Zootaxa 2683: 45–55 (2010) ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2010 · Magnolia Press ISSN 1175-5334 (online edition)

Diabrotica collicola (Coleoptera: Chrysomelidae), a new species of leaf from and key to species of the virgifera group and relatives

NORA CABRERA1,3 & GUILLERMO CABRERA WALSH2 1División Entomología, Museo de La Plata, Paseo del Bosque, s/n, 1900 La Plata, Argentina. E-mail: [email protected] 2South American Biological Control Laboratory, USDA-ARS, Bolivar 1559, B1686EFA- Hurlingham, Buenos Aires, Argentina. E-mail: [email protected] 3Corresponding author

Abstract

The new species Diabrotica collicola Cabrera & Cabrera Walsh is described and illustrated based on specimens collected from Balcozna, Catamarca Province (Argentina). A full description is provided and includes adding morphological characters of the mouthparts, hind wing venation, binding patch, metendosternite, and details of male and female genitalia previously overlooked for the genus. Diabrotica collicola is recognized by the following characters: general color parrot green with yellowish vittae, genal space less than 1/4 maximum length of the eye, antennomeres 2 and 3 subequal in males, 3 longer in females, both antennomeres together more than length 1/2 of 4 in both sexes, prothoracic and mesothoracic tarsi of males with ventral adhesive patch, internal sac of the median lobe with four sclerites. Differences with similar species D. mutabilis Baly, D. fulvofasciata Jacoby, D. mapiriensis gussi Krysan & Smith and D. porracea Harold are discussed. A key to D. collicola and similar species is provided.

Key words: Chrysomelidae, , Diabrotica collicola, Argentina, systematics

Introduction

The genus Diabrotica Chevrolat comprises 338 species arranged in three groups, as proposed by Smith & Lawrence (1967): Diabrotica fucata group, Diabrotica virgifera group and Diabrotica signifera group. Some of the species do not clearly fit within any of these groups and they may need to be placed in others (Cabrera et al. 2008). Members of this genus are distributed throughout Central and , with the exception of continental Chile and southern Argentina, although this distribution may be widened in the future. Only species of the D. fucata and D. virgifera groups are represented in Argentina and they are widely distributed in the central and northern parts of the country, up to the provinces of Ro Negro and Neuquén (ca. 40ºS) (Cabrera 2001 a, b). So far, only three species in the D. virgifera group have been found in Argentina, whereas the D. fucata group is much better represented (Krysan & Smith 1987; Cabrera 2001 a, b; Cabrera & Cabrera Walsh 2004a). However, most species studied are from the subtropical and temperate plains, while the northwestern highlands of Argentina have rarely been explored for galerucines. This region, covering the Yungas, Puna and Prepuna biogeographical provinces (Cabrera & Willink 1980), is characterized by a series of north - south oriented valleys and large plateaus between 1,000 and 4,000 m in elevation, isolated by arid mountainous chains of 4000 – 6000 m in elevation. Annual rainfall ranges from 50 to 600 mm and occurs almost exclusively during summer. Agriculture in this area is limited to the lower parts of the valleys, fringing the floodplains. It is mostly irrigation dependent and consists of small, traditional polyculture farms. During the years 2001 to 2003, we traveled in this region in search for species of Diabrotica and their natural enemies. The purpose of this paper is to provide a full description of a new species, Diabrotica collicola, including morphological characters of the mouthparts, hind wing venation, binding patch, metendosternite, and some

Accepted by M.L. Chamorro: 25 Oct. 2010; published: 22 Nov. 2010 45 details of male and female genitalia previously overlooked in the genus. A key to D. collicola and similar species is provided.

Material and methods

Adult specimens were collected with hand-held aspirators from pumpkin ( maxima Duchense) and (Zea mays L.), and from cucurbitacin-baited cloths during December and March. In the laboratory, the were fed an artificial diet, and offered maize seedlings wrapped in black cotton cloth to obtain eggs (Cabrera Walsh 2001). Morphological terminology follows Cabrera & Cabrera Walsh (2004b), Konstantinov (1998a and b), and Lingafelter & Konstantinov (2000). Measurements were taken using an eye-piece micrometer on a Wild dissecting microscope at 25X magnification. The range is indicated in millimeters, with the average and standard error in brackets. Measurements and abbreviations used in the text were: eye length (eL), genal length (GL), length of antennomeres (A1-A4), length of pronotum (PL), pronotum width (PW), humeral width (HW), elytral length (EL), and elytral width (EW), as defined by Cabrera & Cabrera Walsh (2004b). Other measurements include interocular distance (OD), measured across the vertex between the eyes; and interantennal distance (AD), the distance between the inner margins of the antennal sockets. Body length was measured from the posterior margin of the eyes to the apex of the longest elytron. Relative proportions of the above-mentioned measurements for AD/OD, GL/eL, PW/PL, HW/PW, and EW/HW were computed. Drawings were made with camera lucida on a Leitz compound microscope and a Wild dissecting microscope. Electron micrographs were taken with a Scanning Electron Microscope (SEM) Jeol SMZ 1500. The holotype and paratypes have been deposited in the collection of the Museo de La Plata (MLP), La Plata, Buenos Aires Province, Argentina. The type label includes the species name, the type status and gender.

Results

Diabrotica collicola Cabrera & Cabrera Walsh, new species (Figs. 1−19)

Diagnosis. Small-sized species, body oval, slightly convex. Coloration parrot green with yellowish vittae. Genal space less than 1/4 maximum length of the eye. Antennomeres 2 and 3 subequal in males, 3 longer in females, together more than 1/2 length of 4. Elytra with two weak sulci, basal inner surface of elytra with single binding patch. Pro- and mesothoracic tarsi of males with ventral adhesive patch. Receptacle of spermatheca subcylindrical. Internal sac of the median lobe with four sclerites. Holotype male. (Fig. 1). Color. Head yellowish brown, one-third of mandibles chestnut colored. Antennomeres chestnut colored, with all surfaces of antennomere 1 green. Disc of pronotum green, tinged with yellowish brown. Each elytron parrot green with two yellowish vittae, one medial, widest at middle area, tinged with brown, the other marginal, both vittae partly or entirely confluent at apex; epipleura yellowish; humeral calli yellowish brown. Scutellum amber. Coxae and basal third of femora yellowish brown, tarsi chestnut colored. Venter with prosternum parrot green, meso-metasternum and abdomen yellowish tinged with brown. Head. Vertex finely and sparsely punctate, depressed above antennal calli; antennal calli oval, elevated over surface of vertex, as wide as diameter of the antennal sockets, supracallinal sulcus and midfrontal sulcus deeply impressed, supra-antennal sulcus barely indicated; antennal sockets adjoining anterior margin of eyes, interantennal space smaller than transverse diameter of eye; frontal ridge moderately raised in lateral view; anterofrontal ridge not separated from frontal ridge. Genal space very small, less than 1/4 the maximum ocular length. Antennae inserted approximately at midline of eyes, extending to middle of elytra; antennomere 2 and 3 short, subequal in length, antennomeres 2+3 together longer than antennomere 4; antennomeres 3−10 elongate, similar in length, antennomere 11 apically acuminate. Clypeus with eight

46 · Zootaxa 2683 © 2010 Magnolia Press CABRERA & WALSH preapical setae. Labrum (Figs. 2−3) approximately rectangular, anterior margin with small mesal notch, a row of six long setae at mid length, three short fine setae close to notch apically, twelve short, thick sensilla on each side. Mandibles (Fig. 4) symmetrical, pyramidal, five-toothed apically, only teeth 3−5 visible on external face; tooth 3 and 4 narrow, acute, tooth 3 more than twice as long as 4; tooth 4 almost two times longer than 5, which is small, blunt at apex; inner margin of mandible with two very short denticles; mola longer than wide. Maxillae (Fig. 5) with cardo widened apically and with eight long setae; basistipes with five long setae situated on latero-external margin; galea and lacinia with a fringe-like pilosity apically, galea surpassing lacinia, subcylindrical, apically wider than base. Maxillary palpi well developed; palpomere 1 subrectangular; palpomere 2 and 3 subconical, the latter longer than 2; palpomere 4 subconical, with narrow base, patch of digitiform sensilla (Fig. 9) on the externo-basal corner, formed by twelve embedded sensilla, evident with higher resolution. Labium with four long setae between bases of palps. Labial palp three-segmented, with palpomere 1 subrectangular; palpomere 2 subcylindrical, more than 2.0 times as long as 1; palpomere 3 subconical with very narrow base.

FIGURE 1. Diabrotica collicola Cabrera & Cabrera Walsh, male, dorsal habitus.

Thorax. Pronotum slightly convex, shiny, evenly, finely punctate, rectangular, 1.18 times wider than long, widest near middle, PW 1.55−1.81mm, posterolateral foveae weakly impressed; anterior and posterior margins almost straight, posterolateral margins slightly expanding anteriorly; one long, thin seta on each anterolateral and posterolateral angle. Prosternum convex; procoxal cavities contiguous, positioned midway between anterior and posterior margins of prosternum; intercoxal prosternal process thin, incomplete, extending about one-half length of procoxae. Mesoscutum and scutellum fused; scutellum triangular, nearly as wide as long, rounded at apex. Mesosternum shorter than metasternum at midline, intercoxal mesosternal process thin, extending to one-half length of mesocoxae; mesocoxal cavities inserted on posterior margin, nearly contiguous, open laterally to mesepimeron. Metanotum (Fig. 8) transverse, wider than long, metanotal apodeme “d intersecting apodeme “c posterior to midpoint of “c. Metendosternite (Figs. 7 and 10 ) with stalk longer than wide; lateral arms, thin, divergent, apically deflexed, mesofurcal-metafurcal tendons poorly

NEW SPECIES OF FROM ARGENTINA Zootaxa 2683 © 2010 Magnolia Press · 47 FIGURES 2−8. Diabrotica collicola Cabrera & Cabrera Walsh (2) Labrum, dorsal view. (3) Labrum, ventral view. (4) Mandible, external face. (5) Maxilla, ventral view. (6) Hind wing. (7) Metendosternite, ventral view. (8) Metanotum. Scale bars= 2−5, 7: 0.1mm; 6, 8: 1 mm. Abbreviations: a, metanotal ridge a; AA, anal anterior vein; b2, metanotal ridge b2; bs, basistipes; c, metanotal ridge c; CuA, cubitoanal vein; CuA 3+4, cubito anal vein 3+4; d, metanotal ridge d; ds, ditistipes; ga, galea; lc, lacinia; mg, median groove; mo, mola; MP 1−2, medial posterior vein 1−2; RA, radial vein; RP, radial posterior vein; re, receptacle; RP-MP2, radial posterior-medial posterior vein 2; SC, subcostal vein; sm, setose membrane; th3, tooth 3; th4, tooth 4; th5, tooth 5. developed, inserted close to the apex of the stalk. Hind wings (Fig. 6) with veins RA, MP, CuA well sclerotized, whereas veins SC, CuA2 and AA scarcely sclerotized. Vein SC connected to RA beyond half its length, radial cell darkly pigmented, well developed, subtriangular; RP-MP2 long, reaching r4; AA unbranched, connected to CuA3+4 less than one-half distance from origin of CuA; CuA2 attached to CuA; cubital anal cell closed, elongate. Elytra with surface densely, irregularly punctate, punctures somewhat

48 · Zootaxa 2683 © 2010 Magnolia Press CABRERA & WALSH coarser than on pronotum; elytra slightly wider than pronotum; humeral calli rounded; greatest width near apical third of elytra, EW/HW 1.10−1.36; two weak elytral sulci present; epipleura subvertical, basally broad, gradually narrowed apically. Basal inner surface of elytra with single binding patch (Figs.13−16) narrowed at apex, with stump-shaped spicules occupying anterior middle part, with sharktooth-shaped spicules, sometimes bidentate, on basal, apical and external borders; surface near basal angle covered with thin, sparse microspicules, sometimes bidentate. Legs with metatibiae longer and slenderer than pro- and mesotibiae, apical margins of meso-and metatibiae each with short tibial spur. Tarsomere 1 of prothoracic legs subrectangular, almost equal in length or slightly longer than tarsomeres 2+3 together; tarsomere 1 of metathoracic legs slender, longer than tarsomeres 2+3 together; tarsal claws bifid. Pro- and mesothoracic legs with ventral adhesive patch covering more than half of protarsus, and approximately one-third of mesotarsus.

FIGURES 9−12. Diabrotica collicola Cabrera & Cabrera Walsh (9) Maxilla, detail of digitiform sensillum. (10) Metendosternite, dorsal view. (11) Median lobe, lateral view. (12) Median lobe, dorsal view. Abbreviations: bf, basal foramen; ds, digitiform sensillum; la, lateral arms; op, orificial plate; os, ostium; st, stalk.

Genitalia. Median lobe (Figs. 11, 12 and 17) evenly curved in lateral view, constricted at about the basal fourth, anteriorly slender, tapering slightly toward apex, scarcely deflexed, apically with acute projection; orificial plate elongate, covering approximately half total length of the median lobe, apically acute, separated from apex by less than one-fourth length of the orifice plate; ostium wide, with pair of well developed triangular lobes attached to side. Internal sac (Fig. 18) with four sclerotized plates. Female. Specimens examined similar in color and sculpturing to the males. Antenomere 3 longer than 2, together 1/2 to 3/4 length of antennomere 4. Pronotum wider than in males. Legs without ventral adhesive patches; tarsomere 1 of prothoracic legs slenderer than in males.

NEW SPECIES OF LEAF BEETLE FROM ARGENTINA Zootaxa 2683 © 2010 Magnolia Press · 49 Genitalia (Fig. 19). Sternite 8 weakly sclerotized; apodeme (tignum) slender, wider posteriorly. Vaginal palpi, slender; apex with 10 setae. Vagina + bursa copulatrix large, undivided, with sclerotized area in the posterior part. Spermathecal duct uncoiled, distal part connecting directly to the receptacle. Receptacle of spermatheca subcylindrical, not noticeably separated from pump; pump curved, with pointed appendage at apex.

FIGURES 13−16. Diabrotica collicola Cabrera & Cabrera Walsh (13) Elytron, detail of binding patch and submarginal ribbon. (14) (E) Detail of anterior area of binding patch, surface covered with stump-shaped spicules and sharktooth- shaped spicules. (15) Detail of posterior area of binding patch covered with sharktooth-shaped spicules (16) Detail of submarginal ribbon along the surface near anterior margin, covered with thin microspicules. Abbreviations: bp, binding patch; shs, sharktooth spicules; sr, submarginal ribbon; sts, stump spicules.

Measurements (3♂♂and 3♀♀) Body length 4.55−5.21 mm (4.98+0.28), eL 0.75−0,82 mm (0.76+0.05), OD 0.58−0.66 (0.61+0.04), GL 0.09−0.13 (0.10+0.02), AD 0.42−0.46 (0.42+0.02), PL 1.28−1.51 mm (139+0.05), PW 1.55−1.81 mm (1.66+0.08), EL 4.29−4.95 mm (4.75+0.29), EW 2.64−2.93 mm (2.79+0.09), AD/OD 0.63−0.79 (0.66+0.16), GL/eL 0,11−0,16 (0.13+0.02), PW/PL 1.14−1.22(1.18+0.07), EW/HW 1.10−1.36 (1.26+0.09). Intraspecific variation. This is a relatively uniform species, the main variability being associated with the basic pattern of the elytral vittae. Sometimes, the marking on the humeral callus continues externally of the yellow vittae, or it is interrupted and looks like two or three isolated spots. In some specimens the yellow vittae fade to yellowish brown, or the marginal vittae are absent, or they may be partly or entirely confluent at the apex and medial area. The head varies from parrot green to yellowish brown. The labrum and mouthparts vary from chestnut to dark brown. The antennae (except the basal antennomere) are unicolorous, but varying from cinnamon to dark brown. The scutellum varies from yellowish to amber colored. The femora and tibiae are frequently lime green. In some specimens, the prosternal surface varies from parrot green to yellowish brown, and the meso- and metasternum and abdomen may be green to yellowish brown.

50 · Zootaxa 2683 © 2010 Magnolia Press CABRERA & WALSH FIGURES 17−19. Diabrotica collicola Cabrera & Cabrera Walsh (17) Median lobe, lateral view. (18) Median lobe, detail of apex and internal sac, dorsal view. (19) Female genitalia. Scale bars= 0.1 mm. Abbreviations: bf, basal foramen; os, ostium; pu, pump; re, receptacle; sc1, sclerite 1; sc2, sclerite 2; sc3, sclerite 3; sc4, sclerite 4; ssd, sclerotized spermathecal duct; tg, tignum; tl, triangular lobe; va-bu, vagina-bursa copulatrix; vg, vaginal palpi.

Etymology. Collicola, meaning that lives in the hills, refers to the environment where this species has been found. Biological notes. This species was collected on pumpkin and maize, two of the main hosts of Diabroticina throughout their distribution. Field-collected adults survived in a rearing chamber at 25+1 ºC, 14:10 (L:D) h photoperiod, in 1.5-liter cages for up to 4 weeks, during which period mating was observed, and about 80 eggs were obtained. However, no eggs hatched when incubated at 25+1 ºC in Petri dishes lined with moist tissue

NEW SPECIES OF LEAF BEETLE FROM ARGENTINA Zootaxa 2683 © 2010 Magnolia Press · 51 paper, suggesting this species has an egg diapause, a common trait of the Diabrotica in the D.virgifera group (Krysan 1982). Geographic range. The specimens described herein were collected in the vicinity of Balcozna (27º53’ 30’’ S; 65º44’ 04’’ W), Province of Catamarca, at an altitude of 1300 m, and this is the only area where the authors have found this species so far. Type material. Holotype: male, ARGENTINA: Catamarca: Balcozna, 14-II, 2002, on Cucurbita maxima Duchense (), Cabrera Walsh col. Paratypes: 2 males and 3 females, with the same locality, date and collector as the holotype.

Discussion

Smith & Lawrence (1967) proposed the D. virgifera group, including 35 valid names, and placed another seven as incertae sedis. Krysan & Smith (1987) defined the group based on the following characters: habitus more elongate than convex, size small to moderate, sexual dimorphism of antennomeres 2 and 3: in males antennomeres 2 and 3 equal in length; in females antennomere 3 varying from slightly longer to twice as long as 2, antennomeres 2 and 3 together more than 1/2 length of 4, genal space less than 1/4 the maximal diameter of the eye, thorax bifoveate, elytral disc usually with three or more sulci, internal sac of aedeagus with four characteristic sclerites. Diabrotica collicola conforms to the D. virgifera group in that the genal space is less than 1/4 as great as the maximum eye length, and that it has sexual dimorphism in the antennomere length of males, but it differs from this group in the weakly bifoveate thorax and few elytral sulci, which are barely discernible. The species described herein is similar to D. mapiriensis gussi Krysan & Smith and D. porracea Harold, both in the D. virgifera group, and to D. mutabilis Baly and D. fulvofasciata Jacoby that are not in the D. virgifera group. Diabrotica collicola differs from D. mapiriensis gussi in that the latter is smaller, with a piceous head, and with lime-coloured elytra having three diffuse yellow areas. Diabrotica collicola differs from D. porracea in that the latter is convex, bright green, sometimes lighter in tone, and with 4 or 5 distinct sulci on the elytral disc. Diabrotica collicola is similar in coloration and elytral pattern to female D. mutabilis, a species from and . However, D. mutabilis is more convex than D. collicola, and has the head, antennae, scutellum and tibiae piceous, green elytra with a wide yellow longitudinal band, postmedial and apical black markings on the elytra, and males with antennomeres 4−9 modified. Diabrotica fulvofasciata is also similar to D. mutabilis, from which it differs in having only antennomeres 5 to 9 modified in the male, and lacking the elytral markings. However, it is difficult to place a substantial number of Diabrotica species that shares certain traits with the D. virgifera group. For instance, while the females of some of these species possess D. virgifera characteristics, males present atypical antennal dimorphisms and different sclerites in the internal sac. Even Krysan & Smith (1987) acknowledged that several Diabrotica species that resemble species in the D. virgifera group should eventually be included in new groups. They included some of these species in their key of this group, but assigned them to two other species-groups not formally named; the first group comprised of species with a more convex habitus like D. tortuosa Jacoby, and the second group, with species close to D. tumidicornis Erichson, which present enlarged antennomeres 7 to 9. Diabrotica fulvofasciata and D. mutabilis (latter not included in their key), could eventually be placed in the same species group as D. tumidicornis. Consequently, we modify the key of Krysan & Smith (1987) to include the new species described herein, as well as D. mutabilis and proceed with their number to maintain continuity.

Key to Diabrotica collicola and similar species (modified from Krysan & Smith 1987)

39 (38) Elytra green with no yellow or pale brownish yellow markings (examples from Texas and may have interrupted pale yellow vittae); elytron often with piceous or fuscous spot or vitta at, or originating at humeral angle ...... 44

52 · Zootaxa 2683 © 2010 Magnolia Press CABRERA & WALSH - Elytra green with broad yellow or pale brownish yellow vittae or diffuse yellow spots basally and apically40 40 (38) Yellow elytral maculation forming vittae of approximately uniform width, arising from humeri and meso-basal area near humeri, not from suture, sometimes interrupted medially...... 41 - Elytra tinged with yellow in two areas: one basal and extending apically from humeri, the other on the apical convexity; epipleural fold frequently tinged yellow along length from base to apical convexity; head and pectus piceous...... D. mapiriensis gussi Krysan & Smith. 41 (40) Antennomeres 5 to 10 of male much greater in diameter than 1 through 5 and 11, modified...... 42 - All antennomeres of male approximately equal diameter...... 43 42 (41) Head, antennae and thorax testaceous. Each elytron with a broad yellow vitta; discal elytral sulci numbering no more than 3, usually fewer, sometimes absent. Venezuela...... D. fulvofasciata Jacoby. - Head and antennae (except antennomeres 9 to 11) piceous. Each elytron with a broad yellow vitta, commonly narrowed or interrupted medially (sometimes represented as isolated spots) and two postmedial and basal black spots; elytral disc with 2 sulci. Venezuela and Colombia ...... D. mutabilis Baly. 43 (41) Each elytron yellow to green with a broad yellow vitta not including apical margin, commonly narrowed or interrupted medially. Each elytron with 4 or 5 distinct sulci on disc ...... D. porracea Harold. - Each elytron parrot green with two yellowish vittae, one medial and tinged with brown, the other marginal, the two vittae partly or entirely confluent at apex. Each elytron with 2 very weak sulci on disc.... D. collicola n. sp. 44 (39) Antennomeres 7, 8 and 9 of male distinctly thicker than other segments; disc of elytra very weakly sulcate; (not in D. virgifera group) ...... D. tumidicornis Erichson. - All antennomeres of approximately equal diameter; disc of elytra with 4 to 5 distinct sulci...... 45 45 (44) Antennae (except proximal few), clypeus, tibiae, and tarsi infuscated, often shiny piceous...... 46 - Antennae, vertex, clypeus, tibiae, and tarsi testaceous or greenish brown; elytra often lacking humeral spots...... D. barberi Smith and Lawrence (in part). 46 (45) Elytra with humeral spots or vittae small or absent, when present, confined to humeral callus; vertex and scutel- lum usually testaceous...... 47 - Elytra with humeral spots or vittae often extending beyond middle of elytra, sometimes absent; vertex; vertex and scutellum usually piceous; Kansas and Texas to Arizona and Chihuahua...... D. longicornis (Say) (in part). 47 (46) Eastern United States...... D. barberi Smith and Lawrence (in part). - (not in D. virgifera group) ...... D. piceicornis Baly.

Clave para Diabrotica collicola y especies parecidas (modificada de Krysan & Smith 1987)

39 (38) Elitros verdes sin manchas amarillas o castaño amarillentas pálidas (En ejemplares de Texas y Méjico pueden presentar bandas interrumpidas amarillo pálidas); élitros a menudo con bandas o manchas píceas o castaño oscuras en los ángulos humerales, u originándose en los mismos ...... 44 - Elitros verdes con bandas castaña amarillentas pálidas o manchas difusas amarillas basales y apicales...... 40 40 (38) Manchas elitrales amarillas, bandas de ancho aproximadamente uniforme desde los húmeros y área mesobasal cerca del húmero, no desde la sutura, en ocasiones interrumpida medialmente ...... 41 - Elitros manchados con amarillo en dos áreas: una basal extendiéndose apicalmente desde el húmero, la otra en la convexidad apical; epipleura frecuentemente manchada con amarillo longitudinalmente desde la base hasta la convexidad apical; cabeza y tórax píceos...... D. mapiriensis gussi Krysan & Smith. 41 (40) Antenitos 5 a 10 de los machos de mayor diámetro que del 1 al 5 y el 11, modificados...... 42 - Todos los antenitos de los machos aproximadamente igual diámetro ...... 43 42 (41) Cabeza, antenas y tórax testáceos. Cada élitro con una banda ancha amarilla; no más de 3 surcos discales, usual- mente menos, en ocasiones ausentes. Venezuela ...... D. fulvofasciata Jacoby. - Cabeza y antenas (excepto antenitos 9 a 11) píceos. Cada élitro con una banda amarilla ancha, comúnmente más angosta o interrumpida medialmente (en ocasiones como manchas aisladas) y dos manchas negras postmedial y basal; dos surcos discales. Venezuela y Colombia ...... D. mutabilis Baly. 43 (41) Cada élitro de amarillo a un amplio espectro de verde con una banda amarilla ancha que no incluye el margen apical, comúnmente más angosta o interrumpida medialmente. Cada élitro con 4 o 5 surcos distinguibles en el disco...... D. porracea Harold. - Cada élitro verde brillante con dos bandas amarillentas, una medial manchada con castaño, otra marginal, ambas bandas parcial o enteramente confluentes en el ápice. Cada élitro con dos surcos muy débiles en el disco ...... D. collicola n. sp. 44 (39) Antenitos 7, 8 y 9 de los machos claramente más gruesos que los restantes antenitos; disco elitral muy débil- mente surcado; Per (no en el grupo D. virgifera ) ...... D. tumidicornis Erichson. - Todos los antenitos de aproximadamente igual diámetro; disco elitral con 4 o 5 surcos evidentes ...... 45

NEW SPECIES OF LEAF BEETLE FROM ARGENTINA Zootaxa 2683 © 2010 Magnolia Press · 53 45 (44) Antenas (excepto los antenitos proximales), clípeo, tibias y tarsos castaños oscuros a menudo píceos, brillantes ...... 46 - Antenas, vértex, clípeo, tibias, y tarsos testáceos o castaño verdosos; élitros a menudo sin manchas humerales ...... D. barberi Smith & Lawrence (en parte). 46 (45) Elitros con manchas humerales o bandas pequeñas o ausentes, cuando presentes, confinadas al callo humeral; vértex y escutelo usualmente testáceos...... 47 - Elitros con manchas humerales o bandas pequeñas o ausentes, cuando presentes, confinadas al callo humeral; vértex y escutelo usualmente píceos; Kansas, Texas a Arizona y Chihuahua...... D. longicornis (Say) (en parte). 47 (46) Este de Estados Unidos ...... D. barberi Smith & Lawrence (en parte) - Brasil (no en el grupo D. virgifera)...... D. piceicornis Baly.

Conclusions

Diabrotica is the most numerous and complex genus of Diabroticites and the creation of the species groups was to a point arbitrary. Recent phylogenetic studies of a number of Diabrotica species based on different sources of characters claim that the D. virgifera group is monophyletic (Clark et al. 2001a; Szalanski et al. 2000), whereas the D. fucata group is considered monophyletic by some (Clark et al. 2001a,b; Szalanski et al. 2000), and polyphyletic by others (Eben & Espinosa de los Montero 2004; Cabrera & Durante unpublished results). However, these analyses only include a small number of species, and do not include, among others, those species that Krysan & Smith (1987) relate to the D. virgifera group based on similarity. The D. virgifera group as presently defined is an open question, which requires re-evaluation in future studies. As happens with D. collicola, other species of Diabrotica are difficult to assign to the D. virgifera group based on the morphological traits currently used for its classification. This group should be revised in order to place numerous related species that possess some of its traits. In the future, both morphological and molecular studies will have to be combined and advanced in order to adequately define the Diabrotica species groups.

Acknowledgments

We are grateful to A. S. Konstantinov (USDA/ARS Systematic Entomology Laboratory, Smithsonian National Museum of Natural History, Washington, D.C.), D. G. Furth (Department of Systematic Biology, Smithsonian Institution, Washington, D.C.), and V. Savini (Museo del Instituto de Zoología Agrícola Francisco Fernandez Yepez, Universidad Central de Venezuela, Maracay) for loaning us many museum specimens, and to M. Theiller for the habitus illustration. This work was partially supported by the Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET-PIP n 5486).

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