Calligrapha aladina Bechyné, 1954

Gómez-Zurita, Jesús, 2018, Systematic revision of Central American Calligrapha Chevrolat of the subgenus Erythrographa subgen. nov. (Coleoptera: Chrysomelidae, Chrysomelinae), Zootaxa 4531 (1), pp. 1-58 : 52-56

Calligrapha aladina Bechyné, 1954

( Figs 10d , 11g, 11h , 12)

Calligrapha labyrinthica (pars): Jacoby, 1882. Biol. Centr.-Amer., vol. vi, pt. 1, p. 199.

Calligrapha aladina Bechyné, 1954 . Ent. Arb. Mus. Frey 5, p. 584.

Calligrapha aladina: Wilcox, 1975 . Checklist Chryomelidae, p. 66.

Calligrapha aladina: Gómez-Zurita, 2015 . Zootaxa 3922, p. 8.

Calligrapha aladina: Benítez-García et al., 2017 . Rev. Mex. Biodiv. 88, p. 339.

While being hired at the Museum G. Frey in Munich, Jan Bechyné proposed a new taxon to describe Calligrapha aladina Bechyné, 1954 , based on a specimen externally matching C. labyrinthica Stål in many respects. In fact, the specimen that Jacoby (1882) chose to illustrate C. labyrinthica in his contribution to the Biologia Centrali- Americana is almost certainly C. aladina . This taxon actually represents a bit of a conundrum, which may be hopefully solved in part here. In his work, J. Bechyné referred his new species as unique, also in its colouration, and even reminiscent of Leptinotarsa (" eine auffallend gefärbte, isoliert stehende Art, die eher an eine Leptinotarsa erinnert, [...]"; p. 585 in Bechyné 1954). This is a surprising statement, considering that he knew at least two highly similar species, C. notatipennis Stål and C. suboculata Stål , based on determinations of specimens in the Georg Frey collection as early as 1951. He must have also known C. labyrinthica Stål at least the same year that the description of C. aladina Bechyné was published, based on one specimen determined by him in 1954 at the Royal Belgian Institute of Natural Sciences. However, the explicit mention of the uniqueness of C. aladina , the lack of character-based diagnosis between his new species and the practically indistinguishable C. labyrinthica Stål (diagnoses that often appeared in his works, also in the one discussed here, for example tabulating the differences between C. multiplagata Achard and his new taxon C. melasomina Bechyné, 1954 ; p. 586 in Bechyné 1954), and the actual absence of any reference to C. labyrinthica make me suspect that when he proposed the new taxon, he was not aware of Stål's species. If true, this memory lapse and the lack of an explicit explanation of the reasons that compelled the author to propose a new species, could be responsible for the later confusion with C. labyrinthica or that C. aladina was not recognised in most collections, let alone the possibility of a synonymy. These identification problems are expected, considering that Bechyné's taxon might represent a phenotypic extreme in the evolutionary lineage of C. labyrinthica , which in this case would consist of specimens with fainter punctation on disc of pronotum, elongate scutellum, together with softer, orangish instead of reddish coloration of most body parts, including thickened and highly confluent elytral markings which, as a result, show stronger contrast with their darkened margins. The specimens consistent with the characters of C. aladina , even if in some cases with reasonable doubts about their actual belonging to C. labyrinthica , are in fact geographically concordant, if fully sympatric and very often syntopic on the northwestern range of typical C. labyrinthica Stål ( Fig. 14c ). Considering the geographic concordance of phenotypic diversity of otherwise highly similar organisms, it may be tempting to treat Bechyné's taxon with subspecific rank. However, full sympatry and distribution of morphological traits for both taxa which does not seem to be clinal, together with the existence of diagnostic differences, chiefly the shape of penis, argue in favor of retaining the younger taxon with specific rank.

Holotype, by original designation: Jalisco, Méx. Guadalajara / TYPE Calligrapha aladina m. J. Bechyné det., 1954 ( NMB). The left maxillary palp seems atrophied or scarred. The specimen is a slightly teneral male and the genitalia, which I dissected is poorly sclerotized, but mounted alongside the specimen.

Habitus ( Fig. 10d ). Length: 8.11 mm, width: 5.20 mm. Body elliptical, moderately convex. Head, labrum, six basal antennomeres, mandibles, maxillary palpomeres, pronotum, scutellum, ventral surfaces, legs, epipleura and dark elytral markings orange; pale areas of elytra dark yellow; five apical antennomeres, apex of mandibles, apical border of last maxillary palpomeres and fine margin of dark elytral markings dark brown.

Head large, deeply inserted into pronotum, finely microreticulate, with uniformly distributed fine punctures on frons; area above eyes slightly depressed and uneven, with short supraocular furrow; antennal calli smooth with small setose punctures externally near eyes; frontal suture finely impressed and apically joining equally finely impressed bisinuous frontoclypeal suture. Clypeus short, transverse, microreticulate and unpunctured on disc, with fine punctures, smaller than frontal punctures at sides with short golden hairs. Labrum transverse, rather regularly curved at sides and weakly emarginate at apex, with more or less transverse row of five golden setae at each side. Mandibles thick, strong, relatively short, surpassing apex of labrum by less than labrum length, with strong punctures and long golden setae at sides. Maxillary palpi long; basal palpomere club-shaped, longer than wide at apex; second palpomere with short cylindrical base, slightly narrower than apex of previous segment, elbowed and strongly expanded towards apex; last maxillary palpomere wide and transverse, as wide at base as apex of previous palpomere, expanded laterally on external border and nearly straight at apex. Antennae slender, moderately clavate, reaching slightly beyond humeri; antennomeres 1–6 and base of seventh antennomere smooth, with sparse recumbent hairs; apical 2/3 of seventh antennomere and antennomeres 8–11 rugose and densely covered by fine pale yellowish pubescence; scape relatively short, with posterior border nearly flat and anterior border strongly dilated; pedicel as short club, slightly longer than half scape; third antennomere as long as scape, slender, feebly widened gradually towards apex; fourth antennomere 0.7x as long as third antennomere, feebly clavate; fifth antennomere subequal to and imperceptibly shorter than fourth; sixth antennomere slightly longer than pedicel, strongly widened as base of antennal club; antennomeres 6–10 progressively longer, longer than wide at apex (eighth antennomere, W/L = 0.78), with seventh antennomere about as long as fifth, eighth as long as fourth, ninth 1.1x longer than fourth and tenth nearly 1.2x longer than fourth; apical antennomere longer than scape. Pronotum transverse (W/L = 2.04), convex, wider near basal, obtuse angles, with sides finely margined, weakly converging anteriorly and gently curved at apical half toward strongly produced anterior angles; basal border unmargined, weakly bisinuous with posteriorly produced median lobe occupying 2/3 of base; anterior border feebly convex and finely margined; surface of pronotum very finely microreticulate, nearly smooth, unpunctured on disc except for sparse small punctures at basal third, with strong, deep punctures at sides and row of elongate deep punctures confluent with basal border except medially. Hypomera slightly convex, smooth, glossy, with few transverse impressions in basal depressed area; hypomeral suture wide and deep, parallel to pronotal border and slightly divergent preapically following basal contour of anterior calli. Prosternum short, transverse, with anterior border concave and margined; surface finely shagreened, depressed with imprecise punctures and posteriorly recumbent golden setae before procoxae and strong, individual punctures at base of process; prosternal process narrow, spatulate, with strong punctures. Mesanepisterna small, triangular, punctured. Mesepimera finely shagreened, unpunctured. Metanepisterna finely microreticulate, with dense strong punctures, slightly elongate close to posterior end. Disc of metaventrite glossy, unpunctured, slightly depressed, with fine longitudinal median suture disappearing on anterior process; sides finely shagreened, with small punctures. Scutellum long, about 1.5x longer than wide at base, finely microreticulate, unpunctured. Elytra long, wider than base of pronotum, with humeri relatively prominent, sides gently curved, widest at middle and regularly curved at apical third towards sutural angle, narrowly margined; surface finely microreticulate with relatively dense, very fine punctures, ordered, deeper and larger around and within dark markings, including premarginal line of punctures at apical half of elytra, diverging from border of elytron at level with midlateral spot and becoming progressively confused towards subhumeral area; scutellar row of 14–15 punctures present, slightly confused basally. Elytral markings consisting of: (i) sutural stripe entire, confluent with base of elytron and broadly surrounding scutellum basally, progressively narrowed towards apex, reaching sutural angle; (ii) subsutural stripe completely confluent laterally with sutural stripe from anterior position relative to apex of scutellum, gradually narrowing posteriorly except briefly at apical declivity of elytron, and nearly reaching sutural angle, expanding preapically and fusing with distal end of dark marginal stripe laterally; (iii) arcuate band entire, externally concave, and broadly fused laterally with subsutural stripe, with anterior free divergent lobe twice as long as posterior lobe; (iv) humeral spot large, long, from near base to nearly middle of elytron, entirely fused internally with humeral lunule, externally convex and projecting short lobe towards midlateral spot; (v) humeral lunule large, regularly curved, confluent with base of elytron and reaching slightly beyond middle of elytron, fused with slightly over basal 1/3 of arcuate band; (vi) basal spot enclosed by humeral lunule small, free, round, at level with basal end of subsutural stripe and slightly closer to lunule, almost completely dark brown; posterior spot enclosed by humeral lunule much larger, roundish, at level and broadly fused laterally with middle concavity of lunule; (vii) subhumeral spot large, irregular, broadly fused laterally with lateral convexity of humeral spot and with narrow dark lateral margin of elytron; (viii) midlateral spot large, elliptic, broadly fused laterally with lateral margin of elytron and expanded posteriorly as narrow marginal dark stripe between margin of elytron and premarginal row of punctures, nearly reaching sutural angle, and fusing with apical enlargement of subsutural stripe; (ix) spot of apical declivity large, elliptic elongate, broadly fused laterally with slight preapical widening of subsutural stripe; (x) apical spot large, roundish, narrowly confluent posteriorly with dark marginal stripe; (xi) additional spots on posterior lateral declivity of elytron fused as a single large irregular marking with peripheral lobes in positions recognizable as these of free spots in other species, and separated from surrounding features. Epipleura finely shagreened, unpunctured, weakly slanted ventrally, visible at basal 2/3 of elytra from lateral view. Femora long, enlarged at middle, glossy, with sparse fine punctures and short appressed golden hairs. Tibiae as long as corresponding femora, nearly straight, gradually expanded apically, with gradually widened furrow on apical half of external border; glossy, scarcely punctured at base, finely rugose and densely pubescent at apex, with fringe of golden setae along internal border. Tarsi shorter than corresponding tibiae; onychia with simple claws, divergent at acute angle. First abdominal ventrite shorter than metaventrite at middle, glossy with sparse minute puctures and fine recumbent translucent hairs on disc and finely shagreened, with denser, stronger punctures at sides, behind metacoxae; ventrites 2–4 progressively narrower, glossy, with sparse fine punctures and fine, translucent recumbent hairs; fifth ventrite longer than second, glossy, finely punctured, with apical border margined and bisinuous. Penis ( Figs 11g, 11h ) relatively slender compared to allied species, with sides nearly parallel in ventral view, weakly enlarged postmedially, before operculum, regularly curved dorsally in lateral view, tapering to apex at apical quarter; basal flap of operculum transverse, with anterior border concave; apical border feebly curved, nearly flat at middle, laterally projecting acute teeth with maximal width of penis between tips; distal end of flagellum projecting two sclerotized sinuous arms ventrally.

Distribution. Mexican endemic species distributed in the Pacific slopes of the Sierra Madre Occidental, from Sinaloa and western Durango, to the States of Morelos and Guerrero (Fig. 12).

Other material examined (43 specimens).

MEXICO.

EGRC: (1) four specimens: Mexico, Guerrero, 22 km N Taxco, 26.vii.1987, R. Turnbow coll., Calligrapha labyrinthica Stål J. Gómez-Zurita det. 2011; FSCA: (1) one specimen: Mexico, State of Mexico, 2 mi NE Ixtapan de la Sal, 18.vii.1974, R.L. Mangan & D.S. Chandler coll., Calligrapha aladina Bechyné J. Gómez-Zurita det. 2011; (2) one specimen: Mexico, Guerrero, Hwy95, 23 km N of Xochipala Rd., 24.vii.1972, G.H. Nelson coll., on Mimosa sp., Calligrapha aladina Bechyné J. Gómez-Zurita det. 2011; (3) two specimens: Mexico, Guerrero, 22 km N Taxco, 27.vii.1987, R. Turbow coll., Calligrapha aladina Bechyné J. Gómez-Zurita det. 2011. HNHM: (1) one specimen: Mexico, Calligrapha labyrinthica St. MCZ: (1) one specimen: Mexico, Jacoby 2 nd Coll.; (2) three specimens: Mexico from Durango to the Pacific, Calligrapha aladina Bech. J. Gómez-Zurita det. 2010; (3) two specimens: Guadalajara, Mexico, Dr. Buller, Calligrapha aladina Bech. J. Gómez-Zurita det. 2010; (4) one specimen: Jalisco, Dr. B. MfN: (1) one specimen: 379, Canelas, Calligrapha aladina Bechy. det. Daccordi '79; (2) two specimens: Pilar, Durango; (3) one specimen: Canelas; (4) one specimen: Canelas, Zool. Mus. Berlin, Calligrapha aladina Bechyné det. M. Daccordi 1992 [penis dissected]. NHM: (1) one specimen: Canelas, comparato con il typus MD'80, Brit. Mus. 1984-58, Calligrapha aladina Bech. , det. Daccordi 80; (2) two specimens: Cuernavaca, Mexico, Sallé Coll., Godman-Salvin Coll., Biol. Centr.-Amer. [one with: 662]; (3) one specimen: Huetamo, Michoacan, Höge, Godman-Salvin Coll., Biol. Centr.-Amer.; (4) one specimen: Ventanas, Durango, Höge, Godman-Salvin Coll., Biol. Centr.-Amer.; (5) one specimen: Mazatlan, Mexico, Sallé Coll., 660, Godman-Salvin Coll., Biol. Centr.-Amer.; (6) one specimen: Mazatlan, Mexico, Sallé Coll., Sp. figured, Godman- Salvin Coll., Biol. Centr.-Amer. NMCZ: (1) one specimen: Mexique, Boucard, Coll. Achard Mus. Pragense; (2) one specimen: Sierra de Durango, Coll. Achard Mus. Pragense. NMNH: (1) one specimen: Mexico, Jalisco, 14 km SW Cocula, 28-IX-91, Morris, Wappes, Giesbert, Calligrapha sp. Det. E.G. Riley ’01, Calligrapha aladina Bechyné J. Gómez-Zurita det. 2003. TAMUIC: (1) two specimens: [X0534064, X0534169], Mexico, Guerrero, 15 mi SW Chichihualco, 15.vii.1984, L.E. Carroll, J.C. Schaffner & T.P. Friedlander coll., Calligrapha aladina Bech. J. Gómez-Zurita det. 2011; (2) five specimens: [X0534204, X0534334, X0534712, X0535531, X0538046], Mexico, Michoacán, 10 mi S Uruapan, 29.vii.1988, Ferreira & Schaffner coll., Calligrapha aladina Bech. J. Gómez-Zurita det. 2011. ZSM: (1) one specimen: El Cora, Tepic, Ad. Lüdecke, Polyspila labyrinthica Stål, Staatssammlung München 1975 Erwerb Coll. Machatschke.

UNKNOWN SOURCE. MCZ: (1) one specimen: Jacoby 2nd Coll., Calligrapha aladina Bech. J. Gómez-

Zurita det. 2010; (2) one specimen: 1596bis. NMCZ: (1) one specimen: labyrinthica J. Achard det., Coll. Achard

Mus. Pragense; (2) one specimen: Coll. Achard Mus. Pragense. Variation. The variation displayed by C. aladina Bechyné is less pronounced than found in its likely sister C. labyrinthica Stål , and it can be summarised as different degrees of darkening of margins of elytral markings, the presence in a reduced number of specimens of the narrow pale sutural area also present more often in C. labyrinthica , the large spot enclosed by humeral lunule possibly touching the subsutural stripe or more rarely connected with the smaller, basal spot enclosed by humeral lunule, and finally the confluence of additional spots, which in some cases may show one of the lobes detached or can be laterally confluent with the arcuate band.

Figures

FIGURE 10. Dorsal and lateral views of the lectotype of Calligrapha suboculata Stål (a), lectotype of C. notatipennis Stål (b), lectotype of C. labyrinthica Stål (c), and the holotype of C. aladina Bechyné (d).

FIGURE 11. Ventral and lateral views and dorsal view of apex of the penises of Calligrapha suboculata Stål (a, b), C. notatipennis Stål (c, d), C. labyrinthica Stål (e, f) and C. aladina Bechyné (g, h). Scale bars = 1.0 mm.

FIGURE 14. Distribution of Calligrapha labyrinthica Stål, separating localities with specimens with completely dark and with narrowly pale suture.