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Iheringia. Série Zoologia

Print version ISSN 0073-4721On-line version ISSN 1678-4766

Iheringia, Sér. Zool. vol.105 no.4 Porto Alegre Oct./Dec. 2015

http://dx.doi.org/10.1590/1678-476620151054439452 

Articles

Comparative morphology of the type-species of Isotes and Synbrotica(Coleoptera, Chrysomelidae, Galerucinae), with a new synonymy of species

Morfologia comparada das espécies-tipo de Isotes e Synbrotica(Coleoptera, Chrysomelidae, Galerucinae), com nova sinonímia de espécies

Juliana M. S. Rodrigues1  2 

José Ricardo M. Mermudes1 

1.Laboratório de Entomologia, Departamento de Zoologia, Universidade Federal do Rio de Janeiro, Caixa Postal 68044, 21941-971, Rio de Janeiro, RJ, Brazil. (jrmermudes@gmail.com)

2.Laboratório Nacional e Internacional de Referência em Taxonomia de Triatomíneos, Instituto Oswaldo Cruz, Av. Brasil, 4365, Manguinhos, 21045-900, Rio de Janeiro, RJ, Brazil. (julianamourao@yahoo.com.br)


ABSTRACT

In order to solve the affinities of the species of Isotes Weise, 1922, a detailed morphological comparative study was carried out based on type-species of Isotes and its junior synonym,Synbrotica Bechyné, 1956. Isotes tetraspilota (Baly, 1865) and Isotes borrei (Baly, 1889) had their morphology of mouthparts, endosternites, wings and both male and female genitalia compared by the first time. A new synonymy is established between Isotes borrei (Baly, 1889) and Isotes crucigera (Weise, 1916) syn. nov. based on external and genitalia morphology. New structures for Section Diabroticites Chapuis, 1875 are presented and discussed.

KEYWORDS Taxonomy; Neotropical; Luperini; Diabroticites

RESUMO

Para resolver as afinidades das espécies de Isotes Weise,1922, um estudo morfológico comparado foi realizado com base nas espécies-tipo deIsotes e de seu sinônimo-júnior,Synbrotica Bechyné, 1956. Isotes tetraspilota (Baly, 1865) e Isotes borrei (Baly, 1889) tiveram a morfologia de suas peças bucais, endosternitos, asas e genitálias masculina e feminina comparadas pela primeira vez. Uma nova sinonímia é estabelecida entre Isotes borrei (Baly, 1889) eIsotes crucigera (Weise, 1916) syn. nov., com base na morfologia externa e genitália. Estruturas inéditas para a Seção Diabroticites Chapuis, 1875 são apresentadas e discutidas.

PALAVRAS-CHAVE Taxonomia; Neotropical; Luperini; Diabroticites

The genus Isotes was proposed by Weise (1922) to include a single species, Isotes quadrimaculataWeise, 1922 - type-species by monotypy. The distribution of the genus is Neotropical, occurring in Central and South America. Aslam (1972) studied the holotype of Isotes quadrimaculata Weise, 1922, established synonymy with Synbrotica tetraspilota (Baly, 1865), and consideredSynbrotica Bechyné as junior synonym of Isotes. The synonymy was proposed in a single note, without remarks of any features that led to genera synonymy.

This fact increased a problem because a great number of species were assigned toSynbrotica since the genus was proposed by Bechyné (1956). Additionally, subsequent authors never provided diagnostic features for the genus. Bechyné (1956)only designated Diabrotica borrei Baly, 1889, from Brazil, as type-species of Synbrotica.

Bechyné & Bechyné (1969) provided only six diagnostic features for Synbrotica, but also they affirmed that the genus included many not congeneric species, forming an artificial group, and that they should be transferred. Despite this, even after the synonymy with the genusIsotes, none other study has been conducted to investigate this problem.

Isotes currently includes 179 species (Bechyné & Bechyné, 1970; Wilcox, 1972, 1975; Moura, 2003) and the systematic was resumed to the addition of species, redescriptions, new combinations, citations in catalogue and key to genera. None other contribution provided detailed anatomical study, except Moura (2009) that illustrated the male terminalia of Isotes eruptiva (Bechyné, 1955).

Since that the previous authors synonymized these genera and did not compare both type-species, the aim of this work was to perform a detailed study of the morphology of the type-species Isotes tetraspilota (Baly) and the type-species ofSynbrotica, I. borrei (Baly), including external morphology, mouthparts, wing venation, endosternites, and female and male genitalia. In parallel, a comparative study of Isotes borrei (Baly) and Isotes crucigera (Weise) is performed by first time to determinate if they should be synonymized.

Taxonomic history. Diabrotica tetraspilota was described by Baly (1865) based on a single male from Mexico. Later, Smith & Lawrence (1967) noticed another male from Guatemala with a Baly's label, but could not be considered a syntype because Guatemala was not originally included in the description. This species was assigned to the genus Synbrotica by Smith & Lawrence (1967). After that, it was considered as senior synonym of Isotes quadrimaculataWeise by Aslam (1972).

Diabrotica borrei was described by Baly (1889) to a female from "Tejuca" (= Tijuca, a neighborhood in city of Rio de Janeiro, Brazil), with possible paralectotypes from Petropolis, Brazil (Smith & Lawrence, 1967). Bechyné (1956) designed posteriorly as type-species ofSynbrotica.

Another species, Diabrotica cruciata, was proposed by Baly (1889) in the same paper and page of D. borrei, based on a male from Santa Catarina (Brazil). Both species had short descriptions with a minor variation between them.

Gahan (1891a) finished the uncompleted Baly's work (1890), transcribing the descriptions for D. borrei and noticed that "D. cruciata, Baly (Ent. Mo. Mag., xxv., p. 253) is merely a variety of this species". Finally, Gahan (1891b) and Weise (1916) provided the replacement name crucigera, pointing out D. cruciataBaly as junior homonym of D. cruciataJacoby, 1887.

Weise (1924) considered D. crucigera as an aberration of D. borrei in the catalogue, and Bechyné (1958) placed D. crucigera in the genus Synbrotica. Despite of the overall similarity they were treated as distinct species.

MATERIAL AND METHODS

The material examined was provided by the following institutions (names of curators in parenthesis): DZRJ, Coleção José Alfredo Pinheiro Dutra, Universidade Federal do Rio de Janeiro, Rio de Janeiro, Brazil (J. R. Mermudes); CEIOC, Coleção Entomológica do Instituto Oswaldo Cruz, Rio de Janeiro, Brazil (M. E. Felix); EMEC, Essig Museum of Entomology, University of California, Berkeley, USA (C. B. Barr); MNRJ, Museu Nacional, Universidade Federal do Rio de Janeiro, Rio de Janeiro, Brazil (M. L. Monné); MZSP, Museu de Zoologia, Universidade de São Paulo, São Paulo, Brasil (S. A. Casari); USNM, National Museum of Natural History, Smithsonian Institution, Washington D.C., USA (D. G. Furth); ZMHB, Museum für Naturkunde der Humboldt-Universität, Berlin, Germany (J. Frisch).

The terminology follow Moura (1998b, 2009), Konstantinov (1998), Lingafelter & Konstantinov (1999) and Cabrera & Durante (2003), except for numeration of mandibles' teeth that followSavini & Joly (2006).

RESULTS

Morphological comparison between Isotes borrei,I. crucigera and I. tetraspilota

Integument, general coloration (Figs 1-5). In Galerucinae the pattern of the integument color is highly variable, often being the only feature used to separate species when lacking studies with genitalia. By other hand, the species also exhibit interspecific variation, while other species morphologically distinct show the same color pattern. Both I. borrei (Figs 1-2), I. crucigera (Fig. 3) and I. tetraspilota (Figs 4-5) have elytra with integument yellow-brownish, with two large black or blue-black spots.

Figs 1-5 Habitus: 1, 2, I. borrei (Baly); 3,I. crucigera (Weise) syn. nov.; 4, 5, I. tetraspilota (Baly). Scale bars = 1,0 mm. 

Head (Figs 6-11). The width across the eye is narrower than anterior margin of pronotum. Vertex convex (Figs 8, 11), with variable punctuation, and a weak depression near frontal tubercles. Antennal calli transverse and subtriangular, slightly larger than antennal cavities (Figs 6, 9). Eyes (Figs 6-11) small (by relation between the maximum eye diameter in lateral view, against the interocular distance), and well-marked orbital lines near the outer margin. Antennal cavities rounded, inserted just below the midline of the eye. Frons with frontal ridge and fronto-clypeal suture, surface variably punctate and rugose. Isotes borrei and I. crucigera exhibit elongated genal area (length subequal to half or feebly more than the maximum diameter of the eye), and front as wide as long (Figs 9-11). Isotes tetraspilotahas genal area and front evidently wider than long (Figs 6-8).

Mouthparts. Clypeus (Figs 6, 9) narrow and subrectangular, with a row of setae (variable in number) near the basal margin.Isotes tetraspilota has eight setae (Fig. 6) while I. borrei and I. crucigera have six setae (Fig. 9). Both species have labrum (Figs 6, 9, 12-13) rounded at the sides, feebly notched at apex, and with a row of six submedian long setae in coarse punctate.

Mandibles (Figs 14, 15) symmetrical, subtriangular, outer margin distinctly curved, inner cutting edge with five apical teeth, with short and dense setae on the outer surface. First tooth small, attenuated at the apex; second tooth twice the length of the first; third narrowed and twice the length of the second and fourth teeth; fourth variable at apex, slightly longer than the fifth, this short and blunt; prostheca wide, with abundant setae, extending almost from the basal margin of the mandible to the first apical tooth; mola longer than wide and sculptured with rows of transverse elevations. Both species have similar shape and length of mandibles, except: 1) inner cutting edge concave in I. tetraspilota and distinctly straight in I. borrei; 2) mandibular teeth, in external view: tooth three with inner margin shaped and more projected in I. borrei, and inner margin with slight sinuosity and less projected in I. tetraspilota. While that in ventral view, it is robust and little curved in the last species and narrower and bent to the side in the first one; 3) tooth two and four, ventral view, blunt in I. tetraspilota and pointed in I. borrei; and 4) mola wider in I. borrei than in I. tetraspilota.

Savini & Joly (2006) observed that the presence or absence of mola is not an informative character for the classification when they studied the mandibles of 22 genera of Galerucinae. Thirteen of the 22 genera studied does not have mandibular mola, but when the mola exist, it is variable in the shape, can be formed by parallel blades, either toothed or fused with the body of mandible, and separated by a well-defined suture, an indication that the structure and shape of the mola could be contribute together with other characters to definition of genera or groups of genera in the subfamily. The shape of the mandible, prostheca and number of teeth of the incisive area are other valuable characters that may contribute to the classification of the group.

Maxilla (Figs 16, 17). In all species examined, the maxilla present cardo narrowed at proximal portion, wider apically, and with long setae; basistipe with long setae at the latero-external edge, and dististipe well developed with short setae at the inner margin, and short setae near the lacinia, with apical pubescence fringed and strongly dense; galea subcylindrical and slightly longer than lacinia, with long and dense setae at the apex. Maxillary palps well developed, with four palpomere, exceeding the galea by apical palpomere; palpomere one short, subrectangular, subequal as half the second; palpomere two subconical; palpomere three subcylindrical, twice longer than the apical; apically acute. Distal part of the galea twice the length of the basal, even more curve inI. borrei that in I. tetraspilota. Palpomeres clearly shorter and robust in I. tetraspilota and more elongated and narrower at the base in I. borrei.

Labium (Figs 18, 19). Labium poorly developed, with ligula membranous and not lobated. Labial palps with three palpomeres, basal short and strongly transverse; palpomere two slightly longer than the apical, subcylindrical, and slightly longer than wide; apical narrow to apex. Prementum subrectangular in both species. Ligula reaching the base of the apical palpomere in both species, but it is distinguished in the shape, apex narrowed in I. tetraspilota, and broadly rounded in I. borrei. Mentum (= postmentum Cabrera & Durante, 2003) transverse and subparallel at sides in I. tetraspilota and subtrapezoidal in I. borrei.

Figs 6-19 Head: Isotes tetraspilota (Baly): 6, frontal; 7, dorsal; 8, lateral; I. borrei (Baly): 9, frontal; 10, dorsal; 11, lateral. Labrum: 12, I. tetraspilota; 13,I. borrei. Mandible: 14, I. tetraspilota; 15, I. borrei. Maxillae: 16,I. tetraspilota; 17, I. borrei. Labium: 18, I. tetraspilota; 19, I. borrei (bs, basistipes; cd, cardo; ds, dististipes; ga, galea; ge, gena; lc, lacinia; lg, ligula; lp, labial palp; m, mentum; mo, mola; mp, maxilar palp; prm, prementum; pros, prostheca; I-V, position of the mandible teeth). Scale bars = 0,5 mm. 

Antennae. Antennae with 11 antennomeres, exceeding the middle of elytra. Scape cylindrical, slightly thickened to apex, one third longer than the pedicel and subequal to the antennomere III; antennomere IV longer than the III; V-VI subequal in length, both slightly shorter than the IV; antennomeres VII-X lightly shorter; XI slightly longer than the X, acute at apex.

Thorax. Prothorax (Figs 20-22) wider than long and narrower than the elytral base. Pronotum (Figs 20-21) slightly convex, with variable depressions in the basal half. Isotes borrei exhibits well marked lateral margins, subparallel and slightly sinuous in the basal third (Fig. 21); anterior and posterior angles rounded; each angle with a long setae inserted in a coarse and deep punctuation; and other short setae between angles, unstable in number. In I. tetraspilotathe lateral margins of pronotum is convex, slightly rounded, and wider in the middle (Fig. 20).

The species studied have the pronotum wider than long, but the proportion varies from 1.6 to 1.7 times wider in I. tetraspilota (Fig. 20), while in I. borrei and I. crucigera from 1.8 to 2.0 times (Fig. 21). The type-species Isotes tetraspilota has no depressions on pronotum (Fig. 20). On the other hand, I. borrei has a transverse depression in the posterior half (closer to the posterior margin), reaching the sides, where it is evidently deeper (Fig. 21). Prosternum (Fig. 22) transverse, with a short and laminar prosternal process between the procoxae. Procoxal cavities contiguous and opened behind. Proendosternite (Figs 23, 24) with divergent projections towards posteriorly, slightly curved in I. tetraspilota and linear in I. borrei.

Figs 20-26 Pronotum, dorsal: 20, I. tetraspilota (Baly); 21,I. borrei (Baly). 22, Prosternum and procoxae,I. tetraspilota. Proendosternites, posterior view: 23, I. tetraspilota; 24, I. borrei. Pterothorax, I. tetraspilota: 25, ventral; 26, lateral (gl, gland; msp, mesepimeron; mss, mesepisternum; mts, metepisternum; mtp, metepimeron; pr, proendosternite). Scale bars: 20-24 = 0,5 mm; 25, 26 = 1,0 mm. 

Mesothorax (Figs 25, 27-28). Mesonotum with mesoscutellum elevated, subtriangular, smooth and shiny: in I. tetraspilota it is widely rounded, while I. borrei is more strongly convergent at sides. Mesosternal process subtriangular and narrowed. Mesocoxal cavities rounded, close to each other and closed at sides.

Metathorax (Figs 25, 26, 29-34). Metanotum (Figs 29, 30) transverse, wider than long in I. borrei and as wide as long in I. tetraspilota. The apodeme "d" of metanotum crossing with "c" near the middle of it, apodeme "b" crossing "c" in the proximal region. Postnotum (Figs 31, 32) with sides projected distally in I. borrei and not projected in I. tetraspilota; in posterior view, I. borrei shows subtriangular sides, broader at the base, while in I. tetraspilotathey are narrow and slightly convergent. Metendosternite (Figs 33, 34) with lateral arms fused to lamina, angulated at apex in I. tetraspilota and straight in I. borrei. Both species exhibit elongated peduncle; meso- and metafurcal tendons moderately developed, and inserted in the middle of the lateral arms.

Elytra (Figs 1-5). Elongated, with subparallel sides, emarginated, with apical margin subtruncated. Humeri slightly prominent. Punctuation with variable thickness. Setae present or absent. Epipleurae visible laterally or ventro-laterally, with variable width: in I. tetraspilota it is large only in the basal third, narrow in the apical two thirds, interrupted near the apex of elytra; in I. borrei andI. crucigera it is large only in the basal half, narrowed towards the apex. Integument with setigerous punctures, fine punctuation, or smooth and glabrous are species-specific characteristics.

Figs 27-34 Mesoscutellum: 27, Isotes tetraspilota (Baly); 28,I. borrei (Baly). Metanotum: 29, I. tetraspilota; 30, I. borrei. Post-notum, posterior view: 31, I. tetraspilota; 32, I. borrei. Metendosternite: 33, I. tetraspilota; 34, I. borrei (a-d, metanotum apodemes; ar, arm; es, escutelum; la, lamina; mg, median groove; pd, peduncle; pn, post-notum; te, tendon). Scale bars = 0,5 mm. 

Hindwings (Figs 35, 36). Length of the apical region, from the apex of the Radial cell to apex of the wing, slightly shorter than the basal region, from the base of the wing to the apex of the Radial cell. Costa (C) short, less than half of Subcosta (Sc) in I. tetraspilota, and elongated in I. borrei, at least two thirds of Sc length. Radial (RA) present. Both species with Radial cell (R) wide and closed. Radial cross vein 3 (r3) (= r4 by Cabrera & Cabrera Walsh, 2004a) present and developed. Isotes tetraspilota andI. borrei with Posterior Media 3 (MP3) vestigial in the posterior region. Isotes borrei differs from I. tetraspilota by Radial Posterior (RP) present and Anal Anterior (AA) conspicuously sharpened. The veins Media Posterior 1+2 (MP1+2), radial posterior-media posterior 2 (rp-mp2), Radial Posterior 3+4 (RP3 +4), Cubitus Anterior (CuA), Cubitus Anterior 3+4 (CuA 3+4), Media Posterior 3 (MP3) and Jugal (Ju) present.

Figs 35-36 Hindwings: 35, Isotes tetraspilota (Baly); 36,I. borrei (Baly) (AA, Anal Anterior vein; arc, arculus; C, Costa; CuA, Cubitus Anterior vein; CuA3+4, Cubitus Anterior vein 3+4; E, Empusal; Ju, Jugal; MP1+2, Media Posterior 1+2; MP3, Media Posterior vein 3; R, Radial cell; r3, Radial transversal 3; RA, Radial Anterior vein; RP, Radial Posterior vein; rp-mp2, radial posterior vein - media posterior 2; RP3+4, Radial Posterior vein 3+4; Sc, Subcostal vein). Scale bars = 1 mm. 

Important to note that the wing of I. borrei was severely damaged in the region of the anal and jugal lobe, not allowing the observation of variations in these regions. Apparently the pattern of Isotes in these two species is also similar to four subfamilies illustrated by Chaboo (2007, fig. 70, AC, E, HK) as the R, r3, RP and MP1+2: Lamprosomatinae, Criocerinae, Galerucinae and Cassidinae. Regarding the veins AA, CuA, CuA3 +4 and MP3 it is also very similar to the wing of galerucine Ophraella sp. (Chaboo, 2007, Fig. 79 C). Both wings keep the vein pattern of Diabroticites, as noted by Cabrera & Cabrera Walsh (2004a,b) inDiabrotica calchaqui Cabrera & Cabrera Walsh, 2004 andPlatybrotica misionensis Cabrera & Cabrera Walsh, 2004.

Empusal (E) and arculus (arc) present. Hamilton (1972) designated the first anal vein as Empusal, being followed by researchers of Cerambycidae (Marinoni & Almeida, 1983; Napp, 1994). The arculus, according to Forbes (1922), is a fragment of the Media, in which the base is fused to the Radial and to the apex of Cubitus (= RP, MP1+2 and RA1+2, respectively, according to Kukalová-Peck & Lawrence, 1993). Napp (1994) noted that the arculus is poorly defined among the cucujoids studied, while in chrysomelids and some cerambycids it is clearly distinct. Crowson (1955), Jolivet (1957) and Suzuki (1969) did not mention the arculus to chrysomelids wings. Kukalová-Peck & Lawrence (1993), in turn, conducted a detailed study of the membranous wings of Coleoptera, being followed by the more recent works of Coleoptera (Kukalová-Peck & Lawrence, 2004; Lawrence et al., 2011) and Chrysomelidae (Lingafelter & Konstantinov, 1999; Cabrera & Cabrera Walsh, 2004a,b; 2010 - Galerucinae; Chaboo, 2007 - Cassidinae; Chamorro-Lacayo & Konstantinov, 2011 - Lamprosomatinae). The scheme adopted by these authors agrees with the proposal that the Cubitus vein of Forbes (1922) is designed as Media. Kukalová-Peck & Lawrence (2004), however, gave a different definition for the arculus: alternative term for arm mp-cua present in all Endoneoptera + Hemineoptera and some Blattoneoptera. Herein we follow the term and interpretation adopted by Forbes (1922).

Legs. All legs are similar, slightly increasing in length. Coxae of pro- and mesolegs rounded, and mesocoxae with a glandular opening near the anterolateral area (Fig. 25). Metacoxae transverse. Trochanters subtriangular and reduced. Femora fusiform, tibiae slender, with apical spur - except protibiae of males.

Tarsi with tarsomeres I and II subrectangular (males with adhesive disc on the ventral surface of pro- and or mesotarsomeres); tarsomere III bilobed, with dense pubescence; tarsomere IV reduced on the basis of V, this elongated, with bifid claw at its end.

Abdomen (Figs 37-39). Abdomen with urosternites I-V subequal in length, wider than long, gradually decreasing in width until the urosternite V; this subtrapezoidal, with variable apex between male and female species of Isotes. Apex of the urosternite V of males weakly notched (Fig. 37) in I. tetraspilota and truncated (Fig. 39) in I. borrei and I. crucigera. In females, in turn, both species exhibit the apex of the urosternite V rounded (Fig. 38).

Figs 37-39 I. tetraspilota (Baly), Abdomen, ventral view: 37, male; 38, female. 39, Urosternite V, male, I. borrei(Baly). Scale bars: 37, 38 = 1,0 mm; 39 = 0,5 mm. 

Male genitalia (Figs 40-47). Aedeagus (median lobe and tegmen; Figs 40-41). Median lobe, ventral view, placed laterally in the abdomen, with curvature toward the left side, and subequal in length to the last four urosternites in I. borrei and I. crucigera. In I. tetraspilota it reaches the first urosternite. Both species have variable curvature in the median lobe, as follows: 1) basal third strongly curved, 90° from the rest of the body of the median lobe at the height of the basal constriction, well-marked in I. borrei and I. crucigera (Fig. 40); and 2) basal third slightly curved, less than 45° relative to the apical two thirds of the median lobe, with well-marked constriction; apical third slightly curved in I. tetraspilota (Fig. 41).Isotes borrei and I. crucigera have a basal crest, slightly sclerotized with length equal to half of the basal third of the median lobe; I. tetraspilota also have this crest, but much shorter and narrowed.

According to Moura (2009), the basal orifice (= median foramen Matsumura & Suzuki, 2008) in Luperini is protected by a hood-shaped process, with rounded border. The basis of this process, in ventral view, is similar in both species, being truncated and notched at apex. In lateral view, all have rounded base, being sinuous in I. borrei and I. crucigera (Fig. 40) and straight in I. tetraspilota (Fig. 41). None of them have basal hooks, corroborating the results found for the tribe Luperini byMoura (2009).

The apex of the median lobe is variable in Luperini (Moura, 2009). However the conformation of the apical flap together with the ventral lobe (Figs 42, 43) indicated importance for the determination of species. In this study we found the following relation of forms to the apex of the median lobe: 1) apical flap rounded, with crenulated margin, and ventral lobe narrowed at the apex, which is indented in I. borrei and I. crucigera (Fig. 42), and 2) apical flap rounded, with smooth margin, and ventral lobe with parallel sides that narrow at the apical margin in a rounded projection in I. tetraspilota (Fig. 43).

Figs 40-47 Aedeagus lateral and sclerites of the internal sac, dorsal: 40,I. borrei (Baly); 41, I. tetraspilota (Baly). Apex of median lobe, dorsal and ventral view, respectively: 42, I. borrei (Baly); 43,I. tetraspilota (Baly). Tegmen, dorsal view: 44,I. borrei (Baly); 45, I. tetraspilota (Baly). Spiculum gastrale: 46, Isotes borrei (Baly), Y-shaped, not fusioned; 47,I. tetraspilota (Baly), V-shaped, fusioned (af, apical flap; cr, basal crest of the median lobe; is, internal sac; llb, lateral lobe; o, basal orifice; os, ostium; sc, sclerite of the internal sac; vl, ventral lobe). Scale bars = 0,5 mm; except 40-43 = 1,0 mm and 0,5 mm, respectively. 

The lateral margins of the apical third of aedeagus, in ventral view, may or may not present a pair of lateral lobes (stuck to the sides of the ostium, according toCabrera & Cabrera Walsh, 2004a,b) that vary in shape. Similarly to the apex in dorsal view, this feature is specific. The studied species show the following shapes: 1) subtriangular in I. tetraspilota (Fig. 43), and 2) sinuous in I. borrei and I. crucigera (Fig. 42). Isotes tetraspilotaexhibit a highly sclerotized structure at apex surrounded by a moderately thick and dense punctuation (Fig. 43), not observed in the other species.

The internal sac is a membranous structure supported by the median lobe; during mating it is everted through the median orifice (= ostium Moura, 2009) under pressure from the hemolymph, forming the endophallus (Matsumura & Suzuki, 2008).

The internal sac in these two species varies in the number and shape of the sclerites (Figs 40, 41): one in I. borrei and I. crucigera, in the middle region of the median lobe, dorsally with the anterior half subretangular, rounded edges, and posterior half bifid - with two spiny projections directed towards the apex, the right greater than left; and two in I. tetraspilota, median, highly curved, pointed at the apex.

Tegmen (Figs 44, 45). The length of tegmen, in both species, is subequal to the basal third of the median lobe, and somewhat sclerotized. Two patterns were observed regarding form: 1) V, rounded at the base inI. borrei and I. crucigera (Fig. 44) and 2) U, subtruncated at the base inI. tetraspilota (Fig. 45); both with basal membranous projections. The lateral arms of tegmen meet the median lobe just below the dorsal constriction. Nadein (2006) appointed without performing phylogenetic analysis, that the simple Y-shaped tegmen, with a long basal part and two short branches forming a fork in the apical basal orifice of the penis, is plesiomorphic for representatives of the Tribe Alticini. According to Moura (2009), the tegmen of Metacyclini and Galerucini maintains this pattern, but in the species of Luperini - Paranapicacaba teintuieri (Allard, 1894) and Isotes eruptiva (Bechyné, 1955) - it consists of a subexagonal basal plate with a fork which originate a subparallel pair of arms directed laterally near the apex. The studied species show that Luperini displays various formats of tegmen, different from observed by Moura (2009).

Spiculum gastrale (Figs 46, 47) with length subequal to half of the median lobe; in all species it is somewhat sclerotized. The studied species have two basic conformations: 1) Y-shaped, base united, not fused in I. borrei and I. crucigera (Fig. 46); and 2) V shaped, fused at base in I. tetraspilota (Fig. 47). According to Powell (1941) and Moura (2009), the Luperini species exhibit V-shaped spiculum gastrale, being the vertex (= base) not fused in Paranapiacaba teinturieri and, inLuperodes meraca (Say, 1825), Y-shaped.

Female genitalia (Figs 48-58). Both species present tergite VIII trapezoid shape, with sides slightly rounded, anterior angles rounded and pointed posteriorly. The sides fold toward the structures of the female genitalia, visible in ventral view.

Tergite IX (Figs 48, 49). Structure situated between the tergite VIII and sternite VIII, above the opening of the gut. It is characterized by a small plate, usually little sclerotized (Lingafelter & Konstantinov, 1999). The tergite IX is present in all species, with little observed variation in conformation and the length/width ratio: 1) longer than wide, inner margin sub truncated in I. borrei and I. crucigera (Fig. 48), and 2) as wide as long, robust with narrow sclerotized region in I. tetraspilota (Fig. 49). All tergites IX exhibit basal setae.

Despite being a structure present in most taxa of Chrysomelidae, including most Alticinae, Galerucinae and Chrysomelinae species (Lingafelter & Konstantinov, 1999), it is neglected in descriptions of female genitalia.

Sternite VIII (Figs 50, 51) slightly sclerotized, with apical and lateral setae near the margins. It shows slight variation in specimens of the same species, but the general shape remains stable. Two forms were observed: 1) slightly longer than wide, with parallel sides and rounded apex, truncated at the proximal margin in I. borrei andI. crucigera, and 2) subtrapezoidal, with proximal margin truncated and wide, narrowing to the distal margin and subrounded in I. tetraspilota (Fig. 51).

Figures 48-58 Tergite IX: 48, Isotes borrei (Baly); 49, I. tetraspilota (Baly). Sternite VIII: 50, Isotes borrei (Baly); 51, I. tetraspilota (Baly). Vaginal Palpi: 52, Isotes borrei (Baly); 53, I. tetraspilota (Baly). Spermatheca: 54, Isotes borrei (Baly); 55, I. tetraspilota (Baly).Bursa copulatrix: 56, Isotes borrei (Baly); 57, I. crucigera (Weise, 1916) syn. nov.; 58, I. tetraspilota, female genitalia - arrow: bursa copulatrix with detail of the very dense median region (cor, cornu; dct, spermathecal duct; nd, nodulus; mp, median part). Scale bars: 48, 49, 52-55 = 0,1 mm; 50, 51, 56-57 = 0,5 mm; 58 = 1 mm. 

Spiculum ventrale (Figs 50-51). The spiculum ventrale (=tignumKonstantinov, 1998) is a rod-shaped sclerite of the endoskeleton, begin the attachment of muscles (Torre-Bueno, 1989). It is present in the two species, in connection with the sternite VIII, varying in the degree of sclerotization, size - compared with sternite VIII - and within reach of the same in the sternite. Regarding the sternite VIII connection can reach the area: distal inI. borrei and I. crucigera, or proximal inI. tetraspilota.

Vaginal palpi (Figs 52, 53) are formed by invagination of the dorsal side of the vagina (Konstantinov, 2002). There are two opinions about the function of these structures: the first considers the palpi as adaptations for posture, with sensorial function in selecting the best substrate to lay eggs, and the second believes that they act in mating (Konstantinov, 1998; Moura, 2008). In all species studied, the palpi are subcylindrical (Cabrera & Cabrera Walsh, 2004b), approximately two times shorter than the spiculum ventrale, strongly punctuated in the apical half and with setae apically. The palpi are united at the basis on all species and vary according to it. At the apex they can be united or separated, a characteristic that varies among specimens of the same species. Moreover, they can be entirely sclerotized or not. Separated palpi apex were observed in I. tetraspilota (Fig. 53). There was variation in I. borrei and I. crucigera, from being completely united to moderately separated (Fig. 52). The shape of the sclerotization of the apex varies inter- and intraspecifically. Additionally, the sides of the palpi are sclerotized in all species, except I. borrei and I. crucigera.

Bursa copulatrix (Figs 56-58). In female insects, it works as a purse, being the genital chamber or part of it. In Coleoptera, it is the blind proximal end of the vagina, with which it is connected widely or closely (Torre-Bueno, 1989). In the studied species, the bursa copulatrix varied in relation to form and the presence or absence of sclerotization. The species present different types and forms of sclerotization: 1) I. borrei (Fig. 56) with median region slightly more sclerotized, quadrangular aspect, and with small bands more sclerotized anteriorly; 2) I. crucigera syn. nov. (Fig. 57) with small oval sclerotized area in the median region; 3) I. tetraspilota (Fig. 58) generally differentiated: median membrane very dense, sharply rounded, without sclerotization.

Spermatheca (Figs 54, 55). Considered a sperm receptacle during mating (Torre-Bueno, 1989). Several terminology are used for the same structure in other families of Coleoptera and even within subfamilies of Chrysomelidae: 1) nodulus, middle part and cornu (Middelhouve & Wagner, 2001; Wagner, 2002; Stapel et al., 2008; Hazmi & Wagner, 2010; Curculionidae the median part is called the ramus); 2) receptacle (= proximal half) and pump (= distal half) (Moura, 1998a,b,c; Cabrera & Cabrera Walsh, 2004a,b, 2010; Cabrera et al., 2005, 2008; Reid & Nally, 2008; Chamorro-Lacayo & Konstantinov, 2009 - Cryptocephalinae); 3) vasculum, vellum and ampulla (Rodriguez, 1994; Borowiec & Skuza, 2004; Borowiec & Pomorska, 2009); and 4) no specific terminology (Moura, 2007 - Galerucinae; Chamorro-Lacayo et al., 2006 - Cryptocephalinae).

In this paper, the first terminology was adopted for the comparison of the morphology of the spermatheca of Isotes (Figs 54, 55). Both species have elongated spermathecae with curved cornus and rounded apex. The shape and width from nodulus varies from elongated, with the same width in the median part in I. borrei and I. crucigera (Fig. 54) to strongly enlarged, cylindrically shaped in I. tetraspilota (Fig. 55). The morphology of the spermatheca has been used to separate genera (Cryptocephalinae in Chamorro-Lacayo et al., 2006), species groups (Stolas, Cassidinae in Borowiec & Pomorska, 2009) and species (Alticini, by Leonardi, 1970), however it was clearly neglected in Galerucinae studies that lacking information terminalia. This study showed the importance of using such characteristics and the real possibly can be tested in further phylogenies.

Spermathecal duct and gland. Both species have the duct length slightly greater than the spermatheca, narrower and slightly sclerotized at the base, and extending until the nodulus. The opening of the spermathecal gland is lateral and proximal to it, with a moderately dilated region after the same. I. tetraspilota(Fig. 55) shows a strongly sclerotized duct, similar to the spermatheca, and unlike other species where it is less sclerotized. The spermathecal gland is poorly sclerotized, narrow and elongated.

Material examined. Isotes borrei (Baly, 1889): BRAZIL, ♂, without date, F. C. Bowditch leg. [1st Jacoby Coll.] (USNM); Rio de Janeiro: Petrópolis, 1, 03.II.1952, H. Barth leg. (CEIOC); Teresópolis, ♀, XII.1925, Dirings leg. (MZSP); Mangaratiba, 2♂, VIII.1938 - II.1939, R. C. Shannon leg. (USNM); Espírito Santo, ♀, without additional data [ex coll. Fruhstorfer] (ZMHB); Córrego Itá, ♂, XI.1956, W. Zikán leg. (MNRJ); Guandú, 3♀, X.1920, F. Hoffmann leg. (MNRJ); São Paulo: Agantuba, ♂, I.1922, Azevedo Marques leg. (MNRJ); Santos, Ilha Santo Amaro, ♀, 02.IV.1912, G.E. Bryant leg. (USNM); Minas Gerais: 1, VII.1925, Loreto Moreira leg. (MNRJ); Rio Grande do Sul: São Leopoldo, ♂, ♀, F. Schneider leg. (ZMHB). Isotes crucigera (Weise, 1916): BRAZIL, Minas Gerais: Conceição da Paraíba, Fazenda S. José, ♀, 01.XI.1960, J. C. M. Carvalho leg. (MNRJ); Rio de Janeiro: Parque Nacional da Bocaina, Estrada Paraty-Cunha, 3♀, 24.I.2010, Mattos & Mermudes leg. (DZRJ); Petrópolis, Morro Castelo, 1, 08.VI.1991, Serpa-Filho, A. leg. (CEIOC); São Paulo, ♀, III-1958, Dirings leg. (MZSP); Bananal, Bocaina, ♂, I-1937, D. Mendes leg. (MNRJ); Barueri, ♂, 15.III.1955, K. Lenko leg. [Coleção Campos Seabra] (MNRJ); Paraná: Marumbi, ♀, II.1944, Dirings leg. (MZSP); Santa Catarina: Corupá, ♂, I.1953, A. Maller leg. [Coleção Campos Seabra] (MNRJ); Joinville, ♂, XI.1956, Dirings leg. (MZSP). Isotes tetraspilota (Baly, 1865): MEXICO, 2♀, 2♂, without date, J. Flohr leg. (ZMHB); 3♂, without date, Hoege S. & Kotze G. leg.; ♀, 1906, Stobre leg. (ZMHB); Chiapas: Tumbala, 4♀, 2♂ (ZMHB); Santo Domingo, S. E. Simojovel, 2♂, 08-15.VII.1949, Ray F. Smith leg.; 2♀, 4♂, 8-15.VII.1949, J. A. Chemsak leg. (EMEC); Córdoba: Fortin de las Flores, Sierra Zongolica [Sa Zongola], ♀, ♂, without additional data (ZMHB); Veracruz: Misantla, 2♂, Hoege leg. (ZMHB); Sonora: Cordova, 2♀, Hoege leg. (ZMHB); Jalapa, 3♀, Deppe leg. (ZMHB); San Andrés Tuxtla, ♂, XI.1944, (EMEC); W. Fortin de las Flores, 2♀, 06.VIII.1962 {Cucurbita moschata and C. martinezi} (EMEC); 2♀, 28.VIII.1962, Ray F. Smith leg. (EMEC); GUATEMALA, Guatemala: San Cristobal, 2♀ (ZMHB).

DISCUSSION

The comparison of I. tetraspilota (Baly, 1865) and I. borrei (Baly, 1889), type-species of the genera Isotesand Synbrotica, showed that the species have a very distinct morphology, including the frons (I. borrei as long as wide and gena elongated; I. tetraspilota wider than long and short gena), clypeus (I. borrei with six setae; and, I. tetraspilota with eight setae), mandibles (inner margin concave inI. tetraspilota and substraight in I. borrei; mola wider in I. borrei that in I. tetraspilota), maxillae (distal part of the galea more curved in I. borrei than inI. tetraspilota; with the articles of palp shorter and robust in I. tetraspilota), labium (ligula narrowed at the apex inI. tetraspilota and broadly rounded in I. borrei; mentum subretangular, transverse, with subparallel sides inI. tetraspilota and subtrapezoidal in I. borrei), pronotum (I. borrei twice wider than long and with depression; I. tetraspilota 1.63 to 1.74 times wider than long and without depression), proendosternite (projections posteriorly directed slightly curved in I. tetraspilota and linear in I. borrei), mesonotum (mesoscutellum more broadly rounded in I. tetraspilota, with sides strongly convergent in I. borrei), metanotum (wider than long in I. borrei and as wide as long in I. tetraspilota), postnotum (lateral projected distally in I. borrei and not projected in I. tetraspilota), metendosternite (angular at the apex in I. tetraspilota and straight in I. borrei), epipleura (I. borrei large to the basal half; I. tetraspilota large to the basal third), and hindwings (Costa short, less than half of Subcosta in I. tetraspilota and elongated inI. borrei, at least two thirds of Sc; Posterior Radial and Anterior Anal thinned in I. borrei).

Furthermore, regarding the genitalia: 1) basal third of aedeagus strongly curved inI. borrei and slightly curved in I. tetraspilota, 2) tegmen V-shaped, with rounded base with a projection in I. borrei, and U-shaped, truncated base with a projection inI. tetraspilota, 3) vaginal palpi slightly to moderately separated from the apical third in I. borrei and widely separated in the apical half I. tetraspilota and 4) spermatheca nodulus with the same width as the rest of spermatheca and duct ending in the base in I. borrei, and nodulus strongly broader, cylindrical, and the spermathecal duct reaching the cornus in I. tetraspilota, providing information to revalidate Synbrotica after a further test with cladistical methods.

Morphological characters presented by the type species showed that the synonymy of genera performed by Aslam (1972) was premature, since he did not observe the type species of Synbrotica(subjective synonymy). The characteristics cited by Weise (1922) in original description were confirmed toIsotes, but cannot be considered diagnostic since they include other genera. As initially discussed, Bechyné & Bechyné (1969) reported six diagnostic features forSynbrotica, which did not include all species of the genus. Comparative and more comprehensive studies can reinstatement ofSynbrotica, only those species with the characteristics listed above remain, as pointed out by Bechyné & Bechyné.

Taxonomical remarks and synonymy of species

Isotes borrei (Baly, 1889)

(Figs 1-3, 9-11, 13, 15, 17, 19, 21, 24, 28, 30, 32, 34, 36, 39, 40, 42, 44, 46, 48, 50, 52, 54, 56, 57)

Diabrotica borreiBaly, 1889:253; Gahan, 1891a:449 (redescr.).

Diabrotica cruciataBaly, 1889:253 (non Diabrotica cruciataJacoby, 1887:547).

Diabrotica borrei [ab.] cruciata; Gahan, 1891a:449 (syn.).

Diabrotica crucigeraWeise, 1916:40 (new replacement name for D. cruciata Baly).

Diabrotica borrei [ab.] crucigera; Weise, 1924:28 (cat.); Blackwelder, 1946:680 (cat.).

Synbrotica borrei; Bechyné, 1955:5; Bechyné, 1958:573; Bechyné & Bechyné, 1962:50 (list); Smith & Lawrence, 1967:44, 153 (type-species); Bechyné & Bechyné, 1969:72 (list); Wilcox, 1972:379 (cat.).

Synbrotica crucigera; Bechyné, 1955: 7 (stat. rev.); Bechyné, 1958:573; Bechyné & Bechyné, 1962:30; 1969:72 (list); Smith & Lawrence, 1967:53 (type-species).

Isotes borrei; Wilcox, 1975:690 (cat.).

Isotes crucigera; Wilcox, 1975:690 (cat.). Syn. nov.

In the original descriptions, these species were considered as different species by the following characteristics (I. borrei versus I. crucigera): (1) body shape narrowly oblong-oval vs. oblong oval, (2) body slightly posteriorly enlarged vs. posteriorly enlarged, (3) apex of antenna yellow vs. articles 9 and 10 and base of the 11 whitish, (4) thorax yellow vs. thorax (this tinged dark) yellow, (5) thorax wide, with two foveae vs. thorax subarcuate excavated, groove more deeply impressed on both sides, (6) elytra punctured, margins, apex dilated, wide band near the middle, suture and apex yellow vs. yellow elytra with two large black stripes.

Isotes borrei (Baly) and I. crucigera (Weise) are synonymized in this paper by having the external morphology, the aedeagus (Figs 40, 42, 44, 46) and female genitalia identical (Figs 48, 50, 52, 54), varying only on the sclerotization of bursa copulatrix (Figs 56, 57), and the basal spot of elytra (reaching the elytral suture and scutellum in I. borrei, and not in I. crucigera) (Figs 1-3). Both these features vary among specimens of the same species and specimens with intermediate characteristics were observed.

Acknowledgements.

We thank Dr. Cheryl B. Barr (EMEC), Dr. Márcio E. Felix (CEIOC), Dr. Marcela L. Monné (MNRJ), Dr. Sônia A. Casari (MZSP), Dr. David G. Furth (USNM) and Dr. Johannes Frisch (ZMHB) for the studied material, and the Laboratório de Ecologia de Insetos (UFRJ) to make available the stereomicroscope for taking the photographs. Special thanks to Luciano A. Moura (MCNZ, Museu Ciências Naturais, Fundação Zoobotânica, Porto Alegre, Brazil), who taught Chrysomelidae and review an earlier version of this manuscript. This research was conducted with the aid of FAPERJ and CNPq (Proc. 101.476/2010; 111.389/2010 and 470980/2011-7) for financial support.

REFERENCES

Aslam, N. A. 1972. On the genus Drasa Bryant with some nomenclatural notes on the GalerucinaeJournal of Natural History 6:483-501. [ Links ]

Baly, J. S. 1865. Descriptions of new genera and species of Phytophaga. Transactions of the Entomological Society of London 2:333-357. [ Links ]

_____. 1889. Diagnoses of uncharacterized species ofDiabrotica. The Entomologist's Monthly Magazine 25(10):251-254. [ Links ]

Bechyné, J. 1955. Troisième note sur lez Chrysomeloidea néotropicaux des collections de l'Institut royal des Sciences naturelles de Belgique. Institut Royal des Sciences Naturelles de Belgique 31(5):1-23. [ Links ]

_____. 1956. Reise des Herrn G. Frey in Südamerika: Galerucidae (Col. Phytophaga). Entomologische Arbeiten aus dem Museum Georg Frey 7(1):241-358. [ Links ]

_____. 1958. Notizen zu den neotropischen Chrysomeloidea (Col. Phytophaga). Entomologische Arbeiten aus dem Museum Georg Frey 9:478-706. [ Links ]

Bechyné, J. & Bechyné, B. 1962. Liste der bisher in Rio Grande do Sul gefundenen Galeruciden. Pesquisas Zoologia 15:5-68. [ Links ]

_____. 1969. Die Galerucidengattungen in Südbrasilien. Iheringia, Série Zoologia (36):1-110. [ Links ]

_____. 1970. Beiträge zur Kenntnis der Insektenfauna Boliviens XX. Coleoptera XV. Beiträge zur Galerucidenfauna Boliviens (Col. Phytophaga). Veröffentlichugen der Zoologischen Staatssamlung München 14:121-190. [ Links ]

Blackwelder, R. E. 1946. Checklist of the Coleopterus insects of Mexico, Central America the West Indies, and South America. Part 4. Bulletin of the United States National Museum 185(4):551-763. [ Links ]

Borowiec, L. & Pomorska, J. 2009. The Structure of the Spermathecae of the Genus Stolas (Coleoptera: Chrysomelidae: Cassidinae: Mesomphaliini) and Its Taxonomic Significance. Annales Zoologici 59(2):201-221. [ Links ]

Borowiec, L. & Skuza, M. 2004. The structure of spermatheca in the genus Chelymorpha Chevrolat, 1837 (Coleoptera: Chrysomelidae: Cassidinae) and its taxonomic significance. Annales Zoologici54:439-451. [ Links ]

Cabrera, N. C. & Cabrera Walsh, G. 2004a. Platybrotica misionensis a New Genus and Species of Luperini (Coleoptera: Chrysomelidae: Galerucinae) from Argentina. Annals of the Entomological Society of America 97(1):6-14. [ Links ]

_____. 2004b. Diabrotica calchaqui, a new species of Luperini (Coleoptera: Chrysomelidae: Galerucinae), from Argentina. Annals of the Entomological Society of America 97(5):889-897. [ Links ]

_____. 2010. Diabrotica collicola (Coleoptera: Chrysomelidae), a new species of leaf beetle from Argentina and key to species of the Diabrotica virgifera group and relatives. Zootaxa 2683:45-55. [ Links ]

Cabrera, N. C. & Durante, S. P. 2003. Comparative morphology of mouthparts in species of the genus Acalymma Barber (Coleoptera, Chrysomelidae, Galerucinae). The Coleopterists Bulletin 57(1):5-16. [ Links ]

Cabrera, N.; Sosa, A. J.; Dorado, J. & Julien, M. 2005.Systena nitentula (Coleoptera: Chrysomelidae), a Flea Beetle Injurious to Alternanthera philoxeroides(Amaranthaceae): Redescription, Biology, and Distribution. Annals of the Entomological Society of America98(5):643-652. [ Links ]

Cabrera, N. C.; Sosa-Gomez, D. R. & Micheli, A. 2008. Morphological and molecular characterization of a new species ofDiabrotica. Zootaxa 1922:33-46. [ Links ]

Chaboo, C. S. 2007. Biology and phylogeny of the Cassidinae Gyllenhal sensu lato (tortoise and leaf-mining beetles) (Coleoptera: Chrysomelidae). Bulletin of the American Museum of Natural History 305:1-250. [ Links ]

Chamorro-Lacayo, M. L. & Konstantinov, A. S. 2009. Synopsis of warty leaf beetle genera of the World (Coleoptera, Chrysomelidae, Cryptocephalinae, Chlamisini). ZooKeys 8:63-88. [ Links ]

_____. 2011. Cachiporrini, a remarkable new tribe of Lamprosomatinae (Coleoptera, Chrysomelidae) from South America. ZooKeys 78:43-59. [ Links ]

Chamorro-Lacayo, M. L.; Konstantinov, A. S. & Moseyko, A. G. 2006. Comparative morphology of the female genitalia and some abdominal structures of Neotropical Cryptocephalini (Coleoptera: Chrysomelidae: Cryptocephalinae). The Coleopterists Bulletin60(2):113-134. [ Links ]

Crowson, R. A. 1955. The natural classification of the families of Coleoptera. London, Nathaniel Lloyd. 187 p. [ Links ]

Forbes, W. T. M. 1922 The wing venation of the Coleoptera. Annals of the Entomological Society of America15:328-352. [ Links ]

Gahan, C. J. 1891a. On the South American species ofDiabrotica. Part II. Transactions of the Entomological Society of London1891:1-524. [ Links ]

_____. 1891b. Notes on some species of Diabrotica.Proceedings of the Entomological Society of London 1890:xliii [ Links ]

Hamilton, K. G. A. 1972. The insect wing, part II. Vein homology and the archetypal insect wing. Journal of the Kansas Entomological Society 45:54-58. [ Links ]

Hazmi, I. R. & Wagner, T. 2010. Revision ofArcastes Baly, 1865 from the Oriental Region (Coleoptera, Chrysomelidae, Galerucinae). Zookeys 42:79-99. [ Links ]

Jacoby, M. 1887. Biologia Centrali-Americana, Insecta, Coleoptera, Galerucidae. London. v. 6, part. 1, p. 497-584. [ Links ]

Jolivet, P. 1957. Recherches sur l'aile des Chrysomelidae (Col.). Mémoires Institut Royal des Sciences Naturelles de Belgique2(51):1-80. [ Links ]

Konstantinov, A. S. 1998. On the structure and function of the female genitalia in flea beetles (Coleoptera: Chrysomelidae: Alticinae). Proceedings of the Entomological Society of Washington 100(2):353-360. [ Links ]

_____. 2002. A new genus of flea beetles from the Greater Antilles (Coleoptera: Chrysomelidae). Zootaxa 124:1-24. [ Links ]

Kukalová-Peck, J. & Lawrence, J. F. 1993. Evolution of the hind wing in Coleoptera. The Canadian Entomologist 125:181-258. [ Links ]

_____. 2004. Relationships among coleopteran suborders and major endoneopteran lineages: evidence from hind wing characters. European Journal of Entomology 101(1):95-144. [ Links ]

Lawrence, J. F.; Slipinski, A.; Seago, A. E.; Thayer, M. K.; Newton, A. F. & Marvaldi, A. E. 2011. Phylogeny of the Coleoptera based on morphological characters of adults and larvae. Annales Zoologici 61(1):1-217. [ Links ]

Leonardi, C. 1970. Materiali per uno studio filogenetico del generePsylliodes (Coleoptera Chrysomelidae). Atti della Società italiana di Scienze naturali e del Museo civico di Storia naturale di Milano 110(3):201-223. [ Links ]

Lingafelter, S. W. & Konstantinov, A. S. 1999. The morphology and relative rank of alticine and galerucine leaf beetles: a cladistic analysis using adult morphological characters (Coleoptera: Chrysomelidae). Entomologica Scandinavica 30(4):397-416. [ Links ]

Marinoni, r. C. & Almeida, L. M. 1983. Sobre a venação alar em Coccinellidae e Cerambycidae (Coleoptera). Revista Brasileira de Entomologia 27(3/4):267-275. [ Links ]

Matsumura, Y. & Suzuki, K. 2008. Comparative morphology of internal reproductive systems in leaf beetles of the Donaciinae and Criocerinae (Coleoptera: Chrysomelidae) and its implication for the phylogeny. Zootaxa 1845:1-32. [ Links ]

Middelhouve, J. & Wagner, T. 2001. Revision ofAfrocrania (Coleoptera: Chrysomelidae, Galerucinae). Part I: Species in which the males have head cavities or extended elytral extrusions. European Journal of Entomology98(4):511-532. [ Links ]

Moura, L. de A. 1998a. Novo status deChlorolochmaea (Coleoptera, Chrysomelidae, Galerucinae)., Iheringia, Série Zoologia 84:145-152. [ Links ]

_____. 1998b. Iucetima, gênero novo de Galerucini da Região Neotropical (Coleoptera, Chrysomelidae, Galerucinae). Iheringia, Série Zoologia (85):75-88. [ Links ]

_____. 1998c. Revisão do gênero Neolochmaea(Coleoptera, Chrysomelidae, Galerucinae, Galerucini). Iheringia, Série Zoologia (85):169-188. [ Links ]

_____. 2003. Nova espécie de Zischkaita Bechyné e notas taxonômicas em Galerucini (Coleoptera, Chrysomelidae, Galerucinae). Revista Brasileira de Zoologia 20(3):643-645. [ Links ]

_____. 2007. Nota sobre o gênero ChlamophoraChevrolat, com a descrição de uma nova espécie do sul do Brasil (Coleoptera, Chrysomelidae, Galerucinae). Revista Brasileira de Zoologia 24(3):552-558. [ Links ]

_____. 2008. Análise cladística dos gêneros da Seção Schematizites (Coleoptera, Chrysomelidae, Galerucinae, Galerucini). Tese de doutorado, Universidade Federal do Rio Grande do Sul, Porto Alegre. 97p. [ Links ]

_____. 2009. Morfologia comparada da genitália masculina de Galerucini (Coleoptera, Chrysomelidae, Galerucinae). Revista Brasileira de Entomologia 53:15-22. [ Links ]

Nadein, K. S. 2006. A significance of the tegmen structure for classification of the genus Psylliodes Latreille, 1829 (Coleoptera: Chrysomelidae: Psylliodina). Proceedings of the Russian Entomological Society 77:250-254. [ Links ]

Napp, D. S. 1994. Phylogenetic relationships among the subfamilies of Cerambycidae (Coleoptera - Chrysomeloidea). Revista Brasileira de Entomologia38(2):265-419. [ Links ]

Powell, E. F. 1941. Relationships within the Family Chrysomelidae (Coleoptera) as indicated by the male genitalia of certain species. American Midland Naturalist 25:148-195. [ Links ]

Reid, C. A. M. & Nally, S. C. 2008. Revision of the genusMenippus Clark in Australia (Coleoptera: Chrysomelidae: Galerucinae). Australian Journal of Entomology 47:87-101. [ Links ]

Rodriguez, V. 1994. Function of the spermathecal muscle inChelymorpha alternans Boheman (Coleoptera: Chrysomelidae; Cassidinae). Physiological Entomology 19:198-202. [ Links ]

Savini, V. & Joly, L. J. 2006. Morfología de las mandíbulas de algunos géneros de Alticinae y Galerucinae (Coleoptera: Chrysomelidae). Entomotropica 21(1):23-40. [ Links ]

Smith, R. F. & Lawrence, J. F. 1967. Clarification of the status of the type specimens of Diabroticites (Coleoptera, Chrysomelidae, Galerucinae). University of California Publications in Entomology 45:1-168. [ Links ]

Stapel, H.; Misof, B. & Wagner, T. 2008. A molecular and morphological phylogenetic analysis of Afrotropical Monoleptaspecies and related Galerucinae (Coleoptera: Chrysomelidae). Arthropod Systematics & Phylogeny 66(1):3-17. [ Links ]

Suzuki, K. 1969. Comparative morphology and evolution of the hind wings of the Family Chrysomelidae (Coleoptera). 1. Homology and nomenclature of the wing venation in relation to allied families. Kontyu 37(1):32-40. [ Links ]

Torre-Bueno, J. R. de la. 1989. The Torre-Bueno Glossary of Entomology. New York, New York Entomological Society. 849p. [ Links ]

Wagner, T. 2002. Revision of Afrotropical Monoleptaspecies (Coleoptera, Chrysomelidae, Galerucinae) Part III: Species with red elytra and yellow prothorax, including descriptions of new species. Mitteilungen aus dem Naturkundemuseum Berlin, Deutsche Entomologische Zeitschrift 49(1):27-45. [ Links ]

Weise, J. 1916. Synonymische Mitteilugen. Deutsche Entomologische Zeitschrift 1916:37-41. [ Links ]

_____. 1922. Chrysomeliden der Indo-Malayischen Region. Tijdschrift voor Entomologie 65:39-130. [ Links ]

_____. 1924. Coleopterorum Catalogus Pars 78,1. Chrysomelidae: 13. Galerucinae. s'Gravenhage, W. Junk. 233p. [ Links ]

Wilcox, J. A. 1972. Coleopterorum Catalogus, Supplementa Pars 78,2. Chrysomelidae, Galerucinae, Luperini, Aulacophorina, Diabroticina. s'Gravenhage, W. Junk. 211p. [ Links ]

_____. 1975. Coleopterorum Catalogus, Supplementa. Pars 78,4. Chrysomelidae: Galerucinae, Addenda et Index. s'Gravenhage, W. Junk. 104p. [ Links ]

Received: June 11, 2015; Accepted: November 30, 2015

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