Calligrapha argus

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 : 7-9

Key to species in the group of Calligrapha argus

The group of C. argus currently comprises fourteen species, including two new species described in this work, and one taxon proposed as subspecies. All species can be distinguished by their particular and relatively constant arrangements and shape of elytral spots, as well as coloration in one case. I paid particular attention to the median lobe of the aedeagus (= penis) in these species, based on the idea that this structure usually provides valuable characters for taxonomic identification in insects (e.g., Sharp & Muir 1912; Shapiro & Porter 1989; Schilthuizen 2003). However, in practice, at the assayed level of inquiry, intraspecific variability for this complex character system overlaps with interspecific variability, particularly among closely related species, where this character would prove more useful. The penis in this species group has few prominent features to seek for diagnostic characters and has a rather constant shape characterized by a slightly flattened, ventrally concave tubular shape, gradually tappering in lateral view and widening in dorsal view at apical third, strangled preapically, before an explanate more or less regularly convex apical border with the characteristic lateral denticles, which always have their tips marking the widest point of the penis in dorsal view. Small differences overlapping between species can be seen in the size and proportions, the degree of curvature of the apical border, the retraction and projection of lateral denticles, or the degree of preapical enlargement, among others. However, the latter may have some important value in the supraspecific systematic arrangement of the species group investigated here, as assessed a posteriori based on covariation with other characters. In particular, in C. argus and close allies the preapical enlargement is more pronounced than in other species, whereby the transversal distance between the basal curvatures of lateral denticles is conspicuously wider than the transversal width of penis medially. The spermatheca of all species examined for this structure were essentially identical except for differences in size, matching the description available for other species ( Gómez-Zurita et al. 2004; Montelongo & Gómez-Zurita 2013); therefore, it has not been investigated systematically in this study.

I determined and compiled label information for 2,135 Calligrapha specimens belonging to this group found in the collections above. This lineage is distributed from the southern parts of the California-Rocky Mountain domain in the Southern Nearctic (two species) to the northern regions of the Caribbean NW South American domain in the Neotropics (three species) following the classifications of Morrone (2006) and Escalante et al. (2010, 2013). However the highest species diversity (ten out of 14 species) is concentrated in the Mexican Transition Zone and the Caribbean Mesoamerican domains, and all the species can be found in the latter. The species in the group can be subdivided in three main subgroups, including these of C. diversa , without midlateral spot ( Fig. 12 ), C. multipustulata , generally with numerous additional spots on disc of elytra and spot enclosed by humeral lunule large, occupying most of the space between humeral lunule and subsutural stripe ( Fig. 3 ), and C. argus (Fig. 6).

1 Midlateral spot present ( Figs 2 k, 2l)..................................................................... 2

- Midlateral spot absent................................................................................. 11

2 With many dark spots on disc, generally occupying most of elytral surface ( Fig. 3 a); spot enclosed by humeral lunule large, ovoid, generally occupying most of the available space, emargined anteriorly or encircling a small pale area ( Figs 3 a, 3b, 3c); penis moderately enlarged preapically, narrowed before lateral denticles to average width of median lobe ( Fig. 4 )......... 3

- Fewer, smaller and generally less contiguous spots on disc (Figs 6, 12); spot enclosed by humeral lunule roundish, smaller, less confined; penis conspicuously enlarged preapically, with base of lateral denticles more distant than with of penis at middle ( Fig 9 )................................................................................................. 6

3 Midlateral spot narrow, elongated, free from margin or narrowly reaching it medially through dark suffusions ( Figs 3 a, 3b, 3c).................................................................................................... 4

- Midlateral spot broadly confluent with elytral margin ( Figs 2 l, 3d).............................. C. multipustulata Stål

4 Large spots on disc ( Figs 3 b, 3c, 3d)....................................................................... 5

- Numerous small irregular spots on disc ( Fig. 3 a)............................................... C. consputa Stål

5 Very narrow longitudinal midlateral spot ( Fig. 3 b)....................................... C. geographica s. str. Stål

- Elongated but medially thickened midlateral spot, narrowly confluent with margin of elytron ( Fig. 3 c)................................................................................... C. geographica ssp. famularis Stål stat. nov.

6 Midlateral spot free; basal end of arcuate band blunt (Fig. 6d); legs bicoloured..................... C. fulvitarsis Jacoby

- Midlateral spot confluent with margin; basal end of arcuate band divergent, separated from subsutural stripe (Figs 6a, 6b, 6c, 6e, 6f); legs unicoloured................................................................................ 7

7 Spot enclosed by humeral lunule largely confluent laterally with humeral lunule; apex of arcuate band long, hook-like (Fig. 6f)....................................................................................... C. ancoralis Stål

- Spot enclosed by humeral lunule free; apex of arcuate band short, round (Figs 6a, 6b, 6c, 6e).......................... 8

8 Spot enclosed by humeral lunule and spots on disc of elytra very small; humeral marking short, ending posteriorly well ahead of basal end of arcuate band; apex of arcuate band of irregular profile; blackish colouration to dark parts (Fig. 6a)............................................................................................... C. ramulifera Stål

- Spot enclosed by humeral lunule and discal spots larger; humeral marking longer, ending at level with basal end of arcuate band; apex of arcuate band rounded...................................................................... 9

9 Reddish colouration to dark parts (Fig. 6b)........................................................ C. argus Stål

- Blackish colouration to dark parts...................................................................... 10

10 Base of subsutural stripe nearly entirely confluent with sutural stripe (Fig. 6c).................... C. catarinae sp. nov.

- Base of subsutural stripe divergent and separated for relatively long distance from sutural stripe (Figs 6e, 11)................................................................................................. C. anabelae sp. nov.

11 Arcuate band completely separated from subsutural stripe ( Fig. 12 b)............................. C. elegantula Jacoby

- Arcuate band largely confluent with subsutural stripe........................................................ 12

12 Apical spot on elytra present. Eight or nine free round markings on elytra...................................... 13

- Apical spot on elytra absent. Maculation much reduced..................................................... 14

13 Base of sutural marking broad, blunt; maculation heavier ( Fig. 12 e)................................. C. simillima Stål

- Base of sutural marking narrower, with slightly divergent end of subsutural stripe ( Fig. 12 d)............... C. diversa Stål

14 Five small free round markings on apical half of elytra ( Fig. 12 c)...................................... C. sylvia Stål

- Additional markings on elytra missing ( Fig. 12 a)............................................... C. aeneopicta Stål

Figures

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 3. Dorsal and lateral habitus of Holotype of Calligrapha consputa Stål (a), Lectotype of C. geographica ssp. geographica Stål (b), Lectotype of C. geographica ssp. famularis Stål (c), and Lectotype of C. multipustulata Stål (d). Scale bar = 5.0 mm.

FIGURE 4. Penis in ventral, lateral and dorsal view of apex of the species Calligrapha multipustulata Stål (a, d), C. geographica Stål (b, e) and C. consputa Stål (c, f). Scale bar = 0.5 mm.

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 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.