens also have the large discal spot more rounded and nearly detached from apex of large humeral marking. One of the specimens from Tuxpan at MCZ also shows the arcuate band completely fused to subsutural stripe. The specimen from Río Balsas in the same collection also has the arcuate band nearly entirely fused in basal 2/3 with subsutural stripe, but this specimen is heavily melanic, most noticeably showing an enlarged humeral marking, connected both to base of arcuate band and to the spot enclosed by humeral lunule, which in turn touches the subsutural stripe. The specimen mentioned from Nogales also has a clear transversal stripe attached to apex of arcuate band, pretty much as in C. serpentina , and instead of small irregular spots at apex of elytra, it has a large triangular spot. In general, the additional spots in different specimens or different elytra of the same individual can be broken in smaller spots or fused among them and with surrounding features.

Distribution. Neotropical endemism gradually replacing C. serpentina to the south, reaching from southern Sierra Madre to Yucatán in Central Mexico and recently found in Nicaragua and Costa Rica as well ( Fig. 10 ). It appears associated mainly to Neotropical mountain and dry forests, and herein we provide with records for the following biomes: Sierra Madre Occidental, Trans-Mexican Volcanic Belt and Central American Pine-Oak Forests, Sierra de los Tuxtlas, Oaxacan Montane Forest, and Balsas, Southern Pacific and Yucatán Dry Forests.

Figures

FIGURE 7. Dorsal and lateral views of the lectotype of Calligrapha serpentina (Rogers) (a), lectotype of C. mexicana Stål (b), lectotype of C. serpentina var. discrepans (Achard) (c), holotype of C. serpentina ssp. temaxensis Bechyné (d), holotype of C. sallei (Achard) (e), and lectotype of C. serpens Bowditch (f). Scale = 5.0 mm.

FIGURE 8. Dorsal (apex), ventral and lateral views of the penises of Calligrapha serpentina (Rogers) (a), C. mexicana Stål (b), C. sallei (Achard) (c), C. discrepans (Achard) (d), and C. serpentina ssp. temaxensis Bechyné (e). Scale = 0.5 mm.

FIGURE 10. Distribution of Calligrapha discrepans (Achard) (white circles) and isolated records of specimens that fit the description of C. serpentina ssp. temaxensis Bechyné (gray circles).