ZooKeys 332: 33–54, doi: 10.3897/zookeys.332.4931
Review on the use of sexually dimorphic characters in the taxonomy of Diabroticites (Galerucinae, Luperini, Diabroticina) 1
Laura Rocha Prado 1
1 Museu de Zoologia da Universidade de São Paulo, Av. Nazaré, 481, CEP 04263–000, São Paulo-SP, Brazil

Corresponding author: Laura Rocha Prado (laurarochaprado@gmail.com)

Academic editor: J. Santiago-Blay

received 1 March 2013 | accepted 7 May 2013 | Published 19 September 2013

(C) 2013 Laura Rocha Prado. This is an open access article distributed under the terms of the Creative Commons Attribution License 3.0 (CC-BY), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

For reference, use of the paginated PDF or printed version of this article is recommended.

Citation: Prado LR (2013) Review on the use of sexually dimorphic characters in the taxonomy of Diabroticites (Galerucinae, Luperini, Diabroticina). In: Jolivet P, Santiago-Blay J, Schmitt M (Eds) Research on Chrysomelidae 4. ZooKeys 332: 33–54. doi: 10.3897/zookeys.332.4931


Sexual dimorphism occurs frequently in Chrysomelidae Latreille, 1802 and is particularly variable in subfamily Galerucinae Latreille, 1802. This diversity has been early noted by authors a potential source of taxonomic characters. The section Diabroticites (Luperini Gistel, 1848) is one of the largest assemblies of chrysomelid genera with currently 823 valid species in 17 genera (12 based on dimorphic characteristics), being most diverse in the neotropical region. Apart from a revision work on the type specimens for the section, there are no general taxonomic studies for this group. The occurrence of sexually dimorphic characteristics in the section Diabroticites is revised and their practical taxonomic relevance evaluated. A total of 240 species was studied (145 species with males available), representing 15 out of the 17 genera included in Diabroticites. The analysis of characters was based on the study of specimens in south-american collections, literature and the aid of photos in online databases. Sexual dimorphism occurred in most species analyzed. Dimorphic features were divided in general (i. e., occur in higher taxa) and special characters (those that support the definition of species and genera). Special dimorphism was observed in every tagma, and most modifications occur in antennae. Characters used as diagnostic of genera often do not correspond to the modifications present in species included in them. Many modifications were considered by earlier authors as a single character, probably due to vague definitions. Most generic definitions are, therefore, inaccurate. The study of morphology and the homology assessment of characters are needed to increase understanding of the genera in Diabroticites.


Taxonomy of Coleoptera, rootworms, review, leaf-beetles


Sexual dimorphism has always been a subject of great curiosity amongst naturalists. Since Darwin’s suggestion of his Theory of Sexual Selection, many explanations have been proposed, and several have been successfully tested, for the existence of often peculiar modifications in males. Even though most beetles lack conspicuous sexual dimorphism (Kawano 2006), there are striking examples in all major Coleoptera taxonomic groups (Eberhard 2009). Most sexually dimorphic characters in beetles are described by strongly positive allometries (Kawano 2006), and also modifications found in antennae, tarsi, posterior legs and ventrites (Crowson 1981). Other less common characteristics include the reduction of the wings (Thayer 1992) and the presence of luminescent (Branham and Wenzel 2003) or stridulatory organs (Jansson and Selander 1977).

In Chrysomelidae Latreille, 1802, sexual dimorphism is thought to occur more frequently at the species level (Jolivet and Verma 2002). Common dimorphic features that are the body size (with females usually bigger than males) and the modification of tarsi, usually related to greater adhesion of males to the females’ dorsal surface during copulation (Jolivet and Verma 2002, Hammack and French 2007, Voigt et al. 2008, Nardi et al. 2012).

Within the subfamily Galerucinae Latreille, 1802, sexual dimorphism is particularly variable, as Mohamedsaid and Furth (2011) have illustrated and summarized. This diversity has been early noted by authors as a potential source of taxonomic characters. As Horn (1893) pointed out, many taxonomic issues related to this group could be resolved with the aid of “sexual peculiarities”, which could be a useful guide for understanding the relationships between species. Blake (1958) also stated that the use of such characteristics could help the delimitation of genera in problematic groups such as the tribe Luperini Gistel, 1848.

The section Diabroticites Chapuis, 1875 (Luperini) is one of the largest assemblies of chrysomelid genera with over 900 recorded names in 17 genera (12 based on dimorphic characteristics), being most diverse in the neotropical region. Apart from a revision work on the type specimens for the section (Smith and Lawrence 1967), there are no general taxonomic studies for this group. The most recent catalogue mentions 793 species (Wilcox 1972), but a review of the subsequent literature reveals that the group has currently 823 valid species. Table 1 presents an overview on the current composition of Diabroticites. Prior to 1906, Diabroticites included Diabrotica Chevrolat, 1837 (almost half of the total species in the section), and three other genera, which were all monotypic: Ensiforma Jacoby, 1876, Pseudodiabrotica Jacoby, 1892, and Paratriarius Schaeffer, 1906. Barber (1947) was the first to investigate male genital characters to understand the relationships within the group and described two more genera in the section, Acalymma Barber, 1947 and Amphelasma Barber, 1947. In the subsequent years, the remaining 11 genera were erected, eight of which based on species formerly included in Diabrotica (Anisobrotica Bechyné & Bechyné, 1969, Aristobrotica Bechyné, 1956, Buckibrotica Bechyné & Bechyné, 1969, Cochabamba Bechyné, 1955, Cornubrotica Bechyné & Bechyné, 1969, Gynandrobrotica Bechyné, 1955, Synbrotica Bechyné, 1956, and Paranapiacaba Bechyné, 1958), and most supported primarily on dimorphic features (Table 1). Surprisingly enough, only one genus has a detailed description of genital characters (the most recent genus, Platybrotica Cabrera & Cabrera Walsh, 2004).

Table 1.

Overview on the composition of genera of Diabroticites. The total number of analyzed species includes specimens in collections and online type-specimens in MCZ database.

Genus Dimorphism as diagnostic Number of species in original description Current number of species Number of analysed species (% of genus total)
Acalymma Barber, 1947 6 72 28 (38%)
Amphelasma Barber, 1947 5 11 2 (18%)
Anisobrotica Bechyné & Bechyné, 1969 X 1 5 5 (100%)
Aristobrotica Bechyné, 1956 X 10 17 3 (17%)
Buckibrotica Bechyné & Bechyné, 1969 X 1 1 1 (100%)
Cochabamba Bechyné, 1955 4 10 10 (100%)
Cornubrotica Bechyné & Bechyné, 1969 X 1 2 2 (100%)
Diabrotica Chevrolat, 1837 103 360 114 (31%)
Ensiforma Jacoby, 1876 X 1 9 3 (33%)
Gynandrobrotica Bechyné, 1955 X 23 32 4 (12%)
Isotes Weise, 1922 (= Synbrotica Bechyné, 1956) X 1(51) 182 38 (20%)
Palmaria Bechyné, 1956 X 1 1 -
Paranapiacaba Bechyné, 1958 X 16 59 14 (23%)
Paratriarius Schaeffer, 1906 X 1 51 11 (21%)
Platybrotica Cabrera & Cabrera Walsh, 2004 X 1 1 1 (100%)
Pseudodiabrotica Jacoby, 1892 X 1 1 -
Zischkaita Bechyné, 1956 1 9 4 (44%)
Total 12 177 823 240 (28%)

About 80% of the diabroticites species have been described prior to 1895, mainly by Joseph Sugar Baly, Charles J. Gahan and Martin Jacoby (Smith and Lawrence 1967). Most of those descriptions lack detailed morphological information, and usually depict characters relative to color pattern and, sometimes, punctuation. For many of the genera this is also true, with internal characters being almost completely ignored. General morphology has also been vaguely treated, described usually without any aid of illustrations whatsoever. Such scarcity of information and the apparent uniformity in morphology of some diabroticites resulted in a difficult taxonomic scenario.

The purpose of this study is to summarize the occurrence of sexually dimorphic characteristics in the section Diabroticites, as well as to review these characters chosen by earlier authors to support their definitions of genera and evaluate their practical taxonomic relevance. Systematic research is being conducted on Diabroticites by the author, and the first results are reported here.


The analysis of characters was based on the study of specimens, literature (original descriptions and revision works, when available) and the aid of photos in online databases. A total of 240 species was studied, representing 15 out of the 17 genera included in Diabroticites.

Specimens were obtained from south-american collections listed in Table 2, always in comparison to original descriptions – and many types were available. Out of the total species available, only 145 species had males available or known. These taxa were listed in Appendix. The specimens were examined and illustrated using a Zeiss Discovery.V8 stereomicroscope with a camera lucida attached. Final art was done in Adobe Illustrator®. Photographs were taken using a Leica M205C stereomicroscope with an attached magnifying lens and Leica DFC 295 video camera. Image combination was performed with Leica Application Suite V3.6.0, and subsequent edition was done in Adobe Photoshop®.

Table 2.

Institutions that provided specimens for the study.

Acronym Name City Country Curator
CEAH Coleção Entomológica Adolph Hempel, Instituto Biológico São Paulo Brazil Sérgio Ide
INPA Coleção Sistemática de Entomologia, Instituto Nacional de Pesquisas da Amazônia Manaus Brazil Augusto Henriques
FIOC Fundação Instituto Oswaldo Cruz Rio de Janeiro Brazil Jane Costa von Sydow
IACC Instituto Agronômico de Campinas Campinas Brazil Édson Possidônio Teixeira
MGAP Museu Anchieta Porto Alegre Brazil Fernando Meyer
MCNZ Museu de Ciências Naturais da Fundação Zoo-Botânica do Rio Grande do Sul Porto Alegre Brazil Maria Helena Galileo
DZUP Coleção de Entomologia Padre Jesus Moure, Universidade Federal do Paraná Curitiba Brazil Lúcia Massuti de Almeida
MZSP Museu de Zoologia da Universidade de São Paulo São Paulo Brazil Sônia Casari
MNRJ Museu Nacional, Universidade Federal do Rio de Janeiro Rio de Janeiro Brazil Marcela Monné
MPEG Museu Paraense Emilio Goeldi Belém Brazil Orlando Tobias Silveira
UFVB Museu Regional de Entomologia da Universidade Federal de Viçosa Viçosa Brazil Paulo Sérgio Fiuza Ferreira
MLPA Universidad Nacional de La Plata, Museo de la Plata La Plata Argentina Nora Cabrera

Most taxonomic literature available for Diabroticites was reviewed. The original descriptions of monotypic genera Palmaria Bechyné, 1956, and Pseudodiabrotica, known only for their type-specimens, which could not be loaned, were the only source of characters for comparison. Revisionary works were available only for genera Acalymma (in part) (Munroe and Smith 1980, Cabrera 1999, Cabrera and Durante 2003), Diabrotica (in part) (Marques 1941, Christensen 1943, Krysan and Smith 1987, Cabrera 2000a, Cabrera 2000b) and Synbrotica (in part) (Cabrera 1995).

Due to the peculiarity of genus Isotes Weise, 1922, which was described based on a single species later found to be a senior synonym of the type-species of genus Synbrotica (at that time with over 100 species), the original description of the latter was also included in the analysis for comparative purposes. As a reference to their original descriptions, both names will be used interchangeably throughout the text, even though Isotes is the current valid name.

Characters mentioned in original descriptions and other taxonomic works, when available, were compiled and later compared to specimens. Those characters were then redefined, in order to fulfill uniform homology criteria. The broader studies of Mohamedsaid and Furth (2011) and Mohamedsaid (2004) were used for character comparison with other taxonomic groups.

Also, some taxa that had not enough specimens available in south-american collections were studied by the analysis of photos of type specimens available in the Museum of Comparative Zoology online Type Database, Harvard University, Cambridge, USA.

Results and discussion

Out of the 17 genera that comprise section Diabroticites, 12 have sexually dimorphic features as diagnostic characters mentioned in original descriptions by nine different authors (15 papers) in a period of more than 150 years (Table 1). Most genera were established based on a single or on few species, further taxa being later added, frequently not by their original authors. This information is relevant when checking whether the initial concept proposed for the genus was maintained or not.

Most species studied have some kind of sexual dimorphism. Those modifications were categorized in two distinct groups, concerning its level of taxonomic comprehension: general and special dimorphism.

General dimorphism

Characters that are referred to as of general comprehension are those proposed several times in the literature as being important to species definition, but were actually found to be more generalized, i. e., they in fact occur in higher taxa. General characters are found in all, or most, male of diabroticites analyzed, and might also support the definition of larger taxonomic groups: 1) Smaller body size: considered general for Chrysomelidae, observed for most diabroticites; 2) Bigger eyes (relative to the total size of the head): cited often in species descriptions, but actually observed in most diabroticites; 3) Tarsal adhesive disks: structures present in most Chrysomelidae, with variation found among subfamilies and often among tribes, regarding the number of legs in which they occur and the proportion of the dorsal surface that they occupy (Stork 1980). For Diabroticites, the adhesive disks are present at the first tarsomere in pro- and mesothoracic legs; 4) Emargination on the posterior margin of ventrite V: this characters sometimes supports the definition of the subfamily Galerucinae, and its shape can be diagnostic of tribes, being rounded in Luperini and Galerucini, for example (Bechyné and Bechyné 1962, Wilcox 1965). In diabroticites this emargination is, usually, round; and 5) Prothoracic tibiae with continuous apex, without spurs: probably constant in the whole tribe (Wilcox 1965).

Special dimorphism

Special characters are those used to describe lower taxonomic ranks, i. e., genera and species, and have been or not mentioned as diagnostic features in original descriptions.

With the exception of two genera, Amphelasma Barber, 1947 and Cochabamba all genera in Diabroticites are represented with special dimorphic characters. Of all species analyzed, only 47 were found to have special dimorphic characteristics. This is interesting, since the original definitions of diabroticites genera were usually based in sexual dimorphism. In accordance with the pattern observed for the subfamily (Mohamedsaid and Furth 2011), special dimorphism was observed in every tagma in the analyzed taxa of Diabroticites (Table 3).

Table 3.

Location of diagnostic, sexually dimorphic characters mentioned in original descriptions.

Genus Antennae Legs Head (except antennae) Elytra
Anisobrotica Bechyné & Bechyné X X
Aristobrotica Bechyné X X
Paranapiacaba Bechyné X X
Buckibrotica Bechyné & Bechyné X
Cornubrotica Bechyné & Bechyné X
Ensiforma Jacoby X
Paratriarius Schaeffer X
Platybrotica Cabrera & Cabrera Walsh X
Isotes Weise (ex Synbrotica Bechyné) X*
Palmaria Bechyné X
Gynandrobrotica Bechyné X
Pseudodiabrotica Jacoby X

*: present in the original description of Synbrotica.

Out of the 12 genera recognized by dimorphic characters, 4 are monotypic (Buckibrotica, Palmaria, Platybrotica and Pseudodiabrotica). The diagnostic characters and its validity will be discussed for each tagma.


Most of the dimorphic characters were found in the head, mainly in the antennae, a proportion which agrees with the general pattern observed in the subfamily (Mohamedsaid and Furth 2011).

Gynandrobrotica was described as showing an “excavated clypeus”, without any further details or illustrations. All four species studied have the same kind of modification, which is better described as the frons being elongated and with a shallow, smooth, round concavity (Figure 1), accompanied by sparse, large punctuation. It is interesting to compare this character with the differently excavated type of frons found in species of Cerotoma Chevrolat, 1837 (Figure 2), Eucerotoma Laboissiere, 1939 and Neobrotica Jacoby, 1887 – all of which are usually placed in sister-section Cerotomites Chapuis, 1875. Gynandrobrotica has been suggested to be more related to these taxa in some phylogenetic analyses (Eben and Monteros 2004, Gillespie et al. 2008). Other characteristics common to these genera, such as the small eyes, the elongated frons and different shapes of antennomeres I-III should be further studied in order to verify if they are indeed homologues.

There are also three other species with distinctive head features in the male: Isotes onira (Bechyné & Bechyné), 1961 has an enlarged head from the vertex up to the antennal insertions – Figure 3), Diabrotica serroazulensis Bechyné & Bechyné, 1962 has an enlarged labrum, which is as large as half of the frontal length, and Acalymma cornutum (Baly, 1886), has a deep cleft in the frons, with lateral projections, and also a labrum with an acute anterior projection that reaches the frons (illustrated in Munroe and Smith 1980).

Figure 1.

Gynandrobrotica caviceps (Baly, 1889), head in frontal view (female, left, male, right).

Figure 2.

Cerotoma variegata (Fabricius, 1792) head in frontal view (female, left, male, right).

Figure 3.

Isotes onira (Bechyné & Bechyné, 1961), dorsal view, detail of pronotum and head (female, left, male, right).


Galerucines commonly display filiform antennae, which can show numerous dimorphic variations (Jolivet and Verma 2002, Mohamedsaid 2004). This is also true for Diabroticites, with the main antenna type being filiform and antenommeres mostly subequal in size and shape (Figure 4A).

Non-dimorphic modifications are often related to the length of some antennomeres and sometimes support generic definitions. For instance, the two largest genera in Diabroticites, Diabrotica and Synbrotica are essentially distinguished by the length of antennomere III, described as being subequal to II in the first (same as observed in genus Cochabamba in Figure 4B), and subequal to IV (that is, almost twice as longer as II, as in Figure 4C) in the latter (Bechyné 1956).

Figure 4.

Male modified antennae in lateral view A Isotes borrei (Baly, 1889) B Cochabamba marginata (Ha-rold, 1875) C Isotes onira (Bechyné & Bechyné, 1961) D Aristobrotica angulicollis (Erichson 1878) E Buckibrotica cinctipennis (Baly, 1886) (detail in ventral view) F Ensiforma caerulea Jacoby, 1876. Scale bar = 1 mm.

Eight genera have diagnostic characters based in their dimorphism of antennae (listed in Table 3). Examples of modified antennae are showed in Figure 4C–F. Because of their variability, dimorphic antennae are the most used structure in descriptions, but their modifications have been scarcely detailed. As a result, there are different genera proposed on characters depicted in sentences such as “apical antennomeres modified” (Cornubrotica), “antennomeres VII and IX of complicated shape” (BuckibroticaFigure 4E), “antennomeres V-VII modified” (Paratriarius).

The absence of unified criteria in the understanding of what a “modified” antennomere is has lead, several times, to the establishment of artificial grouping of species, simply because a single “aberrant” antennomere can display an assembly of four different aspects of its morphology. Modifications include change in (in quoting marks, expressions used on original descriptions): length – antennomeres considered “elongated” or “shortened” when compared to the usually fixed antennomeres I and III; width – antennomeres described as “swollen”, “inflated” (homogeneous modification), “distally expanded” (heterogeneous modification), and dorsoventrally “flattened”; shape – those described as having “lateral projections” or “ventral excavations”; structure – antennomeres with “rough punctuation” and different amounts of hairs/sensillae. As variations observed in the species do not always correspond to the diagnosis defined for genera they have been included in, these characteristics do not provide an accurate guide to the identification of taxa in Diabroticites.

For the non-monotypical genera based on antennal dimorphic features, most original definitions do not correspond to their actual characters. In Anisobrotica, for example, the “widened” apical antennomeres do not always appear – Anisobrotica binisculpta Bechyné & Bechyné, 1969 only has in common with the other taxa the excavation present in glabrous ventral surface of antennomeres IX-XI (such as excavations observed in apical antennomeres of Anisobrotica donckieri (Baly, 1889) in Figure 5). The same happens with Paratriarius, which includes several species that do not show “modified antennomeres V-VII” present in type-species Paratriarius dorsata (Say, 1824) (illustrated in Wilcox 1965) such as Paratriarius batesi (Baly, 1859), Paratriarius falvolimbata (Erichson, 1847), Paratriarius verrucosa (Jacoby, 1880), Paratriarius alternans (Weise, 1916), Paratriarius nigrotibialis (Bowditch, 1911), Paratriarius castanea (Bowditch, 1911), and also other four species studied by Mohamedsaid and Furth (2011). Instead, these taxa show antennae very similar to the general pattern seen in Diabrotica. The two species included in Cornubrotica do not show identical antennomeres VIII and IX, although both always have ventral excavations (illustrated in Bechyné and Bechyné 1969 and Moura 2005). In Paranapiacaba, the antennal character chosen was, unfortunately, a general one: male antennomeres III-XI uniformly “thickened” (in contrast with slightly slender antennae of females). Nevertheless, antennae do seem to vary uniformly in one genus. In Aristobrotica, the pattern of antennomeres III-V “thickened” is constantly repeated, followed by an unmentioned presence of larger punctuation (Figure 4D).

Figure 5.

Anisobrotica donckieri (Baly, 1889), detail of ventral surface of apical antennomeres, male.

Although Maulik (1936) suggested that, for indo-asian galerucines, the basal antennomeres are more frequently the altered ones, a result that has been corroborated by Mohamedsaid (2004), that feature does not apply to diabroticites analyzed. Also, no obvious topological pattern is seen in the variation of antennomeres (Table 4).

Table 4.

Selected diabroticites species representing antennal dimorphic variation. Grey cells indicate modification in the antennomere.

Taxon/antennomere I II III IV V VI VII VIII IX X XI
Isotes onira (Bechyné & Bechyné, 1961)                                                       
Ensiforma chiquitoensis (Bechyné, 1958)
Aristobrotica angulicollis (Erichson, 1878)
Isotes simplicipennis (Jacoby, 1889)
Ensiforma asteria (Bechyné & Bechyné, 1962)
Paratriarius dorsata (Say, 1824)
Isotes callanga (Bechyné, 1956)
Platybrotica misionensis Cabrera & Cabrera Walsh, 2004
Buckibrotica cinctipennis (Baly, 1886)
Cornubrotica dilaticornis (Baly, 1879)
Diabrotica samouella Bechyné, 1956
Anisobrotica donckieri (Baly, 1889)

The number of modified antennomeres oscillated between 1 to 6. Antennomere II was recorded as dimorphic only in species of Aristobrotica, such as Aristobrotica angulicollis (Erichson, 1878) (Figure 4D) and in Isotes onira (Figure 4C). Few modifications occur in antennomeres II and XI. The most affected are antennomeres V to IX. However, there is no indication of an explicit dependency of occurrence between any pair of modified antennomeres. This is the opposite of what has been observed for asian Galerucinae species (Mohamedsaid 2004). Although no pattern is observed, some variation can occur in blocks, i. e., one modified antennomere occurs with one or two adjacent antennomeres also modified.

It seems that most, if not all, antennal variations could be regarded as the result of the presence of punctuation and setae in greater number, either for the production and/or reception of chemical compounds (i. e., pheromones) (Jolivet 2007). A study on Diabrotica virgifera Leconte, 1858, for instance, showed that male antennae have a much greater number of sensilla than females, and numerous glandular points linked to the production of chemical compounds allegedly to be attractive to females (Newman Jr et al. 1993). Unfortunately, there is little knowledge on the biology or even on the anatomy of Diabroticites to support this view as a more generalized tendency.


Although a great diversity of dimorphic characters occur in the thorax of several galerucines (Mohamedsaid and Furth 2011), features reported for Diabroticites are limited to elytra, and legs. In elytra, variation occurs only on the apical fifth, and are either depressions or callosities, both which can co-occur with punctuation (as in some species of IsotesFigure 6, and in Pseudodiabrotica – the only genus supported on an elytral diagnostic character). Many species in genus Paratriarius show elytral dimorphic characters (such as the callosities present in Paratriarius batesi), and, although such features were never used to originally describe it, they have been used to support the definition of genus Chanchamayia Bechyné, 1956, now considered to be a subgenus in Paratriarius (Smith and Lawrence 1967, Wilcox 1972).

Figure 6.

Detail of elytral modifications, left, Isotes digna (Gahan, 1891), male, right, Paratriarius batesi (Baly, 1859), male.

Only metathoracic legs lack alterations in male diabroticites. In accordance with the more general pattern, morphological differences in the pro- and mesothoracic legs usually are connected to the augmentation of absolute size in femora and tarsomeres I, the latter which are directly linked to the partial or total covering of the ventral surface by adhesive setae (Figure 7).

Figure 7.

Paranapiacaba prolongata (Jacoby, 1882), detail of ventral surface of tarsomeres of prothoracic leg, male.

Tibiae and femora can also be modified, being greatly enlarged (such as in Zischkaita serrana Moura, 2003 – Figure 8), and frequently with internal margins concave or bearing tubercles, forming the “prehensile organ” (Bechyné 1956). Bechyné’s concept of such structure is based on a combination of multiple adaptations and should be used with caution, since the homology of the “prehensile organs” can be difficult to assess. Aristobrotica, for instance, has been described as with one diagnostic feature: the “special build of the median tibiae in male”. The detailed analysis of species included, however, indications that there are at least two distinct types of “prehensile organs” being treated as the same modification. While type-species Aristobrotica angulicollis (Figure 9) bears only a concave mesotibiae with laterally flattened apex, Aristobrotica mirapeua Moura, 1997 and Aristobrotica capillosa Moura, 2011 (both illustrated in their original descriptions) display small projections of the ventral margin of the mesofemora (apical in Aristobrotica mirapeua and basal in Aristobrotica capillosa), and differently shaped tibiae (with a basal concavity in Aristobrotica mirapeua and slightly concave tibiae with apical flattening in Aristobrotica capillosa).

Figure 8.

Zischkaita serrana Moura, 2003, ventral view, male.

Figure 9.

Aristobrotica angulicollis (Erichson, 1878), detail of mesothoracic leg, male.

Moreover, general characters have been used to support definition of genera such as Cornubrotica and Synbrotica, a genus which is now a synonym of Isotes. The former was supposed to be distinguished by pro- and mesothoracic legs of males without emargination, which is rather common in the tribe, and the latter is characterized by antennomere III elongated and a “uniform pilosity covering the ventral surface of tarsomeres in both sexes” (freely translated from the original, in German) – something that does not accurately identify the males in this group, as they normally have distinctive adhesive disks in their tarsomeres.


The most common abdominal modification seen in some galerucines is the presence of processes with different shapes. Although no abdominal characters aid the definition of diabroticites genera, one character was observed for a single species in the group: a central triangular projection, postero-ventrally oriented, in the posterior margin of the ventrite I, in Zischkaita serrana (Figure 10). A similar alteration is observable in Hemygascelis longicollis Jacoby, 1896, an asian species that belongs to section Phyllobroticites in subtribe Luperina, a group thought to be a sister group of Diabroticina (illustrated in Mohamedsaid and Furth 2011).

Figure 10.

Zischkaita serrana Moura, 2003, schematic abdomen in ventral view, male. Scale bar = 0, 5 mm.


The taxonomic history of taxa in section Diabroticites, which dates to over 150 years of specimen sampling and species descriptions, has been supported primarily by scarcely detailed descriptions of morphological features, specially coloration and striking “sexual aberrations”. Sexually dimorphic characters have been the foundation for the creation of many genera in this group, albeit with vague definitions and a general disregard for comparative morphology. The use of inappropriate terminology by some authors has caused further confusion, since the habit of inferring homology from homonymy is common in the taxonomy of Chrysomelidae (Schmitt, 1996). This might have been the origin of the combination of many species into these scarcely defined genera in Diabroticites. As a result, most generic original definitions, based on dimorphic characteristics, are not correspondent to their actual assembly of species.

Nevertheless, the establishment of homology theories is far from being trivial, and some variations might even occur in patterns that can actually help the definition of certain taxa, although it is clear that many sexually dimorphic features found in this section are possibly singular and autapomorfic. In this case, a broader morphological study is necessary.

The comparison of the dimorphic characters in Diabroticites with their related taxa, such as the Asiatic Aulacophorites Chapuis, 1875 (Luperini) and remaining Galerucinae is desired, in order to understand the evolution of such characters. There are striking similarities among many modifications found in these groups and thus it should be useful in the evaluation of homology as well. Parallelism might be the more parsimonious choice in many cases, but that is yet to be tested. Understanding the morphology is critical for better character definitions. Also, genital characters, which have been generally ignored, should provide important characters, as well as the account of several non-dimorphic characters, usually regarded as too uninformative, without detailed consideration.


The author thanks Sônia Casari (Museu de Zoologia da Universidade de São Paulo), Simeão Moraes and Juares Fuhrmann for reviewing earlier versions of this work and providing valuable suggestions, Bárbara Bitarello for English proofreading this manuscript, the reviewers for their careful reading, Michael Schmitt and LOC organization for the ICE Student Travel Award granted, and CAPES (PROEX) for the financial support provided, so that the author could attend the Biology of Chrysomelidae Symposium in Daegu, 2013. This study was funded by CNPq (134436/2008-3 and 140250/2011-5) and FAPESP (2008/02166-5).

Barber HS (1947) Diabrotica and two new genera (Coleoptera, Chrysomelidae). Proceedings of the Entomological Society of Washington 49(6): 151-161.
Bechyné J (1956) Reise des Herrn G. Frey in Südamerika: Galerucidae (Col. Phytophaga). Entomologische Arbeiten aus dem Museum Georg Frey 7(1): 241-358.
Bechyné J, Bechyné SB (1962) Liste der bisher in Rio Grande do Sul gefundenen Galeruciden. Pesquisas (Zool.) 6(15): 1-63.
Bechyné J, Bechyné SB (1969) Die Galerucidengattungen in Südbrasilien. Iheringia, Série Zoologia 36: 1-110.
Blake DH (1958) A review of some galerucine beetles with excised middle tibiae in the male. Proceedings of the United States National Museum 108(3395): 59-101. doi: 10.5479/si.00963801.108-3395.59
Branham MA, Wenzel JW (2003) The origin of photic behavior and the evolution of sexual communication in fireflies (Coleoptera: Lampyridae). Cladistics 19: 1-22. doi: 10.1016/S0748-3007(02)00131-7
Cabrera NC (1995) Nuevos aportes para el conocimiento del genero Synbrotica Bechyné (Coleoptera: Chrysomelidae: Galerucinae). Neotropica (La Plata) 41(105-106): 9-18.
Cabrera NC (1999) Contribución para el conocimiento del género Acalymma em la Argentina (Coleoptera: Chrysomelidae). Revista de la Sociedad Entomologica Argentina 48(3-4): 91-105.
Cabrera NC (2000a) Estudio sistematico de Diabrotica Chevrolat grupo fucata en la Argentina I (Coleoptera, Chrysomelidae). Physis, Sección C 58(134-135): 47-56.
Cabrera NC (2000b) Estudio sistematico de Diabrotica Chevrolat grupo fucata en la Argentina II (Coleoptera, Chrysomelidae). Physis, Sección C 58(134-135): 57-66.
Cabrera NC, Durante SP (2003) Comparative morphology of mouthparts in species of the genus Acalymma Barber (Coleoptera, Chrysomelidae, Galerucinae). The Coleopterists Bulletin 57(1): 5-16. doi: 10.1649/0010-065X(2003)057[0005:CMOMIS]2.0.CO;2
Christensen JR (1943) Estudio sobre el genero Diabrotica Chev. en la Argentina. Revista de la Facultad de Agronomía y Veterinaria de Buenos Aires 10: 464-516.
Crowson RA (1981) The biology of Coleoptera. Academic Press, London, 802 pp.
Eben A, Monteros AE de los (2004) Ideas on the systematics of the genus Diabrotica Wilcox and other related leaf beetles. In: Jolivet, Santiago-Blay, Schmitt (Eds) New Developments in the Biology of Chrysomelidae. SPB Academic Publishing, The Hague, Netherlands, 59-73.
Eberhard WG (2009) Postcopulatory sexual selection: Darwin’s omission and its consequences. Proceedings of the National Academy of Sciences of the United States of America 106 (suppl. 1): 10025–10032. doi: 10.1073/pnas.0901217106
Gillespie JJ, Tallamy DW, Riley EG, Cognato AI (2008) Molecular phylogeny of rootworms and related galerucine beetles (Coleoptera: Chrysomelidae). Zoologica Scripta 37(2): 195-222. doi: 10.1111/j.1463-6409.2007.00320.x
Hammack L, French BW (2007) Sexual Dimorphism of Basitarsi in Pest Species of Diabrotica and Cerotoma (Coleoptera: Chrysomelidae). Annals of the Entomological Society of America 100(1): 59-63. doi: 10.1603/0013-8746(2007)100[59:SDOBIP]2.0.CO;2
Horn GH (1893) The Galerucini of boreal America. Transactions of the American Entomological Society 20: 57-136.
Jansson A, Selander J (1977) Sound production associated with mating behaviour of the large pine weevil, Hylobius abietis (Coleoptera, Curculionidae). Annales Entomologici Fennici 43 (2): 66-75. doi: 10.1017/S0007485399000024
Jolivet P (2007) Hypertélie: mimétisme, signaux sexuels ou moyens de défense. Un dilemme chez les insectes: vrai ou faux concept? L’Entomologiste 63(2): 57–66. http://www.bio-nica.info/biblioteca/Jolivet2007Hypertelie.pdf
Jolivet P, Verma KK (2002) Biology of Leaf Beetles, 332 pp.
Kawano K (2006) Sexual dimorphism and the making of oversized male characters in beetles (Coleoptera). Annals of the Entomological Society of America 99(2): 327-341. doi: 10.1603/0013-8746(2006)099[0327:SDATMO]2.0.CO;2
Krysan JL, Smith RF (1987) Systematics of the virgifera species group of Diabrotica (Coleoptera: Chrysomelidae: Galerucinae). Entomography 5: 375-484.
Marques MA (1941) Contribuição ao estudo dos Crisomelídeos do gênero Diabrotica. Boletim da Escola Nacional de Agronomia 2: 1-57.
Maulik S (1936) The fauna of British India, including Ceylon and Burma. Coleoptera, Chrysomelidae, Galerucinae. Taylor and Francis, London, 648 pp.
Mohamedsaid MS (2004) Modified antennae of Malaysian Galerucinae and its taxonomic significance. In: Jolivet, Santiago-Blay, Schmitt (Eds) New Developments in the Biology of Chrysomelidae. SPB Academic Publishing, The Hague, Netherlands, 231-247.
Mohamedsaid MS, Furth DG (2011) Secondary Sexual Characteristics in the Galerucinae (Sensu Stricto) (Coleoptera: Chrysomelidae). ISRN Zoology 2011: 1-60. doi: 10.5402/2011/328670
Moura LA (2005) Novos táxons em Galerucini e redescrição de Caraguata circumcincta Clark (Coleoptera, Chrysomelidae, Galerucinae). Revista Brasileira de Zoologia 22(4): 1109-1115. doi: 10.1590/S0101-81752005000400042
Munroe DD, Smith RF (1980) A revision of the Systematics of Acalymma sensu sctricto Barber (Coleoptera: Chrysomelidae) from North America including Mexico. Memoirs of the Entomological Society of Canada 112: 1-92. doi: 10.4039/entm112112fv
Nardi C, Luvizotto Ra, Parra JRP, Bento JMS (2012) Mating behavior of Diabrotica speciosa (Coleoptera: Chrysomelidae). Environmental entomology 41(3): 562-70. doi: 10.1603/EN10284
Newman Jr SM, McDonald IC, Triebold B (1993) Antennal sexual dimorphism in Diabrotica virgifera virgifera (Le Conte) (Coleoptera: Chrysomelidae): male specific structures, ultrastructure of a unique sensillum, and sites of esterase activity. International Journal of Insect Morphology and Embryology 22(5): 535-547. doi: 10.1016/0020-7322(93)90039-4
Schmitt M (1996) The phylogenetic system of the Chrysomelidae - history of ideas and present state of knowledge. In: Jolivet, Cox (Eds) Chrysomelidae Biology, vol. 1: The Classification, Phylogeny and Genetics. SPB Academic Publishing, Amsterdam, 57-96.
Smith RF, Lawrence JF (1967) Clarification of the status of the type specimens of Diabroticites. University of California Press, 45.
Stork NE (1980) A scanning electron microscope study of tarsal adhesive setae in the Coleoptera. Zoological Journal of the Linnean Society 5(3): 173-306. doi: 10.1111/j.1096-3642.1980.tb01121.x
Thayer MK (1992) Discovery of sexual wing dimorphism in Staphylinidae (Coleoptera): “Omaliumflavidum, and a discussion of wing dimorphism in insects. Journal of the New York Entomological Society 100(4): 540–573. http://www.jstor.org/stable/25009991
Voigt D, Schuppert JM, Dattinger S, Gorb SN (2008) Sexual dimorphism in the attachment ability of the Colorado potato beetle Leptinotarsa decemlineata (Coleoptera: Chrysomelidae) to rough substrates. Journal of Insect Physiology 54: 765-76. doi: 10.1016/j.jinsphys.2008.02.006
Wilcox J A (1965) A synopsis of the North American Galerucinae (Coleoptera: Chrysomelidae). Bulletin of the New York State Museum and Science Service 400: 1-226. doi: 10.1086/621376
Wilcox JA (1972) Chrysomelidae: Galerucinae, Luperini: Aulacophorina, Diabroticina. In: Junk (Ed) Coleopterorum Catalogus Supplementa. Netherlands, 221-431.

List of taxa with male specimens available in south-american collections.

Acalymma albidovittatum (Baly, 1889)

Acalymma bivittulum (Kirsch, 1883)

Acalymma bruchii (Bowditch, 1911)

Acalymma carinipenne (Bowditch, 1911)

Acalymma exigua Bechyné, 1958

Acalymma granulipenne (Bowditch, 1911)

Acalymma incum (Bowditch, 1911)

Acalymma innubum (Fabricius, 1775)

Acalymma isogenum Bechyné & Bechyné, 1968

Acalymma punctatum (Jacoby, 1887)

Acalymma rubeolum Bechyné, 1958

Acalymma thiemei (Baly, 1886)

Acalymma vitigera (Boheman, 1859)

Acalymma vittatum (Fabricius, 1775)

Acalymma xanthographum Bechyné, 1955

Anisobrotica binisculpta Bechyné & Bechyné, 1969

Anisobrotica donckieri (Baly, 1889)

Anisobrotica nordenskiöldi (Jacoby, 1907)

Anisobrotica notaticollis (Baly, 1889)

Aristobrotica angulicollis (Erichson, 1878)

Aristobrotica capillosa Moura, 2011

Buckibrotica cinctipennis (Baly, 1886)

Cochabamba chacoensis (Bowditch, 1911)

Cochabamba chrysopleura (Harold, 1875)

Cochabamba diversicolor (Baly, 1890)

Cochabamba erythrodera (Baly, 1879)

Cochabamba marginata (Harold, 1875)

Cochabamba mera Bechyné, 1956

Cochabamba polychroma Bechyné, 1956

Cochabamba rugulosa (Baly, 1886)

Cochabamba variolosa (Jacoby, 1878)

Cochabamba volxemi (Baly, 1889)

Cornubrotica dilaticornis (Baly, 1879)

Cornubrotica iuba Moura, 2005

Diabrotica alegrensis Bechyné & Bechyné, 1962

Diabrotica amoena (Dalman, 1823)

Diabrotica antonietta Bechyné, 1956

Diabrotica aracatuba Bechyné & Bechyné, 1964

Diabrotica arcuata Baly, 1859

Diabrotica atrilineata Baly, 1889

Diabrotica atromaculata Baly, 1889

Diabrotica atrosignata Baly, 1890

Diabrotica boggianii Bowditch, 1911

Diabrotica chloropus Harold, 1875

Diabrotica clarki Weise, 1916

Diabrotica confraterna Baly, 1889

Diabrotica consentanea Baly, 1886

Diabrotica cryptochlora Bechyné, 1956

Diabrotica decaspila Baly, 1890

Diabrotica decempunctata (Latreille, 1813)

Diabrotica deliqua Weise, 1921

Diabrotica distincta Jacoby, 1882

Diabrotica egleri Bechyné & Bechyné, 1961

Diabrotica elata (Fabricius, 1801)

Diabrotica emorsitans Baly, 1890

Diabrotica enae Marques, 1941

Diabrotica fallenia Bechyné, 1956

Diabrotica flava (Olivier, 1791)

Diabrotica funerea Bowditch, 1911

Diabrotica fusibilis Bechyné & Bechyné, 1970

Diabrotica gracilenta Erichson, 1847

Diabrotica graminea Baly, 1886

Diabrotica kirbyi Baly, 1890

Diabrotica lamiina Bechyné & Bechyné, 1969

Diabrotica limitata (Sahlberg, 1823)

Diabrotica lutescens Baly, 1890

Diabrotica manaensis Weise, 1921

Diabrotica nitidicollis Baly, 1889

Diabrotica olivacea Jacoby, 1882

Diabrotica panchroma Bechyné, 1955

Diabrotica paranaensis Marques, 1941

Diabrotica pentazyga Bechyné & Bechyné, 1970

Diabrotica piceicornis Baly, 1889

Diabrotica piceosignata Baly, 1890

Diabrotica poecilenta Bechyné, 1958

Diabrotica propylaea Bechyné & Bechyné, 1969

Diabrotica quinquemaculata (Fabricius, 1801)

Diabrotica recki Marques, 1941

Diabrotica rufolimbata Baly, 1879

Diabrotica samouella Bechyné, 1956

Diabrotica schaufussi Baly, 1890

Diabrotica scripta (Olivier, 1808)

Diabrotica sedata Baly, 1890

Diabrotica serroazulensis Bechyné & Bechyné, 1962

Diabrotica sharpii Kirsch, 1883

Diabrotica sheba Bechyné, 1958

Diabrotica simulata Baly, 1890

Diabrotica sinuata (Olivier, 1789)

Diabrotica speciosa (Germar, 1824)

Diabrotica stenocoryna Bechyné & Bechyné, 1970

Diabrotica tarcisia Bechyné, 1971

Diabrotica tijuquensis Marques, 1941

Diabrotica transversa Baly, 1890

Diabrotica travassosi Marques, 1941

Diabrotica univittata Jacoby, 1899

Diabrotica viridans Baly, 1889

Diabrotica viridimaculata Jacoby, 1878

Diabrotica viridula (Fabricius, 1801)

Diabrotica wartensis Cabrera & Sosa-Gómez, 2008

Diabrotica westwoodi Baly, 1889

Ensiforma asteria (Bechynné & Bechyné, 1962)

Ensiforma caerulea Jacoby, 1876

Ensiforma chiquitoensis (Bechyné, 1958)

Gynandrobrotica caviceps (Baly, 1889)

Gynandrobrotica equestris (Fabricius, 1787)

Isotes agatha (Bechyné & Bechyné, 1969)

Isotes albidocincta (Baly, 1889)

Isotes bertonii (Bowditch, 1912)

Isotes bicincta (Bowdtich, 1912)

Isotes borrei (Baly, 1889)

Isotes brasiliensis (Jacoby, 1888)

Isotes cargona (Bechyné, 1958)

Isotes caryocara (Bechyné, 1956)

Isotes crucigera (Weise, 1916)

Isotes delicula (Erichson, 1847)

Isotes digna (Gahan, 1891)

Isotes eruptiva (Bechyné, 1955)

Isotes ignatia (Bechyné, 1956)

Isotes onira (Bechyné & Bechyné, 1961)

Isotes pollina (Bechyné & Bechyné, 1962)

Isotes puella (Baly, 1886)

Isotes sanguineipennis (Baly, 1891)

Isotes semiflava (Germar, 1824)

Isotes sibylla (Bechyné & Bechyné, 1969)

Isotes taeniolata (Gahan, 1891)

Isotes ternata (Bechyné & Bechyné, 1961)

Isotes valentina (Bechyné, 1956)

Isotes varipes (Boheman, 1835)

Paranapiacaba amplexa (Erichson, 1847)

Paranapiacaba biseriata (Gahan, 1891)

Paranapiacaba costalimai (Marques, 1941)

Paranapiacaba decemverrucata (Gahan, 1891)

Paranapiacaba diametralis (Bechyné, 1956)

Paranapiacaba melanospila (Gahan, 1891)

Paranapiacaba morretesi Bechyné & Bechyné, 1969

Paranapiacaba pereirai Bechyné, 1958

Paranapiacaba prolongata (Jacoby, 1882)

Paranapiacaba seraphina (Bechyné, 1956)

Paranapiacaba significata (Gahan, 1891)

Paranapiacaba subirregularis (Bechyné & Bechyné, 1962)

Paranapiacaba teinturieri (Allard, 1894)

Paratriarius batesi (Baly, 1859)

Paratriarius limbatipennis (Baly, 1889)

Platybrotica misionensis Cabrera & Cabrera Walsh, 2004

Zischkaita serrana Moura, 2003

1 Contribution to the 8th International Symposium on the Chrysomelidae, held August 23, 2012, in Daegu, South Korea