A list of 1298 species and 172 genera of Chrysomelidae from the subfamily Galerucinae (sensu stricto) with the males having at least one form of secondary sexual characteristic (SSC) is presented. The number of species amounts to 24% of the total Galerucinae presently known from all over the world—a very significant amount. The SSCs comprise various types of modified structures found on all parts of the body—head, thorax, and abdomen. They are not variable but species specific. Illustrations from selected 87 species that include 84 images and 15 line drawings showing various types of SSC are provided. The amazing array of SSCs from the Galerucinae offers a large and taxonomically diverse set of data that are not comparable with other subfamilies in the Chrysomelidae and may be useful in phylogenetic analysis of the family.
1. Introduction
Galerucinae (sensu stricto) is the second largest subfamily within the Chrysomelidae (Coleoptera), represented by about 520 genera and between some 5000 species [1], 5500 species [2], and 6300 species [3] and its most diverse distribution concentrated mainly in the tropical and subtropical regions.
The classification of the Chrysomelidae proposed by Lawrence and Newton [4] lumped together the Galerucinae and Alticinae into a single subfamily Galerucinae (sensu lato). Consequently, the Galerucinae (sensu stricto) was placed in the Tribe Galerucini and the Alticinae (sensu stricto) as the Tribe Alticini. Since then, some workers have continued to separate the two subfamilies as in the Seeno and Wilcox [5]. There has been support for Alticinae as a monophyletic group with metafemoral spring as the main character [6]. Likewise, Galerucinae has been considered paraphyletic with Alticinae as a subordinated clade [7]. On the other hand, Kim et al. [8] in their molecular and morphological-based phylogenetic studies showed that there is molecular data to support Alticinae as a tribal ranking (Alticini) within the Galerucinae and neither as a separate subfamily. They suggested that more independent characters are needed. More recently, Gillespie et al. [1] claimed to have the most comprehensive phylogeny estimation and showed consistency with previous molecular phylogenetic reconstruction of Galerucinae, but at the same time agreed that there was a lack of taxon representation from the Old World. There is no doubt that more characteristics need to be utilised from a more complete range of taxa selected from the Old and New Worlds, that is, better taxon sampling, including secondary sexual characteristics.
In this study, we refer to a secondary sexual characteristic (SSC), defined by Mayr [9] as a modified structure of the male members of a species that distinguishes the two sexes of the species but does not function directly in reproduction. The modified structure from the male differs from homologous structures in the female. The SSC may provide an advantage during competition for mates, for example, the aquatic smooth newt, Triturus cristatus [Laurenti] (Amphibia: Salamandridae), is dimorphic with the male having a dorsal crest. Darwin thought that the function of the crest was to attract females, but it was a century later that it was demonstrated that females chose males exhibiting conspicuous dorsal crests [10]. Observational and experimental evidence suggest that feather ornaments of birds play an important role in female choice [11]. Eberhard [12, 13] referred to an SSC as the nongenitalic male contact structure, a product of an intersexual competition [14]. Besides agreeing that SSCs are the result of Darwinian selection, Jolivet [15] noted that they are also a manifestation of biodiversity.
The SSCs are species specific; thus, they are important as diagnostic tools for identification. In the Chrysomelidae, particularly the subfamily Galerucinae, SSCs are found on various parts of the body, including head, thorax, and abdomen. The presence of these modified characteristics was discussed in some earlier works on the Galerucinae by Maulik [16, 17], who provided illustrations of modified antennae present in some members of the genus Agetocera Hope and modified clypeus in Palpoxena Baly. According to Maulik [17], the antennae, clypeus, labrum, and maxillary palpi that have undergone extraordinary modifications possess a highly diagnostic value. Blake [18, 19] noted the modified characteristics, the excised metatibiae of the male, as important generic characters (Deinocladus Blake, Luperosoma Jacoby, and Porechontes Blake). Wilcox [20] noted that in some genera, for example, Malacorhinus Jacoby, species in which the elytra are modified in the male are easily identified by that characteristic. Silfverberg [21] defined Prosmidia Weise and Neolaetana Laboissiere above all by their secondary sexual characteristics, the pronotum, and the basal area of the elytra being modified in the male.
In constructing a phylogeny or a classification, a set of characters from morphological data are tabulated, representing common ancestor (plesiomorphy) and derived states (synapomorphy). The SSC is a derived characteristic. Phylogeny is a statement not only of relationships among taxa but also about the evolution of characters. There is little literature pertaining to the utilization of the SSCs in phylogenetic studies of Chrysomelidae; thus, these characteristics received scant attention. The earliest traceable work was that of Silfverberg [22] in a revision of the galerucine group Prosomidiites, only known from the Afrotropical Region. He employed the modified scutellar area of the elytra as the root for eight genera. Furthermore, according to Silfverberg, the modifications of the male pronotum and elytra are of a type not present in other Galerucinae and can, therefore, be considered apomorphic when found in the Prosomidiites. After more than thirty years, SSCs began to be considered in a phylogenetic analysis. Freund and Wagner [23] constructed a phylogenetic tree of 31 species, which employed 4 SSCs. Barroga and Mohamedsaid [24] in the construction of a phylogeny for genus Aulacophora Chevrolat from Sundaland used 59 characteristics, including 13 SSCs. Recently, Stapel et al. [2] utilized the SSCs from the modified head, antennae, and elytra in their phylogenetic analysis on the Afrotropical Monolepta Chevrolat and related galerucine genera.
The present paper has three objectives: (1) to compile all known of SSCs found in chrysomelid beetles of the subfamily Galerucinae (sensu stricto) from all over the world, (2) to provide some explanation on functions of the various modified structures, and (3) to highlight the myriad forms of SSCs in the Galerucinae and their taxonomic significance in the Chrysomelidae.
2. Materials and Methods
Information sources for SSCs from the subfamily Galerucinae (sensu stricto) were gathered in three ways. Unless otherwise stated, all reference to Galerucinae in this study means sensu stricto. First, male specimens were examined physically for modified structures; second, data was extracted from descriptions and illustrations of modified structure in the literature; third, data was extracted through virtual examination of modified structure displayed on images of the males posted on websites of the Internet. Some of the figures included here were taken directly from websites and, therefore, their clarity and resolution may not be of high quality; however, we feel that they are adequate to illustrate the modifications discussed. For the first phase of this study, the first author examined all the specimens in collection of the Centre for Insect Systematic, Universiti Kebangsaan Malaysia (UKM). Subsequently, three main websites were consulted in the Internet: GBIF Deutschland: http://www.biologie.uni-ulm.de/; MCZ, Harvard University: http://mcz-28168.oeb.harvard.edu/mcz/index.htm; Smithsonian Institution Library for the digital Biologia Centrali-Americana: http://www.sil.si.edu/DigitalCollections/bca/explore.cfm.
In this study, we decided to exclude modified last visible abdominal sternite (the apical sternite) and modified first segment of protarsi. The SSCs resulting from modifications of these characteristics are universal that are present in most males of the Chrysomelidae although the modified apical sternite is much more complex in Galerucinae (sensu stricto) than in other subfamilies.
3. Results
In the course of the present study, the generic names for a number of species are retained because they have not been revised or the revisions are only partially completed. The genus Platyxantha Baly was reduced by Reid [25] to a synonym of Taumacera Thunberg, but this synonymy applies only to species from the Oriental Region. Platyxantha from the African Region has not been revised, and in the present study, 28 species are listed with modified characteristics that possibly belong to another genus than Taumacera. The same is true with Monolepta Chevrolat, where its African fauna has been revised by Wagner [26], but the Oriental and Australasian faunas have not been revised, and we believe some of them could belong to another genus. Therefore, the name Monolepta is retained for 41 species listed here with modified characteristics despite the fact that true Monolepta as redefined by Wagner [27] has no modified characteristics/SSCs. Another genus is Nymphius Weise recently revised by Bezděk [28], but for convenience, the following two species: Calomicrus (Nymphius) buettikeri (Medvedev) and C. (Nymphius) friedmani (Lopatin) with a modified abdomen are retained under Nymphius. This is because the abdomen of these two species has appendages, not a characteristic of the genus Calomicrus Stephens. According to Bezděk, the two species should be placed into a separate new genus. Nevertheless, all these taxonomic changes that would have happened do not affect the total number of modified species from the subfamily Galerucinae and, of course, there would be changes in the total number of genera.
As a result of this study, a list of 172 genera (Appendix A) and 1298 species (Appendix B) of the Galerucinae having SSCs is provided. These SSCs represent various types of modified structures found on all parts of the body head, thorax, and abdomen. Table 1 shows a distribution of various forms of modified structures of the SSCs found on the head, the thorax, and the abdomen. Table 2 shows distribution of the galerucine genera and species with modified head, thorax, and abdomens. Table 3 shows distribution of the galerucine genera and species with various types of modified antennae. Table 4 shows a list of 49 largest genera attributed to the main modified characteristics. Illustrations of 87 selected species, that include images of 84 species and line drawings of 15 species, are provided to show amazing and truly unusual SSCs of the Galerucinae. These SSCs are classified, tabulated, and provided with plausible explanations of their functions. However, it is beyond the scope of the present study to go into a detailed anatomy of the various modified structures.
Distribution of modified structures found on head, thorax and abdomen of the chrysomelid beetles subfamily Galerucinae (sensu stricto).
Head
Antennae with one, double, and triple segments, more than three segments modified, or covered with unusually long hairs. Vertex depressed or excavated, with surface smooth or with structures. Frontal tubercles ase very large. Clypeus is strongly convex, excavated, with surface smooth, or with structures. Labrum are extremely large or deformed. Maxillary palpi with the 3rd segment swollen. Eyes are large.
Thorax
Pronotum
Disc bulged and excavated with structures. Posterior margin with process.
Elytra
Elytra humpback. A pair of tubercles at base or at apex. A pair of cavities behind scutellum, in middle, at sides, or at apex. Humeri with rows of erect hairs.
Legs
Front legs: Femora emarginate. Tibiae thickened, curved, emarginated, excised, or notched. Middle legs: Tibiae curved, excised, or notched. Hind legs: Trochanters with a process, a spine, or blade-like structure. Femora enlarged and excavated. Tibiae broadened, excavated, or curved; apex of tibiae with a process. All tarsal claws bifid (simple or appendiculate in female).
Metasternum
Metasternum with posterior median lobe or covered with long hairs.
Abdomen
First abdominal sternite with an appendage, a tubercle, or a pair of appendages, or a projected covering the following four sternites; first and second sternites each with a pair of appendages; second visible sternite with an appendage; third sternite with one or a pair of long appendages; fourth sternite emarginated, with a triangular process, a pair of spines, or long appendages.
Distribution of galerucine genera and species with modified head, thorax, and abdomen.
List of 49 largest genera with common modified organs.
Genera
No. of species
Common modified organs
(1) Apophylia
148
Tarsal claws bifid
(2) Diacantha
81
Elytra tuberculate
(3) Megalognatha
59
Antennal segments modified
(4) Aulacophora
54
Antennal segments modified
(5) Taumacera
45
Metasternum with posterior process
(6) Paleosepharia
44
Elytra excavated
(7) Palpoxena
41
Head excavated
(8) Cerophysa
33
Antennal segments modified
(9) Monolepta
31
Elytra excavated
(10) Sermyloides
31
Head excavated
(11) Pseudocophora
25
Elytra excavated
(12) Hoplosaenidea
24
Head excavated
(13) Hyphaenia
24
Antennae with long hairs
(14) Xenoda
23
Antennal segments modified
(15) Agetocera
22
Antennal segments modified
(16) Diabrotica
21
Antennal segments modified
(17) Asbecesta
20
Antennal segments modified
(18) Prosmidia
18
Elytra tuberculate
(19) Platyxantha
17
Antennal segments modified
(20) Paridea
15
Elytra excavated
(21) Hoplasoma
13
Abdomen with appendages
(22) Diabrotica
13
Eyes large
(23) Eccoptopsis
12
Antennal segments modified
(24) Afrocandezea
11
Elytra excavated
(25) Haplosomoides
11
Abdomen with appendages
(26) Lamprocopa
10
Head excavated
(27) Laetiacantha
10
Elytra tuberculate
(28) Afrocrania
10
Elytra excavated
(29) Scelolyperus
10
Hind tibiae curved
(30) Macrima
9
Head excavated
(31) Nymphius
9
Abdomen with appendages
(32) Luperosoma
9
Middle tibiae notched
(33) Trichobrotica
9
Middle tibiae notched
(34) Malacorhinus
8
Elytra excavated
(35) Candezea
8
Elytra excavated
(36) Chapuisia
8
Eyes large
(37) Kanahiiphaga
7
Head excavated
(38) Phyllobroticella
7
Antennal segments modified
(39) Samoria
7
Antennal segments modified
(40) Fleutiauxia
6
Head excavated
(41) Parabecesta
6
Antennal segments modified
(42) Metacoryna
5
Antennal segments modified
(43) Schematiza
5
Antennal segments modified
(44) Theopea
5
Antennal segments modified
(45) Cneoranidea
5
Maxillary palpi large
(46) Atrachya
5
Elytra excavated
(47) Neolaetana
5
Elytra excavated
(48) Euliroetis
5
Abdomen with appendages
(49) Jacobya
5
Pronotum excavated
Every part of the body, including the head, thorax, and abdomen, is affected with some forms of modifications (Tables 1, 2, and 3). Table 1 shows various modified structures present on the head, thorax, and abdomen. On the head, almost every aspect has a structure that has undergone modification, including vertex, frontal tubercles, clypeus, labrum, maxillary palpi, eye, and antenna. Table 2 shows distribution of galerucine genera and species with the modified structures. Modifications of head structures are found in 756 species and 119 genera. These modified structures can be described as simple to complex. The simple modified vertex has a smoothly bulged surface, or smoothly, strongly depressed surface (Monolepta flavicollis (Gyllenhal) (Figure 1)). The complex modified vertex has a depressed or excavated area with ridges or other structures (Aulacophora frontalis Baly (Figure 2)).
Modified head: Monolepta flavicollis (Gyllenhal).
Modified head: Aulacophora frontalis Baly.
The clypeus is located anterior to the frontal tubercles. The modified clypeus is found in 181 species and 24 genera (Table 2). The genus Palpoxena Baly has the highest number of modified clypeus, represented by 43 species. A simple modified clypeus has its surface transversely, deeply depressed and without structures (Cerotoma atrofasciata Jacoby (Figure 3)). A complex one has some forms of structures, such as ridges, projections, and hairs. In Aulacophora cornuta Baly (Figures 4(a) and 4(b)), the clypeus is deeply excavated anteriorly and with a process at sides. In an extreme case, the modified clypeus is deformed or disfigured (Palpoxena divisa (Jacoby) (Figure 5), P. sumatrensis (Jacoby) (Figure 6), and Eccoptopsis denticornis (Jacoby) (Figures 7(a) and 7(b))). Mouthparts affected with modifications are labrum and maxillary palpi. Modified labrum in form of extremely large size is found in two genera, Acroxena Baly and Palpoxena Baly. The labrum is either transverse (P. barbata (Baly) (Figures 8(a) and 8(b))) or triangularly shaped (P. coeruleipennis (Baly) (Figure 9) and P. variabilis (Jacoby) (Figure 10)). Maxillary palpi with modification are quite common, and they are found in 51 species and 12 genera. In some species the modification is very extreme and the usually cylindrical-shaped of the palpi is deformed (Palpoxena laeta (Baly) (Figure 11)). Males with large size of eyes are found in 35 species and 12 genera (Table 2).
Modified head: Cerotoma atrofasciata Jacoby.
Modified head: Aulacophora cornuta Baly.
Modified head: Palpoxena divisa (Jacoby).
Modified head: Palpoxena sumatrensis (Jacoby).
Modified head: Eccoptopsis denticornis (Jacoby).
Modified labrum: Palpoxena barbata (Baly).
Modified labrum: Palpoxena coeruleipennis (Baly).
Modified labrum: Palpoxena variabilis (Jacoby).
Modified maxillary palpi of Palpoxena laeta (Baly).
The modified antennae are the most widely distributed, found in 645 species and 119 genera (Table 2). The number of species with modified antenna amounts to 50% of the total modified Galerucinae. The genus Megalognatha Baly has the highest number with modified antenna, represented by 59 species. Generally, all males and females have 11 antennal segments. However, in the following two genera, Oroetes Jacoby and Phyllecthris Dejean, the male has the antenna with 10 segments compared with 11 in females. Every antennal segment, from the first to the eleventh, is affected by some form of modification. A simple modification is an enlargement of a segment, without any cavity, structure, or gland. A complex one affects more than one segment. Besides, some of the modified segments appear totally different from normal feature, being distorted or deformed.
Table 3 shows distribution of galerucine genera and species with various types of modified antenna. There are examples of sets of single, double, triple, and more than three modified segments. Antennal segment 2 is rarely modified but always reduced to a minute size if segment 3 is greatly modified, as in Taumacera spp. However, in Aulacophora diversa Baly and A. dulitensis Barroga and Mohamedsaid, segment 2 is enlarged and dilated at apex and modified together with segments 3–7. The same is with segment 6 that is never singularly modified but always combined with other segments, such as in the double modified segments 5 and 6 (Taumaceroides sinicus Lopatin) and segments 6 and 7 (Taumacera variceps (Laboissiere); and in the triple modified segments 5, 6, and 7 (Paratriarius dorsatus (Say), and segments 6, 7, and 8 (Megalognatha roletti Laboissiere (Figure 34)).
There are 156 species in 41 genera having a single modified antennal segment. Segment 3 is the most affected and found in 40 species and 9 genera. The genus Sermyloides Jacoby has the highest number with the 3rd segment modified, represented by 26 species. The following are some examples of species with the single modified antenna that affect segment 1 (A. cornuta (Figure 4) and Sarawakiola ajaib Mohamedsaid (Figure 12)), segment 3 (Taumacera sucki (Weise) (Figure 13)), segment 4 (Afrocrania kaethae Middelhauve and Wagner (Figure 14) and A. latifrons (Weise)), segment 5 (Apophylia incisitarsis (Laboissiere) (Figure 15)), segment 7 (Duvivieria apicitarsis Weise (Figure 16) and Megalognatha grouvellei (Weise) (Figure 17)), segment 8 (Cerophysa flava Baly (Figure 18) and Cerophysa gestroi Jacoby (Figure 19)), segment 9 (Metacoryna guatemalensis Jacoby (Figure 20) and Agetocera silva (Bezděk) (Figure 21)), segment 10 (Ectmesopus darlingtoni Blake, (Figure 22)) and segment 11 (Aulacophora luteicornis (Fabricius) (Figure 23)).
Modified antennae: Sarawakiola ajaib Mohamedsaid.
Modified antennae: Taumacera sucki (Weise).
Modified antennae: Afrocrania kaethae Middelhauve and Wagner.
There are 106 species in 27 genera having a set of double modified antennal segments with segments 3 and 4 the most affected, which is found in 36 species and 12 genera. But the genus Agetocera Hope with the double modifications of segments 8 and 9 has the highest number, represented by 19 species. Examples of species with double modifications of segments 1 and 3 (Taumacera khalednordini Mohamedsaid (Figure 24)), segments 3 and 4 (Hoplosaenidea aerosa (Laboissiere) (Figure 25) and Cerotoma ruficornis (Olivier) (Figure 26)), segments 4 and 5 (Afrocrania foveolata (Karsch)), segments 5 and 6 (Taumaceroides sinicus), segments 6 and 7 (Cerophysa sumatrensis (Jacoby) and Taumacera variceps); segments 7 and 8 (Kinabalua musaamani Mohamedsaid (Figures 27(a) and 27(b))), segments 8 and 9 (Taumacera occipitalis (Laboissiere) (Figure 28), Metacoryna fulvipes Jacoby (Figure 29) and M. jacobyi Bowditch (Figure 30)); segments 9 and 10 (Agetocera nigripennis Laboissiere (Figure 31)). Apparently in Metacoryna jacobyi the modified segments are fused together and appear as one.
There are 97 species in 21 genera having a set of triple modified antennal segments with a combination of segments 3, 4, and 5 the most commonly affected and found in 36 species and 3 genera. The genus Aulacophora has the highest number of triple modifications, represented by 34 species. Examples of triple modifications are segments 3, 4 and 5 (Aulacophora laevifrons Baly (Figure 32) and A. martia Weise (Figure 33)), segments 5, 6, and 7 (Paratriarius dorsatus (Say)), segments 6, 7, and 8 (Megalognatha rolleti Laboissiere (Figure 34)), segments 7, 8, and 9 (Paratriarius neocrassicornis (Bowditch)) and segments 9, 10, and 11 (Vitruvia clavicornis Weise).
The modified antenna with more than three segments is the most common and widely distributed, found in 232 species and 71 genera. The genus Xenoda Baly has the highest number with more than three segments modified and represented by 23 species. The affected segments are either a combination of three or more basal, apical, or a combination of almost all the antennal segments. Examples are (Malacorhinus antennatus Jacoby (Figure 35), Megalognatha variicornis (Weise) (Figure 36), Exosoma deformicornis (Queddenfledt) (Figure 37) and Megalognatha femoralis Laboissiere (Figure 38)). However, in the genus Xenoda (subgenus Xenoda), all modified segments 3 to 8 are united into more or less one piece (Xenoda pallida Jacoby (Figure 39) and X. ovalis Mohamedsaid (Figure 40)). This union of segments is also found in other species, but it affects two segments, the 8th and 9th, as in Metacoryna jacobyi (Figure 30). Besides affecting the shape of antennal segments, the modified antenna is also the one with the presence of long hairs. There are 55 species in 19 genera having antenna covered with long hairs.
In thorax, these modified structures are found on pronotum, scutellum, elytra, legs, and the ventrite of the third thoracical segment (Table 2). The modified pronotum is the least represented, found in 38 species and 10 genera. The pronotum has either the disc bulged or excavated (Jacobya cavicollis (Fairmaire) (Figure 41)), or the middle of posterior margin bearing a short process or knob (Cannonia meridionalis (Weise) (Figure 42)). In an isolated case, scutellum for three species of Prosmidia is transformed into tongue-liked shape (P. suahelorum Weise (Figure 43)). In addition, these species also have both pronotum excavated and elytra with a pair of tubercles.
Modified pronotum: Jacobya cavicollis (Faimaire).
Modified pronotum: Cannonia meridionalis (Weise).
Modified pronotum: Prosmidia suahelorum Weise.
Modified elytra are the most widely distributed and found in 304 species and 30 genera (Table 2). The number of species with modified elytra amounts to 24% of the total modified Galerucinae. There are two types of modified elytra, one is with the presence of tubercles and the other with cavities. The modified structures are located either at base, posterior to scutellum or between scutellum and humeri laterally, or at the apex of the elytra. In Elyces quadrimaculata Jacoby the humerus is strongly tuberculate. A pair of tubercles is located either at the base of elytra (Laetiacantha distincta (Gahan) (Figure 44) and Laetiacantha ruficollis Laboissiere (Figure 45)), or at the apex of elytra, found in 81 species of the genus Diacantha Chevrolat. The tubercles have various sizes and shapes, such as rounded (Diacantha senegalensis (Laboissiere) (Figure 46)), pointed (D. insignipennis Thomson (Figure 47) and D. varians Weise (Figure 48)), bifurcate (D. cupripennis (Laboissiere) (Figure 49)), semicircular-shaped (D. bituberculata (Fabricius) (Figure 50)), and elongate and parallel to the suture (D. dimidiata (Laboissiere) (Figure 51)). In an extreme case the tubercle forms a long lateral carina, extending from the humeri to apex (Candezea bicostata (Weise) (Figure 52)).
Opposite to the formation of a tubercle, the elytra are excavated. There are various sizes and shapes of the cavities, as well as their depth. The cavities are located on various parts of the disc, posterior to the scutellum, in the middle, at sides, or at apex of the elytra. The most common are located at posterior of the scutellum. Simple excavated elytra have traces of a cavity, or with shallow depression (Afrocrania ubatubae Middelhauve and Wagner (Figure 53)), with a straight groove (Paleosepharia jambuica Mohamedsaid (Figure 54)), an outward oblique at the posterior (P. zakrii Mohamedsaid (Figure 55), P. lawa Mohamedsaid (Figure 56) and P. legenda Mohamedsaid (Figure 57)), or a spindle-shaped cavity (P. tenasserimensis (Maulik) (Figure 58)). In Monolepta kerangas (Mohamedsaid) (Figure 59), the cavity is located in the middle and the elytra humpbacked view from side. A few species have the cavity located at apex of elytra (Paratriarius alternans (Weise); Strobiderus excavatus Jacoby). In other species, the cavity is funnel shaped and located at the base between scutellum and humerus (Neolaetana alternans (Silfverberg)). Pseudocrania basalis Jacoby (Figure 60) has the broadest excavated elytra. In addition, in the following two genera: Malacorhinus Jacoby and Androlyperus Crotch, the cavities are located at the sides. In the former, the cavities are located in the middle and the latter near the apex of the elytra. Examples are M. decempunctatus Jacoby (Figures 61(a) and 61(b)), M. foveipennis Jacoby (Figure 62), M. tripunctatus Jacoby (Figure 63) and A. nataliae Clark (Figure 64). Another type of modified elytra is with the presence of rows of erect hairs on shoulders, which is found in five species of Aulacophora.
Modified elytra: Afrocrania ubatubae Middelhauve and Wagner.
The modified leg includes modifications on any part of it, which include trochanter, femur, tibia, tarsus, and claws. There are 401 species in 88 genera with modified legs (Table 2). The number of species with modified leg amounts to 31% of the total modified Galerucinae. The affected part of the leg is either enlarged, emarginated, excised, with a process, or spine, or having a different set of tarsal claws. The male has bifid tarsal claws in contrast to the female with either appendiculate or simple claws. The genus Apophylia Thomson, represented by 148 species, has a characteristic of bifid tarsal claws differing from females that have appendiculate claws. Another genus with the male having bifid tarsal claws is Erynephala Blake, represented by 5 species.
The modified tibia is the widely distributed modification of the leg, represented by 103 species and 39 genera. Examples of part of the modified leg are metatrochanter with a long spine in Coeligetes borneensis Mohamedsaid (Figure 65); protibia enlarged in Mombasa magna (Weise) (Figure 66) and Cerotoma dilatipes Jacoby (Figure 67); protibia emarginate in Taumacera tibialis Mohamedsaid (Figure 68); mesotibia emarginated in Dreeus distinctus (Shute) and Taumacera midtibialis Mohamedsaid (Figure 69); metatibia curved and with a tooth in the middle in Scelolyperus tejonicus Crotch (Figure 70). In the two rare species, Sastroides tarsalis Mohamedsaid and Momaea distincta Mohamedsaid (Figures 71(a) and 71(b)), two parts of the same leg are modified, the mesotibia and mesotarsus, with the former excavated and covered with a series of pegs and the latter has a long process. A modified first segment of protarsus is common in the Chrysomelidae, but in some Galerucinae, it is strongly dilated as in Taumacera yamamotoi (Mohamedsaid) (Figure 72).
Modified legs: Coeligetes borneensis Mohamedsaid.
Modified legs: Mombasa magna (Weise).
Modified legs: Cerotoma dilatipes Jacoby.
Modified legs: Taumacera tibialis Mohamedsaid.
Modified legs: Taumacera midtibialis Mohamedsaid.
Modified legs: Scelolyperus tejonicus Crotch.
Modified legs: Momaea distincta Mohamedsaid.
Modified legs: Taumacera yamamotoi (Mohamedsaid).
The thorax also has the modified structures on the venter of its third segment, known as the metasternum, and this is found in 60 species and 4 genera. The modified metasternum is present in tubercle (Apophylia nobilitata Gerstaecker (Figure 73)), an elongated lobe (Kinabalua musaamani Mohamedsaid (Figure 74)) or a laminated lobe (Taumacera midtibialis (Figure 75)). Besides, in Cneoranidea hirta (Yang), the metasternum is covered with long hairs.
Modified ventrite of thorax: Apophylia nobilitata Gerstaecker.
Modified ventrite of thorax: Kinabalua musaamani Mohamedsaid.
Modified ventrite of thorax: Taumacera midtibialis Mohamedsaid.
The abdomen has the sternite with modified structures in the present appendages, tubercles, and hairs. There are 66 species in 18 genera with modified abdomen (Table 2). The number of species with modified abdomen amounts to 5% of the total modified Galerucinae. The genus Hoplasoma Jacoby has the highest number with modified abdomen, represented by 13 species. The modified abdominal sternite are either densely covered with long hairs (Hemygascelis longicollis Jacoby (Figure 76)), or bearing one piece (Haplosomoides chinmatra Maulik (Figure 77)), a pair (Coeligetes submetallica Jacoby (Figure 78) and Hoplasoma unicolor (Illiger) (Figure 79)), or two pairs of appendages (H. apicale Jacoby (Figure 80), Androlyperus fulvus Crotch (Figure 81) and Scelida flaviceps (Horn) (Figure 82)). Also a complex abdominal appendage is found in species of Nymphius Weise (N. ensifer (Guillebeau) (Figure 83), N. forcipifer (Weise) (Figure 84), N. gianassoi Bezděk (Figure 85), N. lydius (Weise) (Figure 86), N. ogloblini (Bogachev) (Figure 87), N. pravei (Jacobson) (Figure 88), and N. stylifer (Weise) (Figure 89)).
Table 4 presents a list of 49 largest genera represented by at least 5 species. These genera represent 1007 species or 74% of the total 1298 species presently listed. The genus Apophylia has the highest number of species (148) with modified structures followed by Diacantha (81), Megalognatha (59), Aulacophora (54), and Taumacera (45). However, the most common form of SCC is the modified antennae, followed by the modified elytra and the modified head.
Worldwide distribution of modified Galerucinae both from the Old and New Worlds has the former represented by 88% of the total modified species of the subfamily. The Luperini is the major tribe and represented by 84% of the total modified species in the Galerucinae. The Old World Luperini has the highest representation of SCCs with 87% of the total modified Luperini. The Galerucini from the Old World has the highest representation with 90% of the total modified from the tribe.
4. Discussion
A total of 1298 species and 172 genera of the Galerucinae (sensu stricto) with SSCs are recorded; a huge number. The modified species represent about 24% of the total number of the Galerucinae, based on an estimate of 5500 species [2]. However, we believe that there are many more species with SSCs that either we were not aware of or were not available for our examination. Our observations were mainly based on original descriptions and illustrations and images of types that are posted on websites of various institutions. However, there are many species that were described based on female specimens and/or are the only images posted on websites. There is a high possibility that males belonging to a species that was described based on a female have modified characteristics, but there are no images or subsequent descriptions of males of these species. In certain species, we know that some congeneric species described based on the female have males with modified characteristics. We also know that not all males of congeneric species have SSCs; therefore, we cannot make an assumption that a particular species described based on female has male with modified characteristics without examining the specimen, and such a species is discounted in the present study. Despite these deficiencies, we believe that our compilation of species of the Galerucinae with SSCs is representative for the subfamily. We believe that this study has covered more than 80% of all known species of the Galerucinae from around the world and recorded well the presence of the secondary sexual characteristics.
Among the three body parts (head, thorax, and abdomen), the head with various modified structures (vertex, frontal tubercles, clypeus, labrum, maxillary palpi, and antennae) has the widest representation in the Galerucinae, found in 756 species and 119 genera. The thorax with modified pronotum, elytra, third thoracic sternite, and legs is found in 379 species under 37genera. In the modified thorax, the elytra have the highest representation with 304 species, followed by modified first foretarsal segments with 158 species. The ventrites with the modified third thoracic sternite and abdominal sternites combined are found in 126 species of 20 genera. In the modified head, the antennae have the highest representation with 645 species, followed by modified clypeus with 181 species. Modifications of the head occur in 756 species or 58% of the total modified Galerucinae, and of these 645 species have modified antennae (50% of the total) also higher than modifications found in other parts of the body, such as thorax and abdomen.
Generally, the distribution of modified structures on a body part is associated with their function. Modified structures on the head may be associated with olfaction and to a lesser extent with some modified pronota and elytra having hairs or sensillae. Modified structures on the thorax, especially on the legs, are associated with precopulatory behaviour. Also the modified abdomen is associated with precopulatory behaviour. Details of these modified structures are discussed below. There are certainly many more examples of SSCs of which we are not yet aware but will be discovered in the future. The record of Galerucinae having the SSC is growing. Since 2005, 78 species were described including 9 that were described in 2010 and recently 2 in 2011 (see Appendix B, list of species).
5. Peculiar Form of SSCs
Interestingly, the antennae and tarsal claws are two characters that have some of the most peculiar forms of SSCs in the Galerucinae. Usually the number of antennal segments is the same in both males and females, that is, 11 segments in almost all the Galerucinae, and fewer than that in a few genera of the subfamily Alticinae (sensu stricto), for example, 9 segments in Nonarthra Baly, and 10 segments in Psylliodes Latreille Monotalla Bechyné, and Deciplatus Linzmeier and Konstantinov. However, in a couple genera of Galerucinae, there is a difference in the antennal segments between male and female. In Oroetes Jacoby and Phyllectris Dejean the males have 10 antennal segments (O. flavicollis, P. gentilis (Leconte)) compared with females having 11 segments. There is no apparent explanation as to why the male has less segments than the female.
It is common for both males and females to possess the same characteristics of the tarsal claws, either simple, bifid, or appendiculate. However, in Apophylia, the males have bifid tarsal claws versus appendiculate claws in the females, and in Erynephala Blake, males also have bifid claws, but females have simple claws. The beetles of the genus Apophylia have the elytral surfaces closely punctuate, appearing rugose. The same is true for the elytral surface of Erynephala that have closely but moderately coarse punctures. Thus, it is speculated that the male uses its bifid tarsal claws to hold onto the rugose surface of the female elytra during copulation. Of course, there are many genera with both males and females with the same tarsal claws, either bifid or appendiculate and with the rugose elytra.
6. Functions of SSCs
More than 1000 species have SSCs with more than two modified structures, and some 100 species have three or more modified structures. The presence of more than one type of SSC is also known as having multiple sexual ornaments. There are several hypotheses attempting to explain why a male would have multiple sexual ornaments. One of these is the multiple messages hypothesis, which states that each trait display reflects only one aspect of the overall quality of the male, different ornaments, and therefore signal different kinds of properties in an individual [29]. Additional displays may serve to enhance the accuracy with which observers assess a single quality or serve to provide information about different qualities [30]. According to Moller and Pomiankowski [29], multiple sexual ornaments should be particularly common in taxa with the most intense sexual selection (i.e., lekking and other polygynous taxa). However, the SSCs are apparently not ornaments displayed by males to attract females. There is no variation in the SSCs among individuals within a species that would lead to intraspecific competition. Also, they are not weapons like the horns that are found in the other beetles, such as the Scarabaeidae or the jaws of Lucanidae. In these two families, the horns and jaws commonly function as weapons in the intraspecific battles [31].
The majority of the SSCs are associated with sensory and olfactory functions particularly in the antennae and to some extent in elytra where glands are formed. The other modified structures of the SSCs are associated with copulatory behavior, and in some it is suspected to be involved in sound production.
6.1. Olfaction
Olfaction begins with transduction of the information carried by odor molecules into electrical signals in sensory neurons. The odor causes species-specific signal responses by the olfactory receptor neurons. Insect odor and taste receptors are highly sensitive detectors of food, mates, and oviposition sites, and these receptors have been identified in several insects [32]. The sense organs act as transducers in converting chemical energy into electrical energy of nerve impulses in sensory neurons to stimulate appropriate behavioural responses, such as finding food and mate and avoiding danger.
As an olfactory organ, antennae are all provided with sensory receptors or sensillae. Usually, in a modified segment with gland, the segment is enlarged, oval, or rounded with an opening or pore, as in Cerophysa flava (Figure 18), C. gestroi (Figure 19), Metacoryna guatemalensis (Figure 20), M. fulvipes (Figure 29), Ectmesopus darlingtoni (Figure 22), and Aulacophora luteicornis (Figure 23). The same is true with some modified elytra with cavities found in Malacorhinus spp. (Figures 61–63) and Androlyperus nataliae (Figure 64) that bear sensillae and glands [33]. The sensory receptors are densely located in the excavated antenna of Sarawakia ajaib (Figure 12) and Cerotoma ruficornis (Figure 26), clypeus of Palpoxena spp. (Figures 5, 6, 8, and 9), and Eccoptopsis denticornis (Figure 7).
6.2. Copulatory Behavior
The SSC associated with copulatory behaviour is known as a nongenitalic male contact structure [12]. One common function of this structure is to grasp the female. The nongenitalic male structure, such as modified antennae and legs that are specialized to grasp and stimulate the female, is involved in intersexual competition [13]; thus, the tight grasping by the copulating male resists any attempt by another male to dislodge the copulation.
There is a hypothesis about the interrelation between the genitalia and the secondary sexual characteristics (SSCs), wherein for a small and poorly sclerotized aedeagus, there is a provision for compensation in external structures [12, 13]. For example, recently, Konstantinov and Korotyaev [34] suggested that the male apionid beetle (Trichoconapion hirticone (Korotyaev)) probably used the three thickened apical antennal segments as a clasping organ during copulation since it has smaller aedeagus. In the genus Aulacophora, there is a relationship between the structure of aedeagus and the presence and absence of the nongenitalic characteristics [35]. A male species with a small and poorly sclerotized aedeagus is speculated to use the enlarged modified 3rd, 4th, and 5th antennal segments (Figures 32 and 33) function to grasp the female during copulation. It is speculated that other species in a number of genera with large modified middle antennal segments may possibly use these as a grasping organ (Malacorhinus antennatus (Jacoby) (Figure 35), Megalognatha femoralis (Laboissiere) (Figure 38)).
This speculation is that when the leg functions as a grasping organ there is a larger body size, or there is some form of modification of the leg such as being curved or excised (emarginate or notched). The modified structures are found on various parts of the front, middle, and hind legs. Usually, only one part of a leg is modified, either the femur, tibia, tarsus, or tarsal claws. For example, if the protibia is enlarged (Mombasa magna, Figure 66, Cerotoma dilatipes, Figure 67), the protibia excised (Taumacera tibialis, Figure 68), midtibia excised (T. midtibialis, Figure 69), and the metatibia curved with a spine (Scelolyperus tejonicus, Figure 70). The modified first segment of the protarsus being enlarged or strongly dilated is also known in males of other subfamilies of Chrysomelidae; however, in the Galerucinae, the modified tarsi are extremely large and with various shapes (e.g., Taumacera yamamotoi (Figure 71)). In the genus Apophylia, the presence of bifid tarsal claws, which are appendiculate in the female, possibly functions as grasping organs.
Another nongenitalic male contact structures are found on the ventrite of the third segment of thorax, the metasternum, and the five visible abdominal sternites. In the modified metasternum, there are the median of the posterior margin, an extended process, or a pair of lobes. The function of these SSCs could possibly be associated in precopulatory behaviour; the metasternal process (Figures 73 and 74) is used by the male to stroke the female. However, the appendages on the modified abdomen (Figures 75–88) are more complex, particularly in the genus Nymphius (Figures 82–88). It is also speculated that these appendages are used by the male to stroke the female during copulatory behaviour.
6.3. Sound Production
Sound produced through vibration and stridulation in animals, including insects, provides information used in predator-prey interactions, recruitment to food, mate choice, intrasexual competition, and maternal/brood social interactions [36]. Two examples from various studies on insect sound production are the termite Zootermopsis angusticollis (Hagen) that bangs its head to produce a substrate-borne vibration as an alarm signal of a disturbance to the nest by predators [37], and the leaf-cutter ant, Atta cephalotes (Linnaeus), that produces a stridulation sound using a file and scrapper mechanism for communication. Foraging workers stridulate while cutting a leaf fragment, and the stridulatory vibrations are used by the hitchhikers to locate workers engage in leaf cutting [38].
It is suspected that some of the SSCs in the Galerucinae are involved in producing sound, and that such devices are located in every part of the body, the antenna, pronotum, scutellum, elytron, leg, and abdomen [39]. Stridulation is produced by a file and scraper mechanism. Two species of Kinabalua Mohamedsaid and nine species of Xenoda Baly of the subgenus Xenoda (sensu stricto) have modified antennae with stridulatory devices. Figure 27 shows Kinabalua musaamani Mohamedsaid with antennal 8th segment bearing a long spine and 7th segment bearing a series of pegs on basal third. Xenoda species (Figures 39 and 40) has antennal segment 8 bearing a long spine and segment 9 bearing a series of pegs on the apical area. In the modified antennae mentioned above, the spine functions as a scraper and is pulled over the pegs stridulation is produced.
In some species of the following genera: Cannonia (Figure 42), Laetiacantha (Figures 44 and 45), and Prosmidia (Figure 43), the modified pronotum has a short process in the middle of the posterior margin. It is suspected that when the pronotal process (scrapper) is pulled over the surface of scutellum (file), stridulation occurs. This type of sound could also be produced by modified elytra of Prosmidia that has a pair of tubercles located at the base, between scutellum and humeri. It is speculated that during elytral movement, the basal tubercles come into contact with the posterior margin of pronotum and stridulation occurs.
Generally, as discussed above, the modified structures found on legs are associated with copulatory behaviour. But in some Galerucinae, some are suspected to be involved in sound production using a file and scrapper mechanism. These stridulation devices are found on the metatrochanter, mesotibia, and tarsi. In the following three species Coeligetes borneensis Mohamedsaid (Figure 65), Liroeties spinipes (Oglobin), and Monolepta trochanterina Mohamedsaid, the metatrochanter bears a long pointed appendage. This trochanter which moves vertically allows the spine to rub the abdominal sternite covered with series of pegs to produce sound. In males of Momaea distincta (Figures 71(a) and 71(b)) on the middle leg, the first segment of tarsus bears a long process pointing towards an excavated area at the apex of tibia that is covered with a series of pegs. Stridulation occurs when the tarsal process (scrapper) is pulled over the series pegs on the tibia.
There are cases of modified elytra with a pair of tubercles located at the apex that are suspected to produce vibrational sound. Vibration can provide a channel of communication between males and females during mating when no stridulation occurs. The tubercle is hollow, thus when tapped on the abdomen a type of vibrational sound may be produced. It is like when one cups one’s hand and taps on a surface, it produces a sound that is different compared with tapping an open hand. Amazingly, in the genus Diacantha alone, there are 81 species with males having modified elytra in a series of various-shaped tubercles. These tubercles may produce different vibrational sounds. Elytra with this type of tubercle are also found in some species of Paratriarius and one species of Sonchia.
The presence of sound production devices in some of the Galerucinae (sensu stricto) is very interesting and has great potential for future research. It had been reported in other subfamilies of the Chrysomelidae, such as the Megalopodinae, Zeugophorinae, Criocerinae, Clytrinae, Hispinae, and Cassidinae [40–42].
6.4. Unknown Function
For some SSCs, there is no apparent explanation as to their functions. The following are some examples: first antennal segment enlarged (Aulacophora cornuta (Figure 4) and Taumacera khalednordini (Figure 24)), vertex strongly broadly depressed (Monolepta flavicollis (Figure 1)), vertex longitudinally grooved (Aulacophora frontalis (Figure 2)), frons excavated with a projection (Lamprocopa spp.), clypeus transversely, deeply depressed (Cerotoma atrofasciata (Figure 3)), clypeus excavated with projections or spines (Palpoxena sumatrensis (Figure 6), Eccoptopsis denticornis (Figure 7), Palpoxena coerulipennis (Figure 9)), labrum extremely large (Palpoxena spp. (Figures 8–10)), maxillary palpi large (Palpoxena laeta (Figure 11)), larger eyes in the males of 35 species and 12 genera (Table 2), pronotum excavated with projection (Jacobya cavicollis (Figure 41)), scutellum excavated, with sides raised into high ridges (Halysacantha weisei (Jacoby)), elytra longitudinally carinate (Candezea bicostata (Figure 52)), and elytra with cavities or extrusions in Paleosepharia (Figures 54–58) and some species of Monolepta (Figure 59), Pseudocrania (Figure 60), Paridea spp., and Pseudocophora spp.
7. Utilization of the SSC7.1. Identification of the Sex
The presence of SSCs has long been utilized as tool for sex identification. According to Hinton [43], a rapid and accurate method of sexing bark beetle is valuable, but the absence of SSCs in the adult confused flour beetle, Tribolium confusum Duval, and the rust-red flour beetle, T. castaneum Herbst, means that the beetles could only be sexed in the pupal stage, a procedure which is frequently very inconvenient. Schmitz and Furniss [44] reported the relative accuracy and usefulness of SSCs as sex indicators in Scolytus laricis Blackman. Recently, French and Hammack [45] show that the presence of SSC in the tarsal segment of corn rootworms and bean leaf beetles facilitates sex identification, which is very useful for modelling and management purposes.
7.2. Subfamily Taxonomy
There are 1298 species and 172 genera of Galerucinae with at least one type of SSC. The number of species known to have SSCs represents about 24% of the total known Galerucinae, estimated as 5500 [2]. This is significant representation. A high number of Galerucinae with SSCs indicate a large amount of information or data that can be utilized to understand the subfamily in relation to other subfamilies of Chrysomelidae.
Taxonomically, of course, all modified structures are significant at the species level and to some extent at the generic level. Table 4 provides a summary of modified characteristics found in 49 largest genera. The modified characteristics tabulated in Table 4 are generally distinctive for the respective genera. Not much has been written about the significance of the modified characteristics at the subfamily level, except for the antennae. Jolivet [46] highlighted some Galerucinae with modified antennae and noted that compared to other Chrysomelidae this subfamily possesses the greatest diversity in antennal shape. Mohamedsaid [47] provided a general review on some modified antennae of Malaysian Galerucinae. The degree of antennal modification is much greater, not only in terms of their larger size, but also in terms of shape that to some extent may be extremely distorted or deformed. Much earlier, Maulik [17] concluded that such antennal modifications common in the Galerucinae are not found in any other subfamily, not even in the very closely allied Alticinae. However, the present study provides all types of SSCs in the form of many modified structures, including the antennae, found on all parts of the body, the head, thorax, and abdomen. A summation of all these modified characteristics documents its taxonomic significance with respect to the subfamily Galerucinae and continues to demonstrate additional differences of the Galerucinae (sensu stricto) from other subfamilies, especially the Alticinae.
There are some modified structures present in other subfamilies of Chrysomelidae, but they are not comparable in terms of representation within a subfamily and complexity of the modifications or the numbers of taxa found in the Galerucinae. This begs the question as to why this phenomenon of the SSCs is so prevalent and somewhat unique for these characters of the Galerucinae and that this may reflect its position in the Chrysomelidae. This SSC phenomenon is somewhat similar to the correlation of other morphological systems (e.g., aedeagus, spermatheca, and wing venation) of the Alticinae/Galerucinae that demonstrate either a Galerucinae or Alticinae tendency at the generic level [48]. Various authors have made the general statement that there are no differences between larvae of the Galerucinae (sensu stricto) and the Alticinae (sensu stricto) [49–52], but larval studies of the Galerucinae and Alticinae are far from comprehensive and without worldwide representation, that is, poor taxon sampling [53]. Besides, similarities between larvae of certain alticine and galerucine groups may be more closely tied to convergence such as habitat than to actual ancestral identity, that is, relationships [20].
An adult galerucine with the modified characteristics is clearly different from other congeneric adults without modifications. It is not known whether certain larvae of the modified adults are different from the larvae of nonmodified adults. Despite the apparently similar larval morphology, the second author has always maintained that the characteristic of the metafemoral spring (found in both sexes) is unique to and primarily defines the Alticinae, and it differentiates this subfamily from the Galerucinae (sensu stricto) [6]. This characteristic also has the very useful aspect that it can be used to distinguish genera in the vast majority of cases because of its consistent morphology in all species of each genus [54–56]. One study that tried to demonstrate that the metafemoral spring was variable within a genus [7] failed to recognize that when the spring is taken out of the metafemur and allowed to dry out its very thin chitinous structure shrivels up differently depending on many factors such as how it was treated (e.g., chemicals, temperature, etc.) during extraction; therefore, the conclusion in that reference that the metafemoral spring is not a valid intrageneric character is erroneous.
8. Conclusion
There are 1298 species in the subfamily Galerucinae (sensu stricto) with at least one type of sexual secondary characteristics (SSCs). This figure represents 24% of the total members of the subfamily and is a very significant number. The SSCs are so diverse, with modified structures found on every part of the body (head, thorax, and abdomen) and with functions encompassing olfactory and copulatory behaviour, as well as sound production. The presence of SSCs as stridulatory devices functioning to produce sound is certainly a significant characteristic of the subfamily Galerucinae (sensu stricto). The SSCs are not variable but are species specific. They are variable interspecifically a manifestation of amazing biodiversity of the Galerucinae with SSCs. Modified Galerucinae are unparalleled in other subfamily of the Chrysomelidae. Despite reputed larval similarities, these apomorphic characteristics of adult Galerucinae distinguish them from the Alticinae. A high representation of SSCs indicates that they are not exceptional but a characteristic of the subfamily Galerucinae (sensu stricto). These characteristics should be fully utilized in our attempt to understand the Galerucinae and its relationships with other subfamilies in the Chrysomelidae.
AppendicesA. List of Galerucinae Genera Represented by at Least One Species with Secondary Sexual Character (SSC)
Acroxena Baly, 1879—clypeus, antennae, and labrum
Aelianus Jacoby, 1892—antennae
Afroatrachya Weise, 1904—elytra
Afrocrania Hincks, 1949—clypeus, antennae, and elytra
Agetocera Hope, 1840—antennae
Alphidia Clark, 1865—antennae
Anatela Silfverberg, 1982—abdomen
Androlyperus Crotch, 1873—elytra, abdomen
Anisobrotica Bechyne and Bechyne, 1969—antennae
Antsianaka Duvivier, 1891—antennae
Apophylia Thomson, 1858—tarsal claws, antennae, metasternum, abdomen, and tibiae
Arimetus Jacoby, 1903—antennae
Arthrotus Motschulsky, 1857—antennae
Asbecesta Harold, 1877—antennae
Atrachya Dejean, 1837—elytra
Aulacophora Chevrolat, 1837—antennae, vertex, elytra, and eyes
Austrotella Silfverberg, 1975—antennae and elytra
Azlania Mohamedsaid, 1996—clypeus
Bacteriaspis Weise, 1905—pronotum and elytra
Bangprella Kimoto, 1989—antennae
Bonesia Baly, 1865—antennae
Buckibrotica Bechyne and Bechyne, 1969—antennae
Candezea Chapuis, 1879—elytra
Cannonia Hincks, 1949—pronotum and elytra
Cerophysa Chevrolat, 1837—antennae and tibiae
Cerophysella Laboissiere, 1930—antennae
Cerotoma Chevrolat, 1837—clypeus and antennae
Chapuisia Duvivier, 1885—antennae and eyes
Chthoneis Baly, 1864—vertex, clypeus, antennae, and eyes
Clitena Baly, 1864—antennae
Cneoranidea Chen, 1942—maxillary palpi, metasternum, and abdomen
Coeligetes Jacoby, 1884—abdomen and trocanthers
Coelomera Chevrolat, 1837—vertex and maxillary palpi
Coraia Clark, 1865—tibiae
Cornubrotica Bechyne and Bechyne, 1969—antennae, femora, and tibiae
Cyclotrypema Blake, 1966—tibiae and labrum
Deinocladus Blake, 1966—tibiae and antennae
Dercetina Gressitt and Kimoto, 1963—antennae
Diabrotica Chevrolat, 1837—antennae, maxillary palpi, and eyes
Diacantha Chevrolat, 1837—elytra
Dimalianella Laboissiere, 1940—antennae
Dircemella Weise, 1902—antennae
Doryscus Jacoby, 1887—antennae
Dreeus Shute, 1982—antennae and tibiae
Duvivieria Weise, 1903—antennae and tibiae
Eccoptopsis Blake, 1966—clypeus, antennae, and tibiae
Ectmesopus Blake, 1940—antennae
Eleona Fairmaire, 1902—antennae and tibiae
Elyces Jacoby, 1888—elytra
Ephaenidea Gressitt and Kimoto, 1963—tibiae
Erynephala Blake, 1936—tarsal claws
Erythrobapta Weise, 1902—elytra and antennae
Euliroetis Ogloblin, 1936—abdomen
Eusattodera Schaeffer, 1906—clypeus
Exosoma Jacoby, 1903—clypeus and antennae
Fleutiauxia Laboissiere, 1933—clypeus
Geinula Oglobin, 1936—antennae
Gynandrobrotica Bechyne, 1955—clypeus
Halysacantha Laboissiere, 1922—antennae, scutellum, and elytra
Haplosomoides Duvivier, 1890—abdomen, eyes, and clypeus
Hemygascelis Jacoby, 1896—abdomen
Hoplasoma Jacoby, 1884—tibiae and abdomen
Hoplosaenidea Laboissiere, 1933—clypeus, antennae, and tibiae
Huillania Laboissiere, 1921—antennae
Hylaspoides Duvivier, 1892—antennae
Hymnesia Clark, 1865—antennae
Hyphaenia Baly, 1865—antennae, frontal tubercles, clypeus, and eyes
Hystiopsis Blake, 1966—tibiae and clypeus
Inbioluperus Clark, 1993—abdomen
Jacobya Weise, 1901—pronotum
Japonitata Strand, 1922—antennae
Kanahiiphaga Laboissiere, 1931—clypeus
Kinabalua Mohamedsaid, 1997—antennae and metasternum
Kumbalia Mohamedsaid and Takizawa, 2007—antennae
Laetana Baly, 1864—antennae
Laetiacantha Laboissiere, 1921—pronotum, elytra, and antennae
Lamprocopa Hincks, 1949—frons and antennae
Leptaulaca Weise, 1902—antennae and tibiae
Leptoxena Baly, 1888—antennae and femora
Lesnella Laboissiere, 1931—antennae and elytra
Lilophaea Bechyne, 1958—tibiae
Liroetiella Kimoto, 1989—antennae
Liroetis Weise, 1889—abdomen, trochanters, tibiae, and eyes
Lomirana Laboissiere, 1932—elytra
Luperodes Motschulsky, 1858—antennae, eyes, and tibiae
Luperogala Medvedev and Samoderzhenkov, 1989—abdomen
Luperosoma Jacoby, 1891—antennae, abdomen, and tibiae
Lygistus Wilcox, 1965—femora
Macrima Baly, 1878—clypeus
Mahutia Laboissiere, 1917—femora and antennae
Malacorhinus Jacoby, 1887—elytra, antennae, and vertex
Megalognatha Baly, 1878—antennae
Metacoryna Jacoby, 1888—antennae
Metopoedema Duvivier, 1891—antennae
Metrobrotica Bechyne, 1958—clypeus, antennae, and tibiae
Microlepta Jacoby, 1886—antennae
Miltina Chapuis, 1875—antennae
Mimastra Baly, 1865—abdomen
Momaea Baly, 1865—tibiae
Mombasa Fairmaire, 1884—tibiae
Monolepta Chevrolat, 1837—elytra, vertex, and trochanters
Monoleptocrania Laboissiere, 1940—vertex
Neolaetana Laboissiere, 1921—antennae, pronotum, and elytra
Niasia Jacoby, 1889—antennae
Nirina Weise, 1892—antennae
Nirinoides Jacoby, 1903—antennae
Nymphius Weise, 1900—abdomen
Oidomorpha Laboissiere, 1924—antennae
Oorlogia Silfverberg, 1978—antennae
Ornithognathus Thomson, 1858—antennae
Oroetes Jacoby, 1888—antennae, clypeus, pronotum, and tibiae
Orthoxia Clark, 1865—antennae
Paleosepharia Laboissiere, 1936—pronotum and elytra
Palpaenidea Laboissiere, 1933—clypeus and antennae
Palpoxena Baly, 1861—clypeus, labrum, maxillary palpi, and antennae
Parabrotica Bechyne and Bechyne, 1961—antennae and tibiae
Paracanthina Hincks, 1949—pronotum and elytra
Paraplotes Laboissiere, 1933—antennae
Parasbecesta Laboissiere, 1940—antennae
Paratriarius Schaeffer, 1906—antennae and elytra
Paridea Baly, 1886—elytra
Periclitena Weise, 1902—antennae
Phyllecthris Dejean, 1837—antennae and tibiae
Phyllobrotica Chevrolat, 1837—antennae, femora, and abdomen
Phyllobroticella Jacoby, 1894—antennae and elytra
Pimentelia Laboissiere, 1939—antennae
Platybrotica Cabrera and Walsh, 2004—antennae
Platymorpha Jacoby, 1888—tibiae
Platyxantha Baly, 1864—antennae, tibiae, and eyes
Porechontes Blake, 1966—clypeus, antennae, and tibiae
Prosmidia Weise, 1901—pronotum, scutellum, and elytra
Pseudaenidea Laboissiere, 1938—clypeus
Pseudocophora Jacoby, 1884—elytra
Pseudocrania Weise, 1892—elytra
Pseudoluperus Beller and Hatch, 1932—abdomen
Pseudorupilia Jacoby, 1893—antennae
Pseudoscelida Jacoby, 1894—antennae and eyes
Pseudoshaira Beenen, 2007—antennae
Rachicephala Blake, 1966—clypeus
Rohaniella Laboissiere, 1940—eyes and antennae
Ruwenzoria Laboissiere, 1919—antennae
Samoria Silfverberg, 1982—antennae
Sarawakiola Mohamedsaid, 1997—antennae and vertex
Scelida Chapuis, 1875—abdomen and tibiae
Scelidacne Clark, 1998—abdomen and tibiae
Scelolyperus Crotch, 1874—tibiae
Schematiza Chevrolat, 1837—antennae
Sermyloides Jacoby, 1884—clypeus and antennae
Sesselia Laboissiere, 1931—antennae
Shungwayana Silfverberg, 1975—antennae
Sikkimia Duvivier, 1891—antennae
Simopsis Blake, 1966—antennae and tibiae
Sinoluperoides Kimoto, 1989—antennae
Sonchia Weise, 1901—elytra
Spilocephalus Jacoby, 1888—clypeus
Spilonotella Cockerell, 1905—antennae
Stenellina Cockerell, 1905—antennae
Stenoplatys Baly, 1861—antennae and tibiae
Strobiderus Jacoby, 1884—elytra
Synetocephalus Fall, 1910—antennae and tibiae
Taenala Silfverberg, 1978—antennae and pronotum
Taphinella Jacoby, 1899—antennae
Taumacera Thunberg, 1814—antennae, metasternum, clypeus, and tibiae
Taumaceroides Lopatin, 2009—antennae, femora, and tibiae
Theopea Baly, 1864—clypeus and antennae
Therpis Weise, 1900—antennae
Trichobrotica Bechyne, 1956—tibiae
Trichomimastra Weise, 1922—antennae
Vitruvia Jacoby, 1903—antennae
Xenarthra Baly, 1861—antennae
Xenoda Baly, 1877—antennae
Zinjotella Silfverberg, 1975—antennae and elytra
B. List of Galerucinae (Sensu Stricto) with Secondary Sexual Characters (SSCs)
Acroxena clypeata (Baly, 1888)—clypeus excavated, with a median projection; labrum large; antennae 3rd, 4th, and 5th thickened
A. femoralis Kimoto, 1989—clypeus excavated; antennae 1st enlarged, 3rd thickened
A. fulva Kimoto, 1989—clypeus transversely excavated, with tuft of hairs; antennae 1st enlarged, 3rd thickened
A. indica Jacoby, 1896—clypeus deeply excavated, with a median projection; labrum large subtriangular; antennae 3rd–10th under surface covered with long erect hairs
A. nasuta Baly, 1879—clypeus slightly excavated, with two median projections; labrum large, concave, with tuft of hairs; antennae 3rd, 4th, 5th, and 6th thickened
Aelianus scutellatus Jacoby, 1892—antennae dilated, covered with long hairs
Afroatrachya impressus (Weise, 1904)—elytra with postscutellar cavities
Afrocrania aequatoriana Wagner, 2007—elytra with a shallow heart-shaped postscutellar depression
A. assimilis (Weise, 1903)—clypeus deeply excavated; antennae 4th curved, with long distal edge
A. famularis (Weise, 1904)—elytra with small subscutellar markings, visible as dense micro-sculpture
A. foveolata (Karsch, 1882)—clypeus deeply excavated; antennae 4th and 5th each with a long distal edge
A. kaethae Middelhauve and Wagner, 2001—clypeus shallowly excavated; antennae 4th curved, with a long distal edge
A. kakamegaensis Middelhauve and Wagner, 2001—elytra excavated at base, with two small protruding bulges, and at the middle with oval shaped elevation and shallowed groove
A. latifrons Weise, 1892—clypeus deeply excavated; antennae 4th curved inwards
A. longicornis Middelhauve and Wagner, 2001—elytra with longitudinal postscutellar extrusions
A. luciae Middelhauve and Wagner, 2001—clypeus unevenly, deeply excavated; antennae 4th curved with a long distal edge
A. nigra Wagner, 2007—elytra with a small heart-shaped postscutellar depression
A. occidentalis Wagner, 2007—elytra with a keel-like postscutellar extrusion and shallow depression
A. pallida Wagner, 2007—elytra with a shadow-like spot in basal half
A. pauli (Weise, 1903)—elytra with a small hump-backed postscutellar extrusion
A. ubatubae Middelhauve and Wagner, 2001—elytra with a shallow depression
A. weisei Wagner, 2007—elytra with postscutellar extrusion keel-liked and a shallow drop-shaped depression
A. aeruginosa (Hope, 1831)—tarsal claws bifid; abdomen 1st and 2nd, each with a pair of appendages
A. algie Bezděk, 2008—tarsal claws bifid
A. angolensis Laboissiere, 1921—tarsal claws bifid
A. angustata Allard, 1889—tarsal claws bifid
A. asahinai Chujo, 1962—tarsal claws bifid
A. assamensis (Jacoby, 1891)—tarsal claws bifid
A. aurolimbata Allard, 1888—tarsal claws bifid
A. basilana Pic, 1945—tarsal claws bifid; metasternum with a small tubercle
A. beeneni Bezděk, 2003—tarsal claws bifid; metasternum with a small tubercle
A. bertiae Bezděk, 2003—tarsal claws bifid; antennae 1st–5th covered with very long hairs; metasternum with a large tubercle
A. bifasciata Allard, 1889—tarsal claws bifid
A. blecha Bezděk, 2008—tarsal claws bifid
A. borowiecki Bezděk, 2004—tarsal claws bifid
A. brancucci Medvedev and Sprecher, 1999—tarsal claws bifid
A. carinata Laboissiere, 1922—tarsal claws bifid
A. celebensis Pic, 1927—tarsal claws bifid; metasternum with a bifurcate process
A. cervenki Bezděk, 2005—tarsal claws bifid
A. cheni Bezděk and Zhang, 2006—tarsal claws bifid
A. chloroptera Thomson, 1858—tarsal claws bifid; mesotarsi 1st with deeply incised
A. clavareaui Laboissiere, 1940—tarsal claws bifid
A. clavicornis Samoderzhenkov, 1988—tarsal claws bifid; antennae 9th, 10th, and 11th strongly dilated
A. clypeata Samoderzhenkov, 1988—tarsal claws bifid
A. consanguinea Allard, 1889—tarsal claws bifid
A. crassicornis Laboissiere, 1920—tarsal claws bifid; antennae 4th–7th dilated
A. curvipes Laboissiere, 1920—tarsal claws bifid; metatibiae curved
A. cyaneolimbata Laboissiere, 1922—tarsal claws bifid
A. cyanipennis Laboissiere, 1927—tarsal claws bifid
A. dellacasai Bezděk, 2006—tarsal claws bifid
A. dembickyi Bezděk, 2006—tarsal claws bifid
A. demeyeri Bezděk, 2005—tarsal claws bifid; antennae 5th enlarged
A. denisae Bezděk, 2005—tarsal claws bifid
A. dilaticornis (Jacoby, 1894)—tarsal claws bifid; antennae 3rd and 4th enlarged, oblong, 5th–7th enlarged, dilated at apex
A. disconotata Pic, 1947—tarsal claws bifid
A. elongata (Jacoby, 1896)—tarsal claws bifid
A. elschotziae Chen, 1976—tarsal claws bifid; antennae 8th enlarged
A. eoa Oglobin, 1936—tarsal claws bifid
A. epipeluralis Laboissiere, 1927—tarsal claws bifid
A. eroshkinae Samoderzhenkov, 1988—tarsal claws bifid
A. excavata Bryant, 1954—tarsal claws bifid
A. femorata (Jacoby, 1895)—tarsal claws bifid
A. flavovirens (Fairmaire, 1878)—tarsal claws bifid
A. frischi Bezděk, 2003—tarsal claws bifid
A. furcigera Chujo, 1962—tarsal claws bifid; metasternum with a long bifurcate process
A. ghesquierei Laboissiere, 1940—tarsal claws bifid
A. gloriosa Laboissiere, 1922—tarsal claws bifid
A. grandicornis (Fairmaire, 1888)—tarsal claws bifid
A. grobbelaarae Bezděk, 2006—tarsal claws bifid
A. hajeki Bezděk, 2003—tarsal claws bifid; abdomen 1st and 2nd, each with a pair of appendages
A. haladai Bezděk, 2006—tarsal claws bifid
A. halberstadti Baezdek, 2006—tarsal claws bifid
A. hanka Bezděk, 2005—tarsal claws bifid
A. hebes Weise, 1904—tarsal claws bifid; metasternum with a bifurcate process; metafemora very large
A. holosericea Laboissiere, 1925—tarsal claws bifid
A. incisitarsis (Laboissiere, 1922)—tarsal claws bifid; antennae 5th greatly enlarged
A. jeanneli Laboissier, 1921—tarsal claws bifid
A. jolantae Bezděk, 2007—tarsal claws bifid
A. kaffa Bezděk, 2005—tarsal claws bifid
A. kantneri Bezděk, 2003—tarsal claws bifid; antennae 3rd–7th dilated
A. kaoi Bezděk and Lee, 2009—tarsal claws bifid
A. keniaensis Laboissiere, 1920—tarsal claws bifid
A. kimotoi Bezděk, 2003—tarsal claws bifid
A. kubani Bezděk, 2005—tarsal claws bifid; antennae 9th, 10th, and 11th dilated
A. laotica Bezděk, 2005—tarsal claws bifid
A. laticollis Laboissiere, 1922—tarsal claws bifid; antennae 4th–7th dilated
A. lebongana Maulik, 1936—tarsal claws bifid; antennae 3rd–7th dilated, funnel shaped, 9th dilated, triangular; abdomen 1st with a pair of short appendages
A. leontovitchi Laboissiere, 1940—tarsal claws bifid; antennae 5th enlarged
A. lesnei Laboissiere, 1922—tarsal claws bifid
A. levi Bezděk, 2004—tarsal claws bifid; antennae 9th, 10th, and 11th strongly dilated
A. libenae Bezděk, 2007—tarsal claws bifid
A. lindae Bezděk, 2006—tarsal claws bifid
A. liska Bezděk, 2008—tarsal claws bifid
A. luzonica Bezděk, 2003—tarsal claws bifid
A. maculata Kimoto, 1977—tarsal claws bifid
A. maculicollis (Jacoby, 1895)—tarsal claws bifid; metasternum with a small tubercle
A. marginata Jacoby, 1899—tarsal claws bifid
A. marginicollis Laboissiere, 1940—tarsal claws bifid
A. marginipennis Weise, 1922—tarsal claws bifid
A. marketae Bezděk, 2006—tarsal claws bifid
A. marshalli (Jacoby, 1897)—tarsal claws bifid
A. matrensi Bezděk, 2003—tarsal claws bifid
A. mauritanica Pic, 1944—tarsal claws bifid
A. maynei Labossiere, 1922—tarsal claws bifid; metatibiae curved
A. medvedevi Samoderzhenkov, 1988—tarsal claws bifid
A. melli Gressitt and Kimoto, 1963—tarsal claws bifid
A. metallica Jacoby, 1904—tarsal claws bifid
A. micheli Bezděk, 2001—tarsal claws bifid; abdomen 1st and 2nd, each with a pair of short appendages
A. mikhailovi Bezděk, 2003—tarsal claws bifid; metasternum with a large tubercle
A. mila Bezděk, 2005—tarsal claws bifid
A. mimica Samoderzhenkov, 1988—tarsal claws bifid
A. miyamotoi Kimoto, 1969—tarsal claws bifid
A. neavei Bezděk, 2005—tarsal claws bifid
A. nepalica Bezděk, 2003—tarsal claws bifid
A. nigriceps Laboissiere, 1927—tarsal claws bifid
A. nigricollis Allard, 1888—tarsal claws bifid
A. nigrolimbata Laboissiere, 1940—tarsal claws bifid
A. nilakrishna Maulik, 1936—tarsal claws bifid; antennae 5th–7th dilated; abdomen 1st and 2nd, each with a pair of appendages
A. nobilitata Gerstaecker, 1871—tarsal claws bifid; metasternum with a bifurcate process
A. nodicornis Laboissiere, 1922—tarsal claws bifid; antennae 5th extremely large
A. oborili Bezděk, 2005—tarsal claws bifid
A. pacholatkoi Bezděk, 2005—tarsal claws bifid; antennae 9th, 10th, and 11th dilated
A. pallipes (Jacoby, 1892)—tarsal claws bifid; antennae 9th, 10th, and 11th strongly dilated
A. pavlae Bezděk, 2003—tarsal claws bifid
A. pectoralis Pic, 1927—tarsal claws bifid; metasternum with a bifurcate process
A. pesai Bezděk, 2006—tarsal claws bifid
A. phuphanensis Bezděk, 2006—tarsal claws bifid
A. poggii Bezděk, 2003—tarsal claws bifid
A. porraceipennis (Allard, 1889)—tarsal claws bifid
A. pulchella Bryant, 1952—tarsal claws bifid
A. purpurea (Allard, 1888)—tarsal claws bifid; metafemora enlarged
A. quadristigmata Laboissiere, 1922—tarsal claws bifid
A. raffrayi Pic, 1946—tarsal claws bifid
A. rugiceps Gressitt and Kimoto, 1963—tarsal claws bifid
A. saliens Weise, 1904—tarsal claws bifid; metasternum with a bifurcate process; metafemora very large
A. samoderzhenkovi Medvedev, 1993—tarsal claws bifid
A. savioi Pic, 1931—tarsal claws bifid
A. schawalleri Medvedev, 1992—tarsal claws bifid; abdomen 1st with a pair of short appendages
A. securigera Chujo, 1962—tarsal claws bifid
A. semiobscura Faimaire, 1887—tarsal claws bifid
A. sericea (Fabricius, 1798)—tarsal claws bifid
A. shuteae Bezděk, 2003—tarsal claws bifid
A. sikkimensis Bezděk, 2003—tarsal claws bifid
A. similis Weise, 1909—tarsal claws bifid
A. snizeki Bezděk, 2005—tarsal claws bifid; metasternum with a bifurcate process
A. sosia Laboissiere, 1940—tarsal claws bifid
A. sprecherae Bezděk, 2003—tarsal claws bifid
A. sulcata Laboissiere, 1922—tarsal claws bifid
A. taiwanica Bezděk, 2003—tarsal claws bifid
A. takizawai Bezděk, 2005—tarsal claws bifid
A. tarsalis Laboissiere, 1938—tarsal claws bifid
A. thalassina Faldermann, 1835—tarsal claws bifid
A. trapezicollis Laboissiere, 1940—tarsal claws bifid
A. trinotata Gressitt and Kimoto, 1963—tarsal claws bifid
A. trochanterina Gressitt and Kimot, 1963—tarsal claws bifid
A. velai Bezděk, 2003—tarsal claws bifid
A. variicollis Laboissiere, 1922—tarsal claws bifid
A. vernalis (Allard, 1889)—tarsal claws bifid
A. vicinia Laboissiere, 1940—tarsal claws bifid
A. vietnamica Samoderzhenkov, 1988—tarsal claws bifid
A. viridipennis (Jacoby, 1885)—tarsal claws bifid
A. viridis (Jacoby, 1884)—tarsal claws bifid
A. voriseki Bezděk, 2003—tarsal claws bifid
A.weisei (Jacoby, 1896)—tarsal claws bifid
A. wittei Laboissiere, 1940—tarsal claws bifid
A. yangi Bezděk and Zhang, 2006—tarsal claws bifid
B. serricornis (Thomson, 1858)—antennae 4th–8th serrate
Buckibrotica cinctipennis (Baly, 1886)—antennae 7th and 9th dilated, excavated
Candezea atomaria (Fairmaire, 1875)—elytra with a heart-shaped postscutellar protrusion
C. biocostata (Weise, 1907)—elytra with one pair of short postscutellar protrusions with ridges and one pair of long lateral protrusions with ridges, from humeri to apical third
C. costatipennis Laboissiere, 1931—elytra with a heart-shaped postscutellar protrusion and a pair of lateral protrusions with ridges, from humeri to apical third
C. flaveola (Gerstaecker, 1855)—elytra with a heart-shaped postscutellar protrusion
C. franzkrappi Wagner and Kutrscheid, 2005—elytra with a deep crater-like postscutellar depression
C. haematura (Fairmaire, 1891)—elytra with a heart-shaped postscutellar protrusion
C. irregularis (Ritsema, 1893)—elytra with a deep oval shaped postscutellar depression
C. occipitalis (Reiche, 1847)—elytra with a heart-shaped postscutellar protrusion
Cannonia confusa Silfverberg, 1971—pronotum bulged in the middle, the posterior margin with a tongue-shaped process, overhang scutellum; elytra with a pair of large tubercles at base
C. meridionalis (Weise, 1901)—pronotum bulged in the middle and a tongue-shaped process on posterior margin overhang scutellum; elytra with a pair of large tubercles at base
C. occidentalis (Weise, 1901)—pronotum with a short, concave tongue-shaped process on posterior margin overhang scutellum.
C. petersii (Bertoloni, 1868)—pronotum with a short tongue-shaped process on posterior margin overhang scutellum
C. basalis (Baly, 1874)—elytra with elongate postscutellar depression
C. plagiata Laboissiere, 1930—antennae 3rd–11th covered with long hairs
C. viridipennis (Allard, 1889)—pronotum with a pair of tubercles before transverse depression; elytra with around postscutellar depression
Cerotoma atrofasciata Jacoby, 1879—clypeus deeply excavated, without tubercle; antennae 3rd incised at apex, 4th triangular shaped
C. dilatipes Jacoby, 1888—clypeus depressed; front tibiae dilated at apex
C. ruficornis (Olivier, 1791)—clypeus deeply excavated, with a very prominent median tubercle; antennae 3rd dilated, incised at apex, 4th triangular shaped
C. salvini Baly, 1866—clypeus excavated, with a prominent median tubercle; antennae 3rd dilated, incised at apex, 4th triangular shaped
Dreeus distinctus Shute, 1983—antennae 9th and 11th enlarged; mesotibiae curved
Duvivieria africana (Duvivier, 1892)—metatibiae with a long process at apex
D. apicitarsis (Weise, 1903)—antennae 7th elongate, strongly curved, crescent shaped
Eccoptopsis argentinensis Blake, 1966—clypeus moderately excavated, with two short spines on each side; antennae 3rd and 4th enlarged, excavated
E. boliviensis Blake, 1966—clypeus excavated with spines; antennae 3rdand 4th enlarged, excavated
E. cavifrons Blake, 1966—clypeus excavated, with a small spine in middle; antennae 3rd and 4th enlarged, excavated
E. clara Blake, 1966—clypeus excavated, with spines; antennae 3rd and 4th enlarged, excavated
E. costaricensis Blake, 1966—clypeus deeply excavated, with a pair of long, curved spines at sides; antennae 3rd and 4th enlarged, excavated
E. cyanocositiesa Blake, 1966—clypeus moderately excavated; antennae 3rd and 4th enlarged, excavated
E. denticornis (Jacoby, 1887) clypeus excavated with spines; antennae 3rd and 4th enlarged, excavated
E. laticollis Blake, 1966—clypeus deeply excavated, with a pair of long curved spines at sides; antennae 3rd and 4th enlarged and excavated
E. mexicana Blake, 1966—clypeus excavated, with a long thick spine in the middle; antennae 3rd longitudinally depressed at sides, 4th curved, excavated
E. piceofasciata Blake, 1966—clypeus excavated, with a short spine in the middle; antennae 3rd long, excavated, 4th enlarged, slightly excavated; front tibiae enlarged
E. quadriniaculata Blake, 1966—clypeus deeply excavated, with fringe of hairs overhanging the cavity on both sides; antennae 3rd and 4th enlarged
E. quadrimaculata Blake, 1966—clypeus excavated with spines; antennae 3rd and 4th enlarged
Ephaenidea indochinensis Medvedev, 2004—mesotibiae with a long projection at apex
Erynephala brighti Blake, 1970—tarsal claws bifid
E. glabra Blake, 1936—tarsal claws bifid
E. puncticollis (Say, 1824)—tarsal claws bifid
E. morosa (Leconte, 1857)—tarsal claws bifid
E. maritime (Leconte, 1865)—tarsal claws bifid
Erythrobapta benigseni Weise, 1902—elytra with shallow postscutellar cavities
Euliroetis lameyi (Laboissiere, 1929)—abdomen 4th and 5th excavated in the middle
E. melanocephala (Bowditch, 1925)—abdomen 2nd with a pair of appendages, 4th deeply excavated
E. nigronotum Gressitt and Kimoto, 1963—abdomen 1st with a pair of short appendages, 2nd and 3rd each with a pair of very long appendages, 4th excavated
E. ornata (Baly, 1874)—abdomen 1st and 2nd each with a pair of appendages
E. suturalis (Laboissiere, 1929)—abdomen 2nd with a pair of very broad appendages, 4th deeply excavated
E. deformicornis (Quedenfeldt, 1888)—antennae 3rd–11th thickened, 8th enlarged, oblong
E. discoidale (Jacoby, 1895)—antennae 3rd–8th thickened
E. tripunctata (Jacoby, 18 )—elytra with cavities at sides
Fleutiauxia armata (Baly, 1874)—clypeus transversely excavated, with a projection on posterior ridge
F. bicavifrons Gressitt and Kimoto, 1963—clypeus with a large deep cavity on each side of the middle, with a short projection on anterior and posterior of the cavity
F. cyanipennis Laboissiere, 1933—clypeus with transversely deeply excavated, with a short projection in the middle
F. multifrons Gressitt and Kimoto, 1963—clypeus transversely excavated, with a tall projection, and an anterior projection on posterior ridge
F. septentrionalis (Weise, 1922)—clypeus transversely excavated, with a backward projection on posterior ridge
F. violaceipennis Kimoto, 1989—clypeus transversely excavated, with a long projection
Gynandrobrotica imitans (Jacoby, 1879)—clypeus flattened as single piece; maxillary palpi swollen
G. nigrofasciata (Jacoby, 1887)—clypeus flattened as a single piece; maxillary palpi swollen
G. parambaensis (Bowditch, 1912)—clypeus broadly, smoothly, deeply excavated, sides with rows of long hairs; maxillary palpi swollen
Halysacantha weisei (Jacoby, 1899)—antennae 3rd broadly dilated, deeply excavated; scutellum excavated, with sides raised into high ridges; elytra with a pair of tubercles at base
H. antennalis Mevdvedev, 2000—antennae 4th–7th covered with long hairs
H. appendiculata Laboissiere, 1930—eyes large; abdomen 1st with a long appendage
H. biclavata Jiang, 1988—abdomen 2nd with a pair of short and 3rd a pair of long appendages
H. brushei Jang, 1988—abdomen 1st with a brush-like appendage
H. changi Lee, Bezděk and Staines, 2011—abdomen 1st with a long appendage, the apical third lateral margins with rows of long hairs
H. chengi Lee, Bezděk and Staines, 2011—abdomen 1st with a long appendage and its apical third densely covered with short setae
H. chinmatra Maulik, 1936—abdomen 1st with a long appendage; antennae thickened towards apex
H. costata Baly, 1878—abdomen 1st with a long appendage
H. curvipes Medvedev, 2000—hind tibiae curved
H. egena Weise, 1922—abdomen 1st with a slender appendage
H. flava Laboissiere, 1930—eyes large
H. hainana Jang, 1988—abdomen 1st with a brush like appendage
H. himalayana Medvedev, 2002—antennae 7th–11th thickened; abdomen 5th with a short pointed appendage
H. laticornis Laboissiere, 1930—antennae 7th–11th dilated; abdomen 1st with a long appendage
H. nigricollis Jang, 1988—abdomen 3rd with a pair of appendages
H. ryukyuensis Lee, Bezděk and Staines, 2011—abdomen 1st with a long appendage, the apical third lateral margins with rows of long hairs
Hemygascelis longicollis Jacoby, 1896—abdomen 1st tuberculate, 2nd, 3rd, and 4th with long erect hairs, 5th elongate, densely covered with hairs along the middle
Hoplasoma apicale Jacoby, 1884—abdomen 1st with a pair of short appendages, 2nd with a long pair of appendages
H. bosi Bezděk, 2008—abdomen 2nd with a pair of narrow and very short appendages
H. celebense Jacoby, 1886—abdomen 2nd with a pair of appendages
H. costatipennis Jacoby, 1896—abdomen 2nd with tubercles
H. dilaticornis Jacoby, 1900—antennae 8th, 9th, and 10th dilated; hind tibiae curved
H. luzonica Medvedev, 2002—abdomen 2nd with a pair of appendages, widely separated
H. mindanensis Medvedev, 2002—abdomen 2nd with a pair of appendages
H. minor Gressitt and Kimoto, 1963—abdomen 2nd with a pair of appendages
H. nilgiriensis Jacoby, 1904—abdomen 2nd with a pair of short, flat appendages
H. paradoxum Medvedev, 2007—abdomen 2nd with a pair of appendages
H. philippinensis Jacoby, 1894—abdomen 2nd with a pair of short pointed appendages
H. rostripenne Allard, 1888—abdomen 2nd with a pair of large appendages
H. sulawesianum Medvedev, 2007—abdomen 2nd with a pair of large appendages; hind tibiae dilated at apex
H. unicolor (Illiger, 1800)—abdomen 2nd with a pair of appendages
Hoplosaenidea abdominalis (Jacoby, 1884)—hind tibiae curved, dilated, and compressed at the middle
H. aerosa (Laboissiere, 1933)—clypeus deeply excavated, with a prominent tubercle in the middle; antennae 3rd and 4th enlarged
H. apicipennis Baly, 1888—clypeus broadened, strongly raised, with median longitudinal groove
H. capitata (Jacoby, 1886)—clypeus excavated, with a pair of pointed projections
H. cavifrons (Duvivier, 1885)—clypeus very deeply excavated, with a pair of flap-like projections
H. citrina (Jacoby, 1894)—protibiae dilated at apex
H. coomani (Laboissiere, 1933)—clypeus shallowly excavated
H. cornuta (Laboissiere, 1933)—clypeus deeply, broadly excavated, with a tubercle
H. elegans Kimoto, 1989—clypeus deeply, broadly excavated
H. facialis (Baly, 1888)—clypeus excavated, with a triangular, flat tubercle in basal half
H. fragilis Gressitt and Kimoto, 1963—clypeus very deeply excavated
H. laosensis Kimoto, 1989—clypeus deeply, broadly excavated
H. merah Mohamedsaid, 2001—antennae 4th dilated, compressed
H. monstrosa (Jacoby, 1896)—clypeus deeply excavated, with a short projection on frons
H. nigripennis (Laboissiere, 1940)—clypeus excavated
H. nigrolimbata (Jacoby, 1899)—metatibiae dilated and curved at the middle
H. nitida Gressitt and Kimoto, 1963—clypeus moderately excavated
H. ocellata (Baly, 1888)—clypeus deeply excavated, with 2 pairs of tubercles
H. porrecta (Baly, 1865)—clypeus excavated, with a projection; antennae 1st dilated, with a pointed distal edge
H. pulchella (Laboissiere, 1933)—clypeus shallowly excavated
H. sarah Mohamedsaid, 2009—clypeus deeply excavated, with a projection in middle; antennae 1st dilated, with a pointed distal edge
H. semilimbata (Jacoby, 1894)—clypeus deeply excavated, with a pair of pointed spines on frons
H. semperi (Jacoby, 1894)—clypeus deeply excavated, with a horn-like projection on frons
H. subcostata (Jacoby, 1884)—antennae 3rd and 4th compressed, dilated
H. singaporensis Mohamedsaid, 2002—clypeus excavated; antennae 1st dilated, with a pointed distal edge
H. takizawai Mohamedsaid, 2000—clypeus deeply excavated, with a projection; antennae 1st dilated, with a pointed distal edge
H. touzalini Laboissiere, 1933—metatibiae with a long curved process at apex
H. tripunctata (Jacoby, 1894)—clypeus depressed; eyes large
H. variabilis (Jacoby, 1894)—clypeus with a narrow, elongate cavity
H. violacea (Jacoby, 1892)—clypeus deeply excavated, with a projection in the middle
Hymnesia tranquebarica (Fabricius, 1798)—antennae 8th enlarged, concave on ventral surface
Hyphaenia aenea Laboissiere, 1936—antennae 3rd–11th densely covered with long hairs
H. africana Laboissiere, 1921—antennae 3rd–11th densely covered with long hairs
H. antennalis Kimoto, 1989—antennae 3rd–5th covered with long hairs
H. apicicornis Jacoby, 1896—antennae 3rd–11th covered with long hairs
H. azlani Mohamedsaid, 1998—antennae 3rd–8th covered with long hairs
H. bicolor Medvedev, 2001—antennae 3rd–5th covered with long hairs; clypeus excavated, with triangular tooth at anterior margin
H. clypealis Medvedev, 2001—antennae 3rd–11th covered with long hairs; clypeus flat with long hairs
H. cyanescens Laboissiere, 1936—antennae 3rd–11th covered with long hairs
H. discoidalis Jacoby, 1886—antennae 3rd–8th covered with long hairs
H. frontalis Kimoto, 1989—clypeus excavated, with a projection at posterior margin
H. fulva Kimoto, 1989—antennae 8th and 9th curved
H. indica (Jacoby, 1903)—antennae 3rd–8th covered with erect hairs
H. keralensis Medvedev, 2001—antennae 3rd–6th with covered short erect hairs
H. kimotoi Medvedev, 2001—clypeus deeply excavated
H. mandibularis Medvedev, 2001—antennae 3rd–8th with long hairs
H. nigricornis Kimoto, 1989—clypeus broadly excavated
H. nigrilabris Medvedev, 2001—antennae 4th curved
H. nitidissima Medvedev, 2001—antennae 3rd–11th covered with long hairs
H. obscuripennis Jacoby, 1896—antennae 3rd–8th covered with long hairs
H. oculata Mohamedsaid, 1999—antennae 3rd–8th covered with long hairs; eyes very large; maxillary palpi swollen
H. pallida Medvedev, 2001—antennae 3rd and 4th covered with long hairs
H. pilicornis (Motschulsky, 1858)—antennae 3rd–8th with long hairs
H. rahmani Mohamedsaid, 1999—antennae extremely long, 3rd–8th covered with long hairs and 3rd–11th longitudinally ridged; frontal tubercles prominent, very large; eyes large
H. rubra Medvedev, 2001—antennae 3rd–8th covered with long hairs
H. submetallica Jacoby, 1892—antennae 3rd–8th covered with long hairs
H. tristis Medvedev, 2001—antennae 9th and 10th dilated, excavated; clypeus excavated
H. volkovitshi Lopatin, 2009—antennae 3rd–11th covered with long hairs; clypeus deeply excavated
H. marginalis (Fabricius, 1801)—protibiae enlarged
H. terminalis Blake, 1966—protibiae enlarged
H. zonata Blake, 1966—protibiae thickened
Inbioluperus costipennis Clark, 1993—abdomen 2nd with a short rectangular appendage
Jacobya cavicollis (Fairmaire, 1880)—pronotum deeply, broadly, triangularly excavated, with a tubercle in the middle
J. notabilis Weise, 1902—pronotum with basal half deeply excavated
J. ochracea Weise, 1901—pronotum entirely, deeply excavated
J. pilosa Weise, 1902—pronotum with basal half deeply excavated
J. viridis Weise, 1904—pronotum with middle area deeply excavated
Japonitata abdominalis Jiang, 1989—antennae 9th with a spine at apex, 10th enlarged, 11th larger, longer than 1st
J. antennata Chen and Jiang, 1986—antennae 8th thickened, 9th minute, 10th dilated, 11th larger, longer than 1st
J. hongpingana Jiang, 1989—antennae 8th dilated, 9th dilated, shorter than 9th, 10th dilated, pointed at apex, 11th longer than the first segment
Kanahiiphaga aeneipennis Laboissiere, 1937—clypeus transversely excavated, with a projection
K. carpenteri Laboissiere, 1937—clypeus transversely excavated, with a projection
K. costipennis Laboissiere, 1936—clypeus transversely excavated, with a projection
K. costulata Laboissiere, 1931—clypeus transversely excavated, with a projection
K. frontalis Laboissiere, 1936—clypeus transversely excavated, with a projection
K. orphana (Chapuis, 1879)—clypeus transversely excavated, with a projection
K. similis Laboissiere, 1936—clypeus transversely excavated, with a projection
Kinabalua antennata Mohamedsaid, 1997—antennae 7th enlarged, excavated at apex, 8th enlarged, with a broad, short spine; metasternum with a process
Kinabalua musaamani Mohamedsaid, 2010—antennae 7th enlarged, excavated at apex, 8th enlarged, with a long, curved spine; metasternum with a process
Kumbalia longicornis Mohamedsaid and Takizawa, 2007—antennae extremely long, beyond elytra
Laetana histrio Baly, 1864—antennae with segments 3rd–7th dilated
Laetiacantha amabilis (Laboissiere, 1940)—antennae 3rd–7th dilated; pronotum with a small, elliptical depression on each side of midline at posterior margin; elytra with a pair of tubercles at the base
L. distincta (Gahan, 1893)—antennae 3rd–7th dilated towards apex; pronotum with small sharp elevations at posterior margin; elytra with a pair of blunt tubercles at base
L. elegans Laboissiere, 1923—antennae 3rd–7th dilated; pronotum with sharp elevation at posterior margin; elytra with a pair of blunt tubercles at the base
L. freynei Silfverberg, 1975—antennae 3rd–7th dilated; pronotum with sharp elevation at posterior margin; elytra with a pair of tubercles at base
L. maynei (Laboissiere, 1921)—antennae 3rd–7th dilated; elytra with a pair of pointed tubercles at the base
L. ruficollis Laboissiere, 1921—antennae 3rd–7th dilated; pronotum strongly impressed with sharp elevation at posterior margin; elytra with a pair of blunt tubercles at the base
L. simillima Silfverberg, 1975—antennae 3rd–7th dilated; pronotum with a semicircular projection in the middle at posterior margin; elytra with a pair of tubercles at the the base
L. splendens Silfverberg, 1975—antennae 3rd–7th dilated; pronotum with semicircular projection in middle at posterior margin; elytra with a pair of tubercles at base
L. subsudanica (Weise, 1907)—antenna 3rd–7th dilated; pronotum with a small depression on each side of midline at posterior margin; elytra with a pair of tubercles at the base
L. verax Silfverberg, 1975—antennae 3rd–7th dilated; pronotum with sharp elevation at posterior margin; elytra with a pair of tubercles at base
Lamprocopa antennata (Weise, 1903)—frons excavated, with a projection
L. dalata (Erichson, 1843)—frons excavated, with a projection
L. femoralis (Laboissiere, 1929)—frons excavated, with a projection
L. kunowi (Weise, 1892)—frons excavated, with a projection
L. nigripennis (Laboissiere, 1921)—frons excavated, with a projection
L. occidentalis (Weise, 1895)—frons excavated, with a projection
L. orientalis (Weise, 1903)—frons excavated, with a projection
L. praecox (Klug, 1833)—frons excavated, with a projection
L. rothschildi (Laboissiere, 1920)—frons excavated, with a projection
L. seabrai (Gomez Alves, 1951)—frons excavated, with a projection
Leptaulaca nigra Laboissiere, 1920—antennae with long erect hairs
L. nigricornis Weise, 1902—antennae with long erect hairs
L. pusila Weise, 1912—antennae with long erect hairs
L. undecimpunctata (Klugg, 1833)—antennae with long erect hairs
L. venusta Laboissiere, 1930—antennae with long erect hairs; front tibiae curved
Leptoxena eximia Baly, 1888—antennae 9th, 10 and 11th compressed; hind femora incrassate; hind tibiae channelled, with a process at apex
Lesnella fasciata Laboissiere, 1931—antennae 4th–7th thickened; elytra with elevated postscutellar area at suture
L. pygidialis Laboissiere, 1921—antennae 4th thickened
L. rufus Harold, 1877—antennae 4th thickened
Luperogala mirabilis Medvedev and Samoderzhenkov, 1989—abdomen 4th with a sword-like, curved appendage, 5th with a pair of lateral appendages
L. paradoxa Medvedev and Samoderzhenkov, 1989—abdomen 4th with a curved, stylet-shaped appendage, dilated at apex, 5th with a pair of lateral appendages
Luperosoma atlanta (Bechyne, 1950)—antennae 9th, 10th, and 11th enlarged; mesotibiae notched
L. bechynei (Blake, 1966)—antennae 9th, 10th, and 11th enlarged; mesotibiae notched
L. latifrons (Blake, 1966)—antennae 9th, 10th, and 11th enlarged; mesotibiae notched
L. nigricolle Blake, 1966—antennae 8th–11th thickened; mesotibieae notched
L. parallelum (Horn, 1893)—antennae 11th swollen; mesotibiae notched on inner side
L. parvalum (Jacoby, 1888)—antennae 9th–10th thickened; mesotibiae notched
L. schwarzi (Horn, 1896)—antennae 11th swollen; abdomen 1st with a prominent tubercle; mesotibiae notched on inner side
L. subsulcatum (Horn, 1893)—antennae 11th swollen; mesotibiae notched on inner side
Metrobrotica furcata (Olivier, 1808)—clypeus with a small hole in the middle, surrounded by funnel-shaped structure; antennae 3rd very long; protibia thickened
M. geometrica (Erichson, 1847)—antennae 3rd and 4th enlarged
Miltina dilatata Chapuis, 1875—antennae 4th–10th compressed, each segment with a long projection
Mimastra andrewesi Bezděk, 2010—abdomen 2nd a tubercle in middle
M. gracilis Baly, 1878—antennae 3rd–11th densely covered with long hairs
M. procerula Zhang and Yang, 2006—abdomen 3rd and 4th each with a triangular appendage on the posterior margin
Momaea distincta Mohamedsaid, 1999—mesotibiaee excavated at apex, covered with long hairs; first segment of mesotarsus with a process projecting towards mesotibiae
Mombasa armicollis Fairmaire, 1884—protibia strongly dilated at apex
M. magna (Weise, 1900)—protibiae strongly dilated at apex
M. subinermis Fairmaire, 1884—protibiae strongly dilated at apex
Monolepta armatipennis Medvedev, 2005—elytra postscutellar area with a drop-like impression, delimited by tubercle in front
M. azlani Mohamedsaid, 1998—elytra with postscutellar cavities
M. bicavipennis Chen, 1942—elytra with postscutellar cavities
M. borneensis Mohamedsaid, 1993—elytra with oblique postscutellar cavities
M. cavidorsis Fairmaire, 1893—elytra with broad postscutellar cavities
M. cavipennis Baly, 1878—elytra with longitudinal cavities at sides
M. c-alba (Jacoby, 1899)—elytra with postscutellar cavities
M. cumingi (Weise, 1910)—elytra with posthumeral area strongly elevated, bearing oval shaped cavities
M. danumica Mohamedsaid, 1993—elytra humpbacked, with deep, elongate postscutellar cavities
M. dimidiata Jacoby, 1886—elytra with elongate cavities in middle
M. discoidalis (Jacoby, 1895)—elytra with longitudinal postscutellar cavities, oblique, closed to suture
M. flavicollis (Gyllenhal, 1808)—vertex strongly depressed
M. foveipennis Medvedev, 2005—elytra with postscutellar cavities
M. gantokensis Kimoto, 2004—elytra with large elongate cavities at sides
M. hemorrhoidalis (Fabricius, 1801)—elytra with cavities in the middle
M. impressipennis Oglobin, 1936—elytra with large postscutellar cavities
M. kerangas Mohamedsaid, 1998—elytra hump-backed, with postscutellar cavities
M. laysi Medvedev, 2002—elytra with shallow longitudinal impression at apex near suture
M. malaysiana Mohamedsaid, 1993—elytra with broad postscutellar cavities
M. marginipennis (Jacoby, 1892)—elytra with deep, elongate postscutellar cavities
M. merah Mohamedsaid, 1993—elytra with spindle-shaped cavities
M. murphyi Mohamedsaid, 2002—elytra with cavities at sides
M. posthumeralis Medvedev, 2005 elytra with large and shallow cavities at sides
M. sargaonica Medvedev, 2005—elytra with postscutellar impression
M. scutellaris Kimoto, 1989—elytra with longitudinally shallow postscutellar depression
M. semifovea Mohamedsaid, 1993—elytra with shallow postscutellar cavities
M. shirozui Kimoto, 1965—elytra with postscutellar cavities
M. tarsata Medvedev, 2005—elytra with postscutellar impression
M. tatemizo Kimoto, 2004—elytra with a pair of large longitudinal cavities at sides
M. tibowensis Mohamedsaid, 2000—elytra hump-backed, with postscutellar cavities
M. trochanterina Mohamedsaid, 1997—metatrochanters with a spine at the base
M. vietnamica Kimoto, 1989—elytra with postscutellar longitudinal cavities near suture
Monoleptocrania foveata (Olivier, 1801)—vertex deeply excavated, with margins protruded
Neolaetana alternans Silfverberg, 1975—elytra with a pair of funnel-shaped cavities at base; antennae 3rd–7th dilated
N. basalis Laboissiere, 1921—elytra with a pair of funnel-shaped cavities at base; pronotum excavated at base; antennae 3rd–7th dilated
N. freyi Silfverberg, 1975—elytra with a pair of funnel-shaped cavities at base; antennae 3rd–7th dilated
N. neavei Laboissiere, 1923—elytra with a pair of funnel-shaped cavities at the base; pronotum excavated at the base; antennae 3rd–7th dilated
N. neumanni (Weise, 1907)—elytra with a pair of funnel-shaped cavities at the base; pronotum excavated at the base; antennae 3rd–7th dilated
N. coeruleipennis Jacoby, 1899—antennae 8th–11th enlarged, longitudinally grooved
N. difformis Jacoby, 1899—antennae 8th–11th enlarged, longitudinally grooved
Nirina flavofasciata Laboissiere, 1940—antennae 6th,7th, 8th, and 9th enlarged
N. regalis Laboissiere, 1940—antennae 8th and 9th enlarged
N. imitans (Jacoby, 1894)—antennae 6th, 7th, 8th, and 9th enlarged
Nirinoides abdominalis Jacoby, 1903—antennae very long, 5th–11th shortened, thickened
N. abyssinica (Jacoby, 1886)—antennae very long, 5th–11th shortened, thickened
N. staudingeri Jacoby, 1903—antennae very long, 5th–11th shortened, thickened
Nymphius buettikeri Medvedev, 1996—abdomen 1st with erect hairs, 2nd with a long appendage, truncated at apex, 3rd with a long appendage, rounded at apex; middle tibiae curved, dilated at apex
N. ensifer (Guillebeau, 1891)—abdomen 3rd with a short appendage, blunt at apex, 4th tuberculate, 5th with a pair of long appendages, pointed at apex
N. forcipifer (Weise, 1900)—abdomen 3rd with a broad appendages, bifurcate at apex, 4th with large transverse corrugations, 5th with a pair of long appendages, curved, dented at apex
N. friedmani Lopatin, 2002—abdomen 1st covered with erect hairs, 2nd with a long appendage, 3rd with a long appendage, pointed at apex
N. gianassoi Bezděk, 2008—abdomen 3rd with a broad, long appendage, tapered and setose at apex, 5th with a pair of long appendages, blunt at apex
N. lydius (Weise, 1886)—abdomen 3rd with a pair of very short appendages, 4th excavated, with a broad appendage
N. ogloblini (Bogachev, 1947)—abdomen 3rd with a pair of long appendages, bent dorsally, blunt at apex, 4th with a pair of long appendages, tapered at apex
N. pravei (Jacobson, 1899)—abdomen 3rd with a long appendage, pointed at apex, 4th with a pair of lateral appendages
N. stylifer (Weise, 1899)—abdomen 3rd with a pair of long appendages, bent ventrally, pointed at apex, 4th with a pair of long appendages
P. barioensis Mohamedsaid, 2001—elytra with postscutellar extrusions deep, obliquely outward posteriorly
P. basipennis Gressitt and Kimoto, 1963—elytra with postscutellar swelling and shallow cavities
P. basituberculata Chen and Jiang, 1984—elytra with postscutellar extrusions straight, bordered at the base by a tubercle
P. caudata Chen and Jiang, 1984—elytra with oval shaped postscutellar depression
P. costata Takizawa and Basu, 1987—elytra with postscutellar extrusions deep, curved, parallel to suture
P. excavata (Chujo, 1938)—elytra with postscutellar extrusions narrow, parallel to suture
P. feae (Weise, 1892)—elytra with postscutellar extrusions shalllow
P. flava Mohamedsaid, 2001—elytra with postscutellar extrusions straight, obliquely outward posteriorly
P. fulvicornis Chen, 1942—elytra with postscutellar extrusions subparallel to suture
P. fusiformis Chen and Jiang, 1984—elytra with postscutellar depression shallow, spindle shaped
P. gongshana Chen and Jiang, 1986—elytra with postscutellar extrusions deep, spindle shaped
P. haemorrhoidalis Medvedev, 2001—elytra with postscutellar extrusions deep spindle shaped
P. humeralis Chen and Jiang, 1984—elytra with postscutelar extrusions obliquely outward, then inward posteriorly
P. insignata Chen and Jiang, 1984—elytra with postscutellar extrusions straight then perpendicularly outward posteriorly
P. jambuica Mohamedsaid, 1996—elytra with postscutellar extrusions deep, obliquely outward posteriorly
P. joliveti Mohamedsaid and Constant, 2007—elytra with postscutellar extrusions subparallel to suture
P. kolthoffli Laboissiere, 1938—elytra with postscutellar extrusions obliquely outward posteriorly
P. kubani Medvedev, 2004—elytra with postscutellar extrusions deep, obliquely outward posteriorly
P. lambirica (Mohamedsaid, 1993)—elytra with postscutellar extrusions deep, obliquely outward posteriorly
P. lamrii Mohamedsaid, 1999—elytra with postscutellar extrusions obliquely outward posteriorly
P. lawa Mohamedsaid, 2001—elytra with postscutellar extrusions deep, obliquely outward posteriorly
P. legenda Mohamedsaid, 1996—elytra carinate at sides, with postscutellar extrusions deep, obliquely outward posteriorly
P. limbangica (Mohamedsaid, 1993)—elytra with postscutellar extrusions shallow, obliquely outward posteriorly
P. lineata Mohamedsaid, 2000—elytra with postscutellar extrusions semicircular shaped
P. lingulata Chen and Jiang, 1984—pronotum with a tongue like projection at the middle of posterior margin
P. liquidambara Gressitt and Kimoto, 1963—elytra with postscutellar suture strongly raised
P. malayana Mohamedsaid, 1996—elytra with postscutellar extrusions broad, deep, spindle shaped
P. marginata Mohamedsaid, 1996—elytra with postscutellar extrusions obliquely outward posteriorly
P. medvedevi Bezděk, 2008—elytra with postscutellar extrusions deep, obliquely outward posteriorly
P. orbiculata Chen and Jiang, 1984—elytra with oval shaped postscutellar depression
P. persimilis Kimoto, 1989—elytra with postscutellar extrusions obliquely outward posteriorly
P. piceipennis Kimoto, 1989—elytral with postscutellar extrusions deep and obliquely outward posteriorly
P. posticata Chen, 1942—elytra with postscutellar suture strongly raised, obliquely outward posteriorly
P. quercicola Chen and Jiang, 1984—elytra with postcutellar extrusions straight, then obliquely outward postseriorly
P. reducta Medvedev, 2001—elytra with postscutellar extrusions small
P. rompinica Mohamedsaid, 1996—elytra with postscutellar extrusions narrow, hook-like
P. rubromarginata Medvedev, 2001—pronotum shallowly grooved in the middle, covered with short hairs; metatibiae emarginate at apex
P. scutellaris Kimoto, 1989—elytra with postscutellar extrusions shallow, parallel to suture
P. subnigra Gressitt and Kimoto, 1963—elytra with postscutellar suture slightly raised
P. tarsalis Mohamedsaid, 1996—elytra with postscutellar extrusions deep, hook like
P. tenasserimensis (Maulik, 1936)—elytra with postscutellar extrusions broad, obliquely outward posteriorly
P. truncata Laboissiere, 1936—elytra with postscutellar extrusions obliquely outward posteriorly
P. verticalis Chen and Jiang, 1984—elytra with postscutellar extrusions long, obliquely outward, then parallel to suture
P. vietnamica Medvedev, 2004—elytra with postscutellar extrusions obliquely outwards posteriorly
P. zakrii Mohamedsaid, 1996—elytra with postscutellar extrusions deep, obliquely posteriorly
Palpaenidea labeonis Laboissiere, 1933—clypeus deeply excavated, with a bifurcated projection hanging from frons; maxillary palpi swollen; antennae 4th with a spine at apex
P. pallipes (Fabricius, 1801)—clypeus deeply excavated, with a pair of sharp projection hanging from frons; maxillary palpi swollen
P. albicans (Jacoby, 1900)—clypeus excavated; maxillary palpi swollen
P. apicalis (Jacoby, 1889)—clypeus excavated; maxillary palpi swollen
P. barbata (Baly, 1879)—clypeus deeply excavated, with tuft of hairs and a small process; labrum extremely large; maxillary palpi swollen
P. carinata Bryant, 1960—clypeus excavated; maxillary palpi swollen
P. clavareaui (Jacoby, 1903)—clypeus excavated; maxillary palpi swollen
P. cocinnea (Jacoby, 1899)—clypeus deeply excavated, with a tooth-like projection in the middle; maxillary palpi swollen
P. coeruleipennis (Baly, 1888)—clypeus deeply excavated; front with tuft of very long hairs hanging on the cavity; labrum extremely large; maxillary palpi swollen
P. crassipalpis (Jacoby, 1892)—clypeus excavated, bituberculate anteriorly; maxillary palpi swollen
P. dilaticornis (Jacoby, 1896)—clypeus openly excavated, with channels at sides; antennae 3rd–11th longitudinally ridged
P. divisa (Jacoby, 1894)—clypeus deeply excavated, with a pointed projection hanging from frons; maxillary palpi swollen
P. ertli (Weise, 1903)—clypeus excavated; maxillary palpi swollen
P. eximia (Baly, 1879)—clypeus deeply excavated, with projection hanging from frons and two coils of hairs from cavity; labrum extremely large
P. facialis (Baly, 1886)—clypeus deeply, broadly excavated; labrum extremely large; antennae 3rd enlarged, dilated at apex
P. fissipes (Laboissiere, 1924)—clypeus excavated; maxillary palpi swollen
P. flava (Laboissiere, 1939)—clypeus excavated; maxillary palpi swollen
P. gracilis (Jacoby, 1889)—clypeus excavated; middle tibiae with elongate appendage at apex
P. hauseri (Weise, 1903)—clypeus excavated; maxillary palpi swollen
P. latifrons (Baly, 1904)—clypeus openly, deeply excavated, with an upright process in the middle; labrum very large
P. longicornis (Jacoby, 1895)—clypeus deeply excavated, with lateral boundaries channelled
P. marginata (Laboissiere, 1920)—clypeus deeply excavated, with a broad projection in the middle
P. modesta (Jacoby, 1896)—clypeus flat, antennae 3rd–8th thickened
P. nasika Maulik, 1936—clypeus deeply excavated, with wedge-shaped structure in middle; labrum very large
P. nasuta (Westwood, 1837)—clypeus deeply excavated, divided by a middle concave structure; labrum very large
P. nigromarginata (Jacoby, 1895)—clypeus deeply excavated, divided by a central ridge
P. pallida (Jacoby, 1896)—clypeus openly excavated; labrum very large; antennae 4th–11th keeled on inner surface
P. patrizii (Laboissiere, 1937)—clypeus deeply excavated, with a narrow projection in the middle
P. pilicornis (Jacoby, 1896)—clypeus deeply excavated, with projection hanging from frons; labrum very large, excavated on each side
P. praetoriae (Gahan, 1892)—clypeus excavated; antennae 1st enlarged
P. rufipennis (Jacoby, 1887)—clypeus deeply excavated, with a projection hanging on frons
P. rufofulva (Jacoby, 1896)—clypeus slightly excavated, with orifice bearing erect hairs; maxillary palpi swollen
P. sabahensis Mohamedsaid, 1997—clypeus deeply excavated, with a broad projection hanging on frons; labrum large; maxillary palpi swollen
P. sumatrensis (Jacoby, 1884)—clypeus deeply excavated, with a narrow projection in the middle; antennae 7th enlarged
P. truncatipennis (Jacoby, 1896)—clypeus deeply excavated, with a projection
P. ugandensis (Laboissiere, 1937)—clypeus deeply excavated; maxillary palpi swollen
P. variabilis (Jacoby, 1886)—clypeus deeply excavated with a broad triangular projection hanging on frons; labrum very large, triangular; maxillary palpi swollen
P. violaceipennis (Jacoby, 1896)—clypeus extensively excavated, with coiled bunch of hairs at sides; labrum very large; maxillary palpi greatly swollen
P. viridis (Hope, 1831)—clypeus openly excavated, with a tuft of hairs; maxillary palpi swollen
Platybrotica misionensis Cabrera and Walsh, 2004—antennae 6th–10th enlarged, excavated ventrally
Platymorpha homoia Blake, 1966—protibiae broadened; mesotibiae with a shallow notched
P. variegata Jacoby, 1888—protibiae strongly dilated at apex
Platyxantha apicicornis Jacoby, 1903—antennae 10th and 11th dilated
P. bicolor Jacoby, 1906—antennae curved, with long hairs
P. borlei Laboissiere, 1931—antennae thickened
P. calabariensis Laboissiere, 1931—antennae slightly curved, with long hairs
P. carinata Weise, 1912—protibiae curved
P. coerulescens (Weise, 1922)—antennae with long hairs
P. citernii Jacoby, 1899—antennae 5th–10th curved
P. conradti Jacoby, 1903—metatibiae with a long process
P. curvicornis Jacoby, 1894—antennae 4th–8th curved; eyes large
P. discoidalis Jacoby, 1895—antennae 4th–8th thickened
P. fascialis Jacoby, 1899—antennae with long hairs
P. fuscitarsis Weise, 1903—antennae 3rd–9th curved
P. kraatzi Weise, 1903—antennae 4th–6th curved
P. livingstoni Jacoby, 1899—antennae with long hairs
P. longicornis Jacoby, 1903—antennae 3rd–8th curved, with long hairs
P. lukunguensis Jacoby, 1899—antennae with long hairs
P. lusingana Bryant, 1958—antennae 3rd–6th curved
P. minor Weise, 1903—antennae 4th, 5th, and 6th curved, with long hairs
P. nigromarginata Jacoby, 1895—antennae 5th–8th curved
P. pallipes Laboissiere, 1940—antennae 4th–8th curved
P. pauli Weise, 1903—antennae 3rd–9th thicker, curved
P. sutteri Laboissiere, 1939—antennae 3rd–10th dilated
P. tenella Weise, 1903—antennae with long hairs
P. trichroa Laboissier, 1931—antennae 3rd–9th thickened; hind tibia with an elongated process at apex
P. versicolor Laboissiere, 1921—antennae 3rd–9th thickened
P. verticalis Laboissiere, 1931—antennae 3rd–10th thickened
Porechontes albiventris (Blake, 1958)—clypeus with a pore in middle, below antennal socket; middle tibiae notched
P. wilcoxi Blake, 1958—clypeus with a pore in the middle, below antennal socket; antennae 3rd compressed, 8th and 9th triangularly dilated; mesotibiae notched
P. limbella (Weise, 1921)—clypeus with a deep pore in the middle, below antennal socket; antennae 9th broadened; mesotibiae notched; eyes large
Prosmidia bispinosa (Fabricius, 1798)—pronotum with oblique ridge at the base on each side of midline; elytra with a pair of tubercles at the base
P. chevrolati (Guerin-Menevile, 1849)—pronotum with an elevation along midline in posterior half; scutellum tongue shaped; elytra with a pair of tubercles at base
P. concinna (Weise, 1905)—pronotum with ridge-like elevation at posterior margin; elytra with a pair of tubercles at the base
P. conifera (Fairmaire, 1882)—pronotum with a sharp, oblique ridge at the base on each side of midline; elytra with a pair of tubercles at the base
P. decempunctata (Laboissiere, 1926)—pronotum with a raised knob at the middle of posterior margin; elytra with a pair of tubercles at the base
P. dregei (Chapuis, 1876)—pronotum with a raised knob at the middle of posterior margin; elytra with a pair of tubercles at the base
P. excavata (Weise, 1909)—pronotum with a kidney shaped cavity; scutellum tongue-shaped; elytra with three tubercles at the base
P. hastata (Laboissiere, 1921)—pronotum with a raised knob at the middle of posterior margin; elytra with a pair of tubercles at the base
P. magna (Weise, 1904)—pronotum with a pair of projections at the middle of posterior margin; elytra with a pair of tubercles at the base
P. passeti (Allard, 1888)—pronotum with a raised knob at the middle of posterior margin; elytra with a pair of tubercles at the base
P. prasina Silfverberg, 1972—elytra with a pair of tubercles at the base
P. sarcedos Silfverberg, 1973—pronotum with a raised knob at middle of posterior margin; elytra with a pair of tubercles at the base
P. semifasciata Silfverberg, 1973—pronotum with an elevation on midline in posterior half; elytra with a pair of tubercles at the base
P. sexplagiata (Jacoby, 1894)—pronotum with a raised knob at the middle of posterior margin; elytra with a pair of tubercles at the base and at apex
P. suahelorum Weise, 1901—pronotum with the posterior margin forms a short, blunt process; scutellum narrow, tongue shaped; elytra with a pair of tubercles at base
P. suturalis (Jacoby, 1908)—pronotum with a raised knob at the middle of posterior margin; elytra with a pair of tubercles at the base
P. vicina (Gahan, 1909)—elytra with a pair of tubercles at the base
P. zavattarii Laboissiere, 1938—pronotum with an elevation along midline in posterior half; elytra with a pair of tubercles at the base
Pseudaenidea limbata Laboissiere, 1938—clypeus deeply excavated, with long hairs
P. monardi Laboissiere, 1939—clypeus deeply excavated, with long hairs
Pseudocophora ambusta (Erichson, 1834)—elytra with deep postscutellar cavities bearing tubercles
P. apicalis Laboissiere, 1932—elytra with deep postscutellar cavities bearing tubercles
P. bicolor Jacoby, 1887—elytra with deep postscutellar cavities bearing tubercles
P. brunnea Baly, 1886—elytra with deep postscutellar cavities bearing tubercles
P. buquetii (Guerin-Meneville, 1830)—elytra with deep postscutellar cavities bearing tubercles
P. carinata Yang, 1991—elytra with deep postscutellar cavities and bordered with sharp edge
P. cochleata Yang, 1991—elytra with deep postscutellar cavities bearing pointed tubercles
P. distincta Baly, 1888—elytra with deep postscutellar cavities bearing tubercles
P. erichsoni Baly, 1888—elytra with deep postscutellar cavities bearing tubercles
P. flaveola Baly, 1888—elytra with deep postscutellar cavities bearing tubercles
P. flavipes Weise, 1913—elytra postscutellar cavities with tubercles
P. javanensis Laboissiere, 1932—elytra postscutellar cavities with tubercles
P. inornata Jacoby, 1893—elytra postscutellar cavities with tubercles
P. madoni Laboissiere, 1940—elytra postscutellar cavities with tubercles
P. monticola Weise, 1913—elytra postscutellar cavities with tubercles
P. nicobarica Jacoby, 1898—elytra postscutellar cavities with tubercles
P. nitens Allard, 1887—elytra postscutellar cavities with tubercles
P. pectoralis Baly, 1888—elytra postscutellar cavities shallow
P. perplexa Baly, 1888—elytra postscutellar cavities with tubercles
P. philippinensis Laboissiere, 1940—elytra postscutellar cavities with tubercles
P. praeusta Allard, 1889—elytra postscutellar cavities with tubercles
P. sumatrana Jacoby, 1896—elytra postscutellar cavities with tubercles
P. uniplagiata Jacoby, 1884—elytra postscutellar cavities with tubercles
P. ventralis Weise, 1913—elytra postscutellar cavities with tubercles
P. wallacei Baly, 1888—elytra postscutellar cavities with tubercles
Pseudocrania basalis Jacoby, 1907—elytra with large, deep, spindle-shaped postscutellar cavities
P. semifulva Bryant, 1953—elytra with narrow extrusions curving obliquely outward posteriorly
Pseudoluperus tuberculatus (Blake, 1942)—abdomen 2nd with a pair of tubercles
S. collarti (Laboissisre, 1932)—antennae 3rd–7th dilated, compressed
S. fastuosa Silfverberg, 1982—antennae 3rd–7th dilated, compressed
S. jeanelli (Laboissiere, 1918)—antennae 3rd–7th dilated, compressed
S. oculata (Laboissiere, 1891)—antennae 3rd–7th dilated, compressed
S. opulenta (Peringuey, 1892)—antennae 3rd–7th dilated, compressed
S. violacea (Allard, 1888)—antennae 3rd–7th dilated, compressed
Sarawakiola ajaib Mohamedsaid, 1997—antennae 1st rounded, extremely large, deformed; vertex deeply, transversely grooved; frontal tubercles larged, with a pair of spine
Scelida balyi Jacoby, 1878—abdomen 2nd with a pair of large appendages
S. flaviceps (Horn, 1893)—abdomen 2nd with a pair of large appendages
S. nigricornis (Jacoby, 1888)—abdomen 1st with a broad, deep depression in the middle; metatibiae curved
Scelidacne andrewi Clark, 1998—abdomen 3rd with trilobed appendage, the lateral densely pubescent, the median glabrous and bifurcate at apex; metatibiae with a curved appendage at apex
Scelolyperus bimarginatus (Blake, 1928)—metatibiae straight, with a thin lamellate lobe at apex
S. curvipes Wilcox, 1965—metatibiae strongly curved, with inner side glabrous
S. hatchi Wilcox, 1965—metatibiae straight, with a flat lobe at apex
S. kroliki Borowiec, 2005—metatibiae curved
S. liriophilus Wilcox, 1965—metatibiae with a lamellate lobe at apex
S. loripes Horn, 1893—metatibiae curved, with inner side pubescent
S. meracus (Saya, 1826)—metatibiae with a lamellate lobe at apex
S. megalurus Wilcox, 1965—metatibiae curved
S. nigrocyaneus (Leconte, 1879)—metatibiae curved, with a ridged lobe at apex
S. nigrovirescens (Fall, 1910)—metatibiae straight, with a thin lamellate lobe at apex
S. ratulus Wilcox, 1965—metatibiae curved
S. schwarzi Horn, 1893—metatibiae strongly curved
S. smaragdinus (Leconte, 1859)—metatibiae curved
S. tejonicus Crotch, 1874—metatibiae strongly curved, with a prominent tooth
Taumacera apicalis (Baly, 1864)—antennae 9th–10th dilated, 11th enlarged, oblong; metasternum with posterior process; metatibiae with a long, pointed process at apex
T. auripennis (Laboissiere, 1933)—antennae 3rd enlarged, globose, 8th reniform
T. azurea (Laboissiere, 1933)—antennae 3rd enlarged, globose, 8th arcuate
T. bella (Weise, 1922)—antennae 8th and 9th enlarged, excavated; clypeus excavated, with a thin acute ridge in the middle
T. bifasciata (Jacoby, 1889)—antennae 9th enlarged, 10th enlarged, excavated, 11th longitudinally excavated; metasternum with posterior process
T. bicornuta (Medvedev, 2001)—clypeus deeply excavated, bordered with conical horn
T. centromaculata Medvedev, 2008—antennae 3rd enlarged, oblong, excavated; metasternum with posterior process
T. constricta Mohamedsaid, 2002—antennae 3rd enlarged, oblong, excavated; metasternum with posterior process
T. costatipennis (Jacoby, 1896)—antennae 3rd elongate-cylindrical; metasternum with posterior process
T. dekatevi Reid, 2001—antennae 3rd enlarged, ovoid, deeply excavated; metatibiae excavated in the middle; metasternum with posterior process
T. deusta Thunberg, 1814—antennae 3rd enlarged, oblong ovate, excavated; metasternum with posterior process
T. dohertyi (Jacoby, 1894)—antennae 5th and 6th enlarged; metasternum with posterior process
T. duri Mohamedsaid, 2000—antennae 3rd enlarged, oblong, with a spine; metasternum with posterior process
T. evi Reid, 1999—antennae 3rd enlarged, oblong, deeply excavated; metasternum with posterior process
T. frontalis Mohamedsaid, 2001—antennae 3rd strongly dilated; clypeus deeply excavated, with a projection; metasternum with posterior process
T. fulvicollis Jacoby, 1881—antennae 3rd enlarged, oblong; metasternum with posterior process
T. indica (Jacoby, 1889)—antennae 3rd broadly triangular, 8th dilated, 9th enlarged; clypeus deeply excavated; metasternum with posterior process
T. insignis (Baly, 1864)—antennae 9th extremely large, globose, 10th enlarged, longer than broad, oblong; metasternum with posterior process; metatibiae with a short process at apex
T. insularis (Gressitt and Kimoto, 1963)—antennae 6th enlarged, 9th and 10th constricted in the middle, oblong; clypeus transversely depressed
T. khalednordini Mohamedsaid, 2010—antennae 1st and 3rd extremely large; metasternum with posterior process
T. kinabaluensis (Mohamedsaid, 1999)—metasternum with posterior process; metatibiae with a very long process at apex
T. laevipennis (Jacoby, 1886)—antennae 3rd enlarged, oblong; metasternum with posterior process
T. maculata (Baly, 1886)—antennae 3rd enlarged, oblong, deeply excavated; metasternum with posterior process
T. magenta (Gressitt and Kimoto, 1963)—antennae 6th and 7th enlarged
T. midtibialis Mohamedsaid, 1998—antennae 3rd enlarged, globular, 4th–11th longitudinally ridged; mesotibiae excavated; metasternum with posterior process; metatibiae with short process at apex
T. mohamedsaidi Reid, 1999—antennae 3rd enlarged, globular, 4th distorted and angularly excavated; clypeus excavated; protibiae excavated in the middle; metasternum with posterior process
T. monstrosa (Jacoby, 1899)—antennae 9th very large, 10th enlarged, excavated, 11th longitudinally excavated; metasternum with posterior process
T. multicostata (Jacoby, 1896)—metasternum with posterior process; metatibiae with short process at apex
T. nagaii Mohamedsaid, 1998—antennae 9th–10th enlarged, oblong; metasternum with posterior process; metatibiae with a short process at apex
T. nigripennis (Jacoby, 1884)—antennae 9th very large, oblong, 10th enlarged, excavated, 11th longitudinally excavated; metasternum with posterior process
T. occipitalis (Laboissiere, 1933)—antennae 3rd enlarged, broadly triangular, 8th and 9th thickened; metasternum with posterior process
T. philippina (Weise, 1913)—antennae 3rd enlarged; metasternum with posterior process
T. rubida (Allard, 1889)—antennae 6th–9th enlarged, dilated at apex; metasternum with posterior process; metatibiae with a short process at apex
T. rubripennis (Duvivier, 1884)—metasternum with posterior process
T. seminigra Reid, 1999—antennae 3rd enlarged, ovoid; metasternum with posterior process
T. smaragdina (Duvivier, 1888)—antennae 9th enlarged, oblong, 10th dilated at apex, triangular; metasternum with posterior process; metatibiae with a short process at apex
T. subapicalis Mohamedsaid, 1993—antennae 3rd enlarged, ovoid, 4th angulate; metasternum with posterior process
T. sucki Weise, 1922—antennae 3rd extremely large, ovoid; metasternum with posterior process; mesotibiae excavated at apical half, dilated at apex
T. sumatrana (Jacoby, 1899)—antennae 9th–10th enlarged, oblong, dilated at apex; metasternum with posterior process; metatibiae with a short process at apex
T. tibialis Mohamedsaid, 1994—antennae 3rd enlarged, kidney shaped, deeply excavated, 4th narrowed, curved; protibiae excavated in the middle; metatibiae with a process at apex; metasternum with posterior process
T. uniformis (Jacoby, 1891)—antennae 3rd enlarged, strongly dilated at apex, medially angulate; metasternum with posterior process.
T. variceps (Laboissiere, 1933)—antennae 3rd enlarged, much longer than broad, 6th and 7th thickened, each with a tubercle; metasternum with posterior process
T. ventralis (Baly, 1864)—antennae 10th and 11th enlarged, deeply excavated; metasternum with posterior process
T. warisan Mohamedsaid, 1998—antennae 3rd enlarged, cylindrical, 4th–11th longitudinally ridged; metasternum with posterior process; metatibiae with a short process
T. yamamotoi (Mohamedsaid, 1998)—antennae 9th–10th very enlarged, triangular, excavated; metasternum with posterior process; metatibiae with a short process at apex
Zinjotella stefaninii (Laboissiere, 1927)—antennae 3rd–7th dilated towards apex; elytra with scutellar area elevated.
Acknowledgments
This is to acknowledge that several images were downloaded from the following websites, which were then cropped and edited: GBIF Deutschland: http://www.biologie.uni-ulm.de/; MCZ, Harvard University: http://mcz-28168.oeb.harvard.edu/mcz/index.htm; Smithsonian Institution Library for the digital Biologia Centrali-Americana: http://www.sil.si.edu/DigitalCollections/bca/bca/explore.cfm. The authors wish to thank Karie Darrow (Department of Entomology, National Museum of Natural History, Washington, DC) and Shahnon Mohamed Salleh (Subang Jaya) for their assistance in enhancing some of the figures. Thanks are also extended to Roslan Yusop, Scientific Officer, Centre for Insect Systematic, Universiti Kebangsaan Malaysia (UKM) for the permission to examine specimens in the UKM Collection, Jan Bezděk, Czech Republic and Shawn Clark, USA for providing reprints of their publications where line drawings are used in the present study.
GillespieJ. J.TallamyD. W.RileyE. G.CognatoA. I.Molecular phylogeny of rootworms and related galerucine beetles (Coleoptera: Chrysomelidae)20083721952222-s2.0-3884919845610.1111/j.1463-6409.2007.00320.xStapelH.MisofH.WagnerT.A Molecular and morphological analysis of Afrotropical Monolepta species and related Galerucinae (Coleoptera: Chrysomelidae)2008661317ScherzX.WagnerT.Revision of Afrocandezea Wagner & Scherz, 2002 from tropical Africa (Coleoptera: Chrysomelidae: Galerucinae)20071174161183LawrenceJ. F.NewtonA. F.PakalukJ.SlipinskiS. A.Families and subfamilies of Coleoptera1995Warszawa, PolandMuseum i Instytut Zoologii PAN7791006SeenoT. N.WilcoxJ. A.Leaf beetle genera (Coleoptera: Chrysomelidae)198211221FurthD. G.SuzukiK.FurthD. G.Character correlation studies of problematic genera of Alticinae in relation to Galerucinae (Coleoptera: Chrysomelidae)Proceedings of the 3rd International Symposium on the Chrysomelidae1994Beijing, JapanBackhuys116135LingafelterS. W.KonstantinovA. S.LeeJ. E.Systena Chevrolat (Coleoptera: Chrysomelidae: Alticinae): notes on nomenclature, redescription of the genus, and a preliminary discussion of characters and phylogenetic relationships199810034674832-s2.0-0031902637KimS. J.KjerK. M.DuckettC. N.Comparison between molecular and morphological-based phylogenies of galerucine/alticine leaf beetles (Coleoptera: Chrysomelidae)200334153642-s2.0-0038703339MayrE.1969New York, NY, USAMcGraw–HillHedlundL.Factors affecting differential mating success in male crested newts, Triturus cristatus1990220133402-s2.0-0025209544AnderssonM.1994Princeton, NJ, USAPrinceton University PressEberhardW. G.1985Cambridge, Mass, USAHarvard University PressEberhardW. G.Postcopulatory sexual selection: darwin's omission and its consequences2009106supplement 110025100322-s2.0-6765033087410.1073/pnas.0901217106EberhardW. G.Copulatory courtship and cryptic female choice in insects19916611312-s2.0-0026079211JolivetP.Hypertelie: mimetisme, signaux sexuels ou moyens de defense. Un dilemma chez les insects: vrai ou faux concept?20076325766MaulikS.On a structure in the antennae of beetles of the chrysomelid genus Agetocera1932102494395610.1111/j.1096-3642.1932.tb01572.xMaulikS.1936London, UKTaylor & FrancisBlakeD. H.A review of some galerucine beetles with excised middle tibiae in the male195810859101BlakeD. H.More new galerucine beetles with excised middle tibiae in the male1966118233266WilcoxJ. A.A synopsis of the North American Galerucinae (Coleoptera: Chrysomelidae)19654001226SilfverbergH.A revision of the genus Prosmidia Weise (Coleoptera, Chrysomelidae)1973139154SilfverbergHA generic revision of the group Prosmidiites (Coleoptera: Chrysomelidae)198213173182FreundW.WagnerT. H.Revision of Bonesioides Laboissière, 1925 (Coleoptera; Chrysomelidae; Galerucinae) from continental Africa20033716191519762-s2.0-004321227810.1080/00222930110096519BarrogaG. F.MohamedsaidM. S.Phylogeny of the genus Aulacophora chevrolat (Coleoptera: Chrysomelidae: Galerucinae) in sundaland20068943383582-s2.0-34547336653ReidC. A. M.Reappraisal of the genus Taumacera Thunberg with descriptions of two new species from South-East Asia (Coleoptera: Chrysomelidae: Galerucinae)1999381192-s2.0-0012760393WagnerT.An introduction to the revision of the afrotropical Monolepta and related taxa (Coleoptera, Chrysomelidae, Galerucinae)19992152152202-s2.0-0033367301WagnerT.Monolepta Chevrolat, 1837, the most speciose galerucine taxon: redescription of the type species Monolepta bioculata (Fabricius, 1781) and key to related genera from (Chrysomelidae, Coleoptera)2007411–4811002-s2.0-3394732158810.1080/00222930601127384BezděkJ.bezdek@mendelu.czNew species and subspecies of Nymphius (Coleoptera: Chrysomelidae: Galerucinae) from Iran and Turkey20084817993MollerA. P.PomiankowskiA.Why have birds got multiple sexual ornaments?19933231671762-s2.0-002787964810.1007/BF00173774JohnstoneR. A.Multiple displays in animal communication:'backup signals' and 'multiple messages'199635113373293382-s2.0-0030049667EberhardW. G.Beetle horn dimorphism: making the best of a bad lot19821193420427HallemE. A.DahanukarA.CarlsonJ. R.Insect odor and taste receptors2006511131352-s2.0-3084443679010.1146/annurev.ento.51.051705.113646ClarkS.The western North American genus Androlyperus Crotch, 1873 (Coleoptera: Chrysomelidae: Galerucinae)1999133-4217222KonstantinovA. S.KorotyaevB. A.Sexual dimorphism and size of aedeagi in apionid weevils (Coleoptera: Apionidae) and flea beetles (Coleoptera: Chrysomelidae): why some masculine males have small aedeagi200410623243382-s2.0-1942540600MohamedsaidM. S.BarrogaG. F.A study on the morphology of antenna and aedeagus in males of Aulacophora Chevrolat (Coleoptera: Chrysomelidae: Galerucinae)201044405414HillP. S. M.Vibration and animal communication: a review2001415113511422-s2.0-33746323079HowseP. E.The significance of the sound produced by the termite Zootermopsis angusticollis (Hagen)1964122-32843002-s2.0-0000476349RocesF.HolldoblerB.Vibrational communication between hitchhikers and foragers in leaf-cutting ants (Atta cephalotes)19953752973022-s2.0-002899535110.1007/s002650050194MohamedsaidM. S.The Galerucinae with sound production devices (Coleoptera: Chrysomelidae)20101606259262SchmittM.ZuniniM.BellesX.BlasM.Stridulatory devices of leaf beetles (Chrysomelidae) and other Coleoptera1992Torino, ItalyAEC263280SchmittM.JolivetP.Stridulation in leaf beetles (Coleoptera, Chrysomelidae)1994Dodrecht, The NetherlandsKluwer Academic319325SchmittM.FurthD.G.The position of the Megalopodinae and Zeugophorinae in a phylogenetic system of the Chrysomeloidea (Insecta: Coleoptera)Proceedings of the 3rd International Symposium on the Chrysomelidae1994Leiden, The NetherlandsBackhuys3844HintonH. E.Secondary sexual characters of Tribolium194214937835005012-s2.0-6544278238SchmitzR. F.FurnissM. M.Secondary sex characters of Scolytus laricis19686116261627FrenchB. W.HammackL.Sexual dimorphism of basitarsi in pest species of Diabrotica and Cerotoma (Coleoptera: Chrysomelidae)2007100159632-s2.0-3384771702410.1603/0013-8746(2007)100[59:SDOBIP]2.0.CO;2JolivetP.What is a Chrysomelid?2001182135146MohamedsaidM. S.JolivetP.Santiago-BlayJ. A.SchmittM.Modified antennae of Malaysian Galerucinae and its taxonomic significance2004The Hague, The NetherlandsSPB Acadamic231247FurthD. G.SuzukiK.BiondiM.DaccordiM.FurthD. G.Studies on Oriental and Australian Alticinae genera, based on the comparative morphology of the metafemoral spring, genitalia, and hind wing venationProceedings of the 4th International Symposium on the Chrysomelidae1998Torino, ItalyAtti Museo Regionale Scienze Naturali91124BövingA. G.Descriptions of larvae of the genera Diabrotica and Phyllobrotica, with a discussion of the taxonomic validity of the subfamilies Galerucinae and Halticinae (Coleoptera: Chrysomelidae)1927299193205BövingA. G.Beetle larvae of the subfamily Galerucine1929752148BövingA. G.CraigheadF. C.An illustrated synopsis of the principal larval forms of the order Coleoptera1931111351LawsonF. A.StehrF.Chrysomelidae19912Dubuque, La, USAKendall-Hunt568585FurthD. G.LeeJ. E.Similarity of the Blepharida-group genera using larval and adult characters (Coleoptera: Chrysomelidae: Alticinae)20001081-226512-s2.0-0034550832FurthD. G.Inter-generic differences in the metafemoral apodeme of flea beetles (Chrysomelidae: Alticinae)19805263271FurthD. G.The metafemoral spring of flea beetles (Chrysomelidae: Alticinae)198271127FurthD. G.JolivetP.PetitpierreE.HsiaoT.The jumping apparatus of Flea Beetles—the metafemoral spring1988chapter 17Dodrecht, The NetherlandsKluwer Academic