Afrotropical flea beetle genera: a key to their identification, updated catalogue and biogeographical analysis (Coleoptera, Chrysomelidae, Galerucinae, Alticini)

Abstract A revision of the Alticini genera from the Afrotropical region is reported. The paper includes the following for the flea beetle fauna occurring in Sub-Saharan Africa and Madagascar: a key to their identification; habitus photos of all the genera; microscope and scanning electron micrographs of many diagnostic morphological characters; and an updated annotated catalogue with biogeographical notes that include new distributional data. The following new synonymies are proposed: Aphthona Chevrolat, 1836 = Ethiopia Scherer, 1972 syn. n.; Sanckia Duvivier, 1891 = Eugonotes Jacoby, 1897 syn. n.; Eurylegna Weise, 1910a = Eurylegniella Scherer, 1972 syn. n.; Kimongona Bechyné, 1959a = Mesocrepis Scherer, 1963 syn. n.; Diphaulacosoma Jacoby, 1892a = Neoderina Bechyné, 1952 syn. n.; Sesquiphaera Bechyné, 1958a = Paropsiderma Bechyné, 1958a syn. n.; Podagrica Chevrolat, 1836 = Podagricina Csiki in Heikertinger and Csiki 1940 syn. n.; Amphimela Chapuis, 1875 = Sphaerophysa Baly, 1876a syn. n. The following new combinations are proposed: Blepharida insignis Brancsik, 1897 = Xanthophysca insignis (Brancsik, 1897) comb. n.; Blepharida multiguttata Duvivier, 1891 = Xanthophysca multiguttata (Duvivier, 1891) comb. n.; Hemipyxis balyana (Csiki in Heikertinger and Csiki 1940) = Pseudadorium balyanum (Csiki in Heikertinger and Csiki, 1940) comb. n.; Hemipyxis brevicornis (Jacoby, 1892a) = Pseudadorium brevicornis (Jacoby, 1892a) comb. n.; Hemipyxis cyanea (Weise, 1910b) = Pseudadorium cyaneum (Weise, 1910b) comb. n.; Hemipyxis gynandromorpha Bechyné, 1958c = Pseudadorium gynandromorphum (Bechyné, 1958c) comb. n.; Hemipyxis latiuscula Bechyné, 1958c = Pseudadorium latiusculum (Bechyné, 1958c) comb. n.; Hemipyxis soror (Weise, 1910b) = Pseudadorium soror (Weise, 1910b) comb. n. The genera Buphonella Jacoby, 1903aand Halticopsis Fairmaire, 1883a are transferred to the tribe Galerucini; the genus Biodontocnema Biondi, 2000 stat. prom. is considered to be valid and reinstated at generic level. Finally, a zoogeographical analysis of the flea beetle fauna in the Afrotropical region is provided.


Introduction
The Chrysomelidae is one of the largest phytophagous insect families and includes approximately 37,000 to 40,000 species (Jolivet and Verma 2002). The monophyletic tribe Alticini is closely related to the tribe Galerucini, both contained within in the subfamily Galerucinae (Bouchard et al. 2011). The relationship between these two tribes, often considered as different subfamilies, is an area of active research regarding Chrysomelidae phylogeny (Duckett et al. 2004;Gómez-Zurita et al. 2007;Ge et al. 2011Ge et al. , 2012. In our paper, Alticini and Galerucini are considered to be separate suprageneric taxa because of the metafemoral spring in Alticini, as well as specific structures of the spermatheca, median lobe of aedeagus and hind wing venation of these two groups (cf. Suzuki 1994, 1998). In our opinion, some of the recently established groupings, based on DNA sequences, still need further in-depth analysis because they are phylogenetically and biogeographically incomplete (cf. Ge et al. 2011Ge et al. , 2012. The Alticini is a tribe composed of minute to medium sized beetles, whose enlarged hind femora and renowned jumping ability have earned them the common name of 'flea beetles'. They are highly specialised phytophagous insects. Both the adult and larval stages feed on stems, leaves or roots, and rarely on flowers, in almost all the higher plant families (Konstantinov and Vandenberg 1996). The tribe Alticini in-cludes 4,000 to 8,000 species, grouped in approximately 500 genera. This taxon has a world-wide distribution, but occurs mainly in the tropical regions of South America, Africa and Asia (Konstantinov and Vandenberg 1996;Santiago-Blay 2004;Biondi and D'Alessandro 2010a).
We recently published an annotated catalogue of the Afrotropical flea beetle genera, based largely on data from the literature . Subsequent to a deeper and more extensive examination of type material, and the study of new Afrotropical flea beetle material preserved in the institutions listed below, it was possible to compile an updated catalogue that contains new synonymies, new combinations, new genera and new distribution records. Even so, many details concerning the composition of the Afrotropical flea beetle fauna remain incomplete . The discrepancy in the number of morphogenera, and morphospecies in particular, preserved in public and private collections of African entomological material, and those that have been officially described, highlights this shortcoming. Current scientific literature includes over 300 research papers dedicated as a whole, or in part, to Sub-Saharan and Madagascan Alticini. These publications include contributions on taxonomy, faunistics and ecology . The chronological trend in the number of publications over time is summarized in Fig. 1, the first appearing as early as 1830. However, the first significant contribution on the Afrotropical (including Madagascar) flea beetle fauna was made by the English coleopterist, Joseph Sugar Baly (1816Baly ( −1890. Subsequently, in the twenty years following Baly 's death (1890−1910), there were three respected entomologists working on this fauna: Léon Fairmaire (1820Fairmaire ( −1906, a French specialist on Coleoptera and Hemiptera; Julius Weise (1844Weise ( −1925, a German coleopterist that, during his life, published a large number of scientific papers, not only on Chrysomelidae but also on Coccinellidae, Curculionoidea and others; and Martin Jacoby (1842Jacoby ( −1907, a German musician and coleopterist, who published 150 articles on leaf beetles after moving to London. A decrease in the number of publications on the Afrotropical flea beetle fauna followed, until a revival in 1930−1940, initiated by the English coleopterist Gilbert Ernest Bryant (1878Bryant ( −1965 and the French chrysomelid specialist Victor Laboissière (1875Laboissière ( −1942. Jan Bechyné (1920Bechyné ( −1973 and Gerhard Scherer (1929-2012, specialists on the Alticinae, then published many monographs (see References) on the flea beetle fauna of Sub-Saharan Africa and, to a lesser extent, Madagascar. They described many new genera and species between 1950 and 1970. More recently, contributions on the Afrotropical flea beetle fauna were published by Gerhard Scherer, Maurizio Biondi, Paola D'Alessandro, Manfred Döberl, Serge Doguet, and Elizabeth Grobbelaar (see References).

Materials and methods
The catalogue is arranged alphabetically by generic names. Names in bold refer to flea beetle genera primarily occurring in the Afrotropical region; those in square brackets refer to: synonymies; genera incorrectly reported in the Afrotropical region; in some cases genera transferred to Galerucini; or genus-group names that are unavailable. The rules of the International Commission on Zoological Nomenclature (1999) are adhered to the Fourth Edition of the International Code of Zoological Nomenclature.
In addition to the author and date of publication, each genus-group name is accompanied by: a) synonymies, exclusively those for the Afrotropical region; b) bibliographic references, including the original description, other important taxonomic contributions, and distribution data; c) type species, including the method of species assignment; d) geographic distribution in the Afrotropical region (cf. Graf and Cummings 2007) and other zoogeographical regions (cf. Sclater 1858); e) ecological remarks, mainly hostplants and/or habitat preferences in the Afrotropical region; f) notes, including the number of Afrotropical species and significant taxonomic information.
Specimens were examined and dissected using WILD MZ12.5 and LEICA M205C binocular microscopes. Photomicrographs were taken using a Leica DFC500 camera and the Auto-Montage Pro 2006 software (license number: 15224*syn2459*153a2112_ maurizio_266836). Scanning electron micrographs were taken using a PHILIPS SEM XL30 CP and HITACHI TM-1000. Morphometric measures were taken using the image analysis software Image-Pro Insight 8.0 (license number: 03080000-5385).
The type material examined during this study is preserved in the following institutions: BAQ: collection of M. Biondi

Key to the identification of afrotropical flea beetle genera
A new key for the identification of the Afrotropical flea beetle genera is proposed. In comparison with the key previously proposed by , our key includes all the known flea beetle genera occurring in Sub-Saharan Africa, Madagascar, Seychelles and the Mascarene Islands. The first key below identifies eight different generic groups, labelled with an uppercase letter (group A, group B etc.). Our generic groups are, on the whole, similar to the seven numbered groups (group 1, group 2 etc.) proposed by . For widespread genera, we have primarily concentrated on the morphological and chromatic variability displayed by those species of the genus in question, known to occur in the Afrotropical region. Pronotum with a distinct but poorly defined median depression near each lateral margin with surface more strongly punctate (Fig. 135)  Pronotum with ante-basal transverse sulcus (Figs 120,121,126,206,209,212,225,246 Pronotum with anterior angles distinctly produced towards anterior and distinctly thickened; posterior angles rounded (Fig. 236). Hind tibiae often curved towards inside (Fig. 237). First metatarsomere wide and subtriangular (Fig. 237). Apical tarsomere of metatarsus moderately swollen (Fig. 237)  Pronotum with anterior angles not thickened, dentiform apically, indistinctly produced forwards anterior; posterior angles dentiform apically (Fig. 199).

Ecology. No information.
Notes. Endemic to Madagascar and comprises about ten known species. The Neotropical genus Gioia Bechyné (1955d: 77) is very similar to Abrarius, and may well be a synonym. Distribution. Republic of South Africa (KwaZulu-Natal, Limpopo, and Mpumalanga) (Fig. 279).

Ecology. Afroaltica subaptera was collected in an open field on Poaceae (also known as Gramineae) (Biondi and D'Alessandro 2007).
Notes. Two species have been described.
Ecology. Some species of this genus have been collected from plants in the family Apiaceae in South Africa (personal data).
Notes. Only one species is known. Distribution. All zoogeographical regions (Fig. 283). Ecology. Polyphagous. This genus has been found associated with herbaceous plants, shrubs and trees belonging to several plant families (cf. Jolivet and Hawkeswood 1995).
Notes. About fifty species are known from Madagascar and Sub-Saharan Africa. Distribution. Afrotropical (including Madagascar), Australian, Eastern Palaearctic and Oriental regions (Fig. 284).

Ecology. No information.
Notes. There are about twenty known species (personal data). Distribution. Afrotropical (including Madagascar), Australian, Nearctic, Oriental and Palaearctic regions (Fig. 288). All the species from the Neotropical region described as Aphthona should be attributed to different genera (cf. Konstantinov and Vandenberg 1996;Konstantinov 1998).

Argopistes
Ecology. Many species of this genus are associated with Oleaceae in Sub-Saharan Africa, especially with Olive trees [Olea europaea var. africana (Mill.)], on which the larvae are leaf miners and adults defoliators (cf. Jolivet and Hawkeswood 1995;personal data).
Notes. The Afrotropical region has four described species.

References
Ecology. No information.
Ecology. No information.
Notes. About ten species are known from the Afrotropical region (personal data). Distribution. Namibia (Fig. 296). Ecology. Biodontocnema brunnea is the only species in this genus, and it is associated with moist habitats (Biondi 2000).
Notes. Four species have been described. Furth and Suzuki (1994: 123, Fig. 10) report a metafemoral spring for this genus, including a drawing, but re-examination of the specimens they examined revealed that they belong to the genus Hespera Weise. Dissection of a variety of Buphonella specimens did not reveal any structure comparable to a metafemoral spring either. We therefore transfer the genus Buphonella Jacoby to the tribe Galerucini. Type species. Chaetocnema rugiceps Baly, 1877: 308 (Madagascar), by subsequent designation by Bechyné (1954b: 683).
Distribution. Central and Southern Africa, and Madagascar (Fig. 298). Ecology. Species of this genus live in moist habitats and are probably associated with plants of the family Cyperaceae (personal data).
Ecology. This genus is mainly associated with plants in the families Chenopodiaceae, Cyperaceae, Juncaceae, Poaceae (also known as Gramineae), and Polygonaceae (cf. Jolivet and Hawkeswood 1995). In the Afrotropical region some Chaetocnema are serious pests of rice (Biondi and D'Alessandro 2008a).
Notes. Over one hundred species are known from Madagascar and Sub-Saharan Africa. Distribution. Namibia and the Republic of South Africa (Fig. 303).
Ecology. This genus is strictly associated with the plant family Proteaceae (mainly Protea spp. and Leucadendron spp.) (Biondi 1998b Distribution. Central African Republic, Democratic Republic of the Congo, Kenya, Nigeria, Republic of South Africa, Sudan, Tanzania and Zimbabwe (Fig. 304).
Ecology. This genus is associated with Poaceae (also known as Gramineae) in moist meadows and forest clearings (Biondi and D'Alessandro 2004).
Notes. There are six known species.

Decaria
Notes. About twenty species have been described.
Ecology. A genus associated mainly with plants in the family Lamiaceae, but also with plants in the Scrophulariceae, Asteraceae and Apicaeae (cf. Jolivet and Hawkeswood 1995).

Ecology. No information.
Notes. Only one species is known. Distribution. Republic of South Africa (Free State and KwaZulu-Natal) (Fig. 311). Ecology. Drakensbergianella rudebecki is the only species in this genus. It lives in alpine meadows (> 2,000 m) on the Drakensberg and was collected on the inflorescences of Senecio and Helichrysum (Asteraceae) (Biondi and D'Alessandro 2003).
Distribution. All zoogeographical regions (Fig. 313). Ecology. The genus Epitrix is mainly associated with plants in the family Solanaceae. Some species can be harmful to plants of economic importance (cf. Jolivet and Hawkeswood 1995).
Notes. About a dozen species are known from Madagascar and Sub-Saharan Africa.
Ecology. Polyphagous. This genus has been associated with several plant families (cf. Jolivet and Hawkeswood 1995;personal data).
Notes. About one hundred and fifty species are known to occur in Sub-Saharan Africa. According to Biondi and D'Alessandro (2003), Gabonia is closely related to Luperomorpha . The latter genus is widespread and prevalent in the Australian and Oriental regions, and is probably a synonym of the former genus. Many species currently attributed to Gabonia do not show any significant differences from Luperomorpha species. The diagnostic character reported by Scherer and Boppré (1997) for separating these two genera is the apical spur on the hind tibia: long and straight in Gabonia; very short in Luperomorpha. However, this character is not always reliable. In view of the wide spectrum of variability displayed by the genus Gabonia, and the need to consider various other genera, synonymy with Luperomorpha may be confirmed by a detailed and careful comparative study of this complicated African genus in the future Doguet 1979).
Ecology. The only species in this genus was collected at a high altitude (3100 m) .
Notes. A single species is known. Distribution. Democratic Republic of the Congo, Guinea, Nigeria and Rwanda (Fig. 319).
Ecology. No information.
Ecology. Polyphagous. This genus has been associated with herbaceous plants and shrubs belonging to many plant families (cf. Jolivet and Hawkeswood 1995).
Notes. About thirty species are known from Sub-Saharan Africa. Six species of Hemipyxis, known from Madagascar, are here transferred to the genus Pseudadorium Fairmaire (see Notes in Pseudadorium).
Ecology. The Afrotropical species of this genus are mainly associated with plants in the families Anacardiaceae and Ericaceae (cf. Jolivet and Hawkeswood 1995;personal data).
Notes. About thirty species are known from Sub-Saharan Africa.

Ecology. No information.
Notes. This genus, with about ten species known to occur in Madagascar, was transferred from the subfamily Galerucinae to the Alticinae (currently the tribe Alticini) by Biondi and D'Alessandro (2010a). The reason being the presence of a metafemoral spring, very similar to that described for the Oriental genus Mandarella Duvivier (1892b: 433) (cf. Furth and Suzuki 1994).
Ecology. A genus that has been associated with a variety of Acacia species, trees in the family Fabaceae (Cox 1997).
Distribution. Mascarene Islands (probably introduced), Oriental region and South-Eastern part of the Palaearctic region (Fig. 325).
Ecology. This genus is associated with plants in the families Verbenaceae and Lamiaceae (cf. Jolivet and Hawkeswood 1995).
Ecology. No information.
Distribution. Democratic Republic of the Congo, Republic of South Africa (Mpumalanga), Rwanda, and Tanzania (Fig. 329).

References.
Distribution. Democratic Republic of the Congo, Equatorial Guinea, Republic of the Congo and Tanzania (Fig. 330).
Ecology. No information.
Ecology. No information. Notes. Four species are known (personal data).
Ecology. This genus is polyphagous and has been associated with several plant families, particularly the Boraginaceae, Asteraceae, Lamiaceae and Scrophulariaceae (cf. Jolivet and Hawkeswood 1995;personal data).
Notes. Over one hundred species known from Madagascar and Sub-Saharan Africa (personal data). Distribution. Cameroon, Democratic Republic of the Congo, Equatorial Guinea, Ethiopia (!) [Oromia region (BAQ)], Nigeria, and Saudi Arabia, Socotra Island (Yemen) and the Australian, Eastern Palaearctic and Oriental regions (Fig. 333).
Notes. Two species have been described for the Afrotropical region: Luperomorpha vittula (Weise, 1915) [described as Jamesonia Jacoby but then transferred to Luperomorpha by Bechyné (1959a)] and Luperomorpha biondii Döberl. Concerning the presence of this genus in the Afrotropical region, we refer to the comments reported for Gabonia Jacoby. Distribution. Afrotropical (including Madagascar), Australian, Eastern Palaearctic, and Oriental regions (Fig. 334).
Notes. About ten species are known from Madagascar and Sub-Saharan Africa . Bechyné (1968Bechyné ( : 1718 considered Escaleriella Weise and Lypnea Baly to be separate genera because of the difference in the shape of their elytral epipleura: expanded in Lypnea, but straight and narrow in Escaleriella. Distribution. Malawi and the Republic of South Africa (Eastern Cape Province, Free State, Gauteng, KwaZulu-Natal, Limpopo, Mpumalanga, and North-West Province) (Fig. 335).
Ecology. This genus is probably polyphagous and has been reported mainly from the following plant families: Epacridaceae, Urticaceae, Cyatheaceae, Asteraceae and Arecaceae (cf. Jolivet and Hawkeswood 1995).
Notes. About fifteen species described from Sub-Saharan Africa.

[Mantura Stephens, 1831]
Not present in the Afrotropical region. Distribution. Madagascar and the Mascarene Islands (Fig. 337). Ecology. The species in this genus are probably polyphagous, and have been associated mainly with plants in the Ericaceae, Asteraceae and Polygonaceae (cf. Jolivet and Hawkeswood 1995).
Notes. Seven species are known from the Mascarene Islands and about ten, as yet undescribed, from Madagascar (personal data).  (Fig. 338) Ecology. The species of this genus generally live at altitudes above 2,500 m in mixed bamboo forests.
Notes. Six species have been described. Distribution. Madagascar and Sub-Saharan Africa (absent in the southern part of SAF) (Fig. 339).
Ecology. Some species of this genus were collected from Spathodea sp. (Bignoniaceae) in Eastern Africa (cf. Jolivet and Hawkeswood 1995).
Ecology. No information.
Ecology. Species in this genus are mainly associated with plants in the family Malvaceae (cf. Jolivet and Hawkeswood 1995).
Notes. About seventy species of this genus are known to occur in Madagascar and Sub-Saharan Africa. Type species. Notomela cyanipennis Jacoby, 1899b: 357 (Cameroon), designation by monotypy.
Notes. Four species have been described.

Ecology. No information.
Notes. A single species has been described.  Jacoby (1906) was first attributed to this Palaearctic genus. However, examination of the type material resulted in the transfer of this species to the genus Bechuana Scherer (Biondi and D'Alessandro 2010a).

Type species. Haltica ruficollis
Ecology. The species in this genus are mainly associated with plants in the family Euphorbiaceae, but also with Leguminosae and Malvaceae (cf. Jolivet and Hawkeswood 1995;Pollard 1957).
Notes. About twenty-five species have been recorded in Sub-Saharan Africa and sixteen from Madagascar.  (Fig. 346).
Ecology. Species in this genus are associated with plants from the family Lamiaceae in South Africa (personal data).
Ecology. Species in this genus are mainly associated with plants in the family Asclepiadaceae (cf. Jolivet and Hawkeswood 1995).
Notes. About thirty-five species have been described in Sub-Saharan Africa. The Madagascan species previously attributed to this genus have been transferred to the genus Pseudophygasia Biondi and D'Alessandro, in press Fabricius, 1787: 78 (Europe), by subsequent designation by Chevrolat (1845: 6).
Notes. About forty species are known from the Arabian Peninsula, Madagascar and Sub-Saharan Africa. Distribution. Democratic Republic of the Congo, Kenya, Republic of South Africa (Mpumalanga and KwaZulu-Natal), Sudan and Tanzania (Fig. 351).
Ecology. The species in this genus are mainly associated with plants in the family Malvaceae (cf. Jolivet and Hawkeswood 1995: 128); some are known to cause damage to cotton crops, Gossypium sp. (Malvaceae).
Distribution. Sub-Saharan Africa (absent in the south-western part of SAF and Madagascar), Saudi Arabia, and Yemen (Fig. 356).
Ecology. This genus is generally associated with plants in the family Anarcadiaceae (cf. Jolivet and Hawkeswood 1995;Chaboo, Grobbelaar and Larsen 2007).
Ecology. The species of this genus live in forests (300-700 m) (Maulik 1931). Notes. There are eight known species.

Pseudophygasia Biondi & D'Alessandro, in press
very close to that of Amphimela Chapuis. The synonymy proposed by  between Pydaristes and Blepharida Chevrolat is unconvincing, because of the absence of dentiform emargination on the hind tibiae in Pydaristes (according to the original description), such emargination is characteristically present in Blepharida.  (Fig. 361).
Ecology. No information.
Ecology. Species in this genus are known to be associated with plants in the family Asteraceae (Biondi and D'Alessandro 2010b).
Notes. Nineteen species have been described.
Distribution. Afrotropical (including Madagascar) and Australian, Oriental, and Palaearctic regions. The species of Sphaeroderma reported from the Neotropical and Nearctic regions should be attributed to different genera (cf. Savini and Furth 2001) (Fig. 366).
Ecology. Species in this genus are mainly associated with plants in the families Asteraceae and Ranunculaceae (cf. Jolivet and Hawkeswood 1995).
Notes. Over fifty species have been recorded from Sub-Saharan African and about ten from Madagascar. Distribution. Republic of South Africa (Western Cape Province) and Tristan da Cunha (Fig. 367).
Ecology. This genus lives in moist habitats generally associated with plans from the family Juncaceae (Biondi 2002b).
Ecology. The only species in this genus was collected from Cotton plants, Gossypium sp. (Malvaceae) (Bryant 1922a).
Notes. Only one species is known. This genus is particularly interesting because it is the only representative of the tribe Monoplatini in the Afrotropical region. This tribe occurs almost exclusively in the Neotropical and southern part of the Nearctic regions. The only exceptions are the genera Zomba and Opisthopygme Blackburn (1896: 41); this last genus is occurring with two species in the Australian region.

Discussion
There are 99 flea beetle genera known from the Afrotropical region. Of these, 83 occur in continental Sub-Saharan Africa, with the different regions each having a varying number of species: CAF: 63; EAF; 55; SAF: 58; and WAF: 44. Furthermore, 39 genera are known from Madagascar, 9 from the Mascarene Islands and Table 1. Occurrence of Alticinae genera of the Chrysomelidae in different areas of the Afrotropical region and other zoogeographical areas (see text for abbreviations) (updated from .

Genus
Our preliminary analysis indicates that this fauna is distinct and can be separated from the faunas of other zoogeographical regions. In Fig. 381, a dendrogram obtained from the cluster analysis [Coincidence index and Weighted Pair Group Method using Arithmetic averaging (WPGMA) (cf. Biondi 2006; Biondi and D'Alessandro 2010a)] performed, using the presence/absence data of flea beetle genera in each of the OGUs (Operational Geographical Unit) considered, is reported. Results reveal a main Sub-Saharan cluster, including the four continental OGUs, these in turn form two subclusters, namely a central-western (WAF-CAF) subcluster and a southern-eastern (EAF-SAF) subcluster. Madagascar (MAD) also forms part of the Sub-Saharan cluster, completing the "Afrotropical group".
The Seychelles (SEY) and the Mascarene islands (MAS) are more closely associated with the other zoogeographical regions [(NAR-NTR)(PAR-ORR)AUR)] because of the occurrence of a high percentage of widespread genera that characterize the flea beetle fauna of these two archipeligoes. Moreover, faunistic similarity based on the widespread flea beetle genera also clusters the other zoogeographical regions together in two distinct groups, namely the Palaearctic-Oriental-Australian regions [(PAR-ORR) AUR] and Nearctic-Neotropical regions (NAR-NTR).
The geographic distribution of Afrotropical flea beetle genera is therefore well characterized and it has distinct Malagasy and Sub-Saharan African components (Fig. 381).
The percentage of genera occurring in both the Afrotropical and another zoogeographical region is 32.0%, with the cosmopolitan component significant and well represented [8.0% of the total of 99] (Fig. 379). The Afrotropical region shares the highest percentage of genera with the Oriental (27.0%) and Palaearctic (27.0%) regions (Fig. 380). The copresence of genera in both the Afrotropical and Oriental regions, often called Palaeotropical distribution, may be due to a possible Gondwanian origin., Sanckia is such an example, although the genus occurs mainly in Madagascar, species are also found in Sub-Saharan Africa and the southern part of the Oriental region. Other examples are the genera Argopistoides and Jacobyana which occur in Sub-Saharan Africa and the Oriental region and are absent from Madagascar; and Amphimela, Chabria, Nisotra, and Paradibolia, which occur

Genus WAF CAF EAF SAF MAD SEY MAS PAR NAR NTR ORR AUR
in the Afrotropical, Oriental and the Australian regions. Other genera, such as Bikasha, Hemipyxis, Luperomorpha, Lypnea, Manobia, Philopona and Trachytetra occur not only in the Afrotropical (including Madagascar, although infrequently), Oriental and, generally Australian regions, but also in the eastern part of the Palaearctic region. More specifically, the Palaeartic region shares 27 flea beetle genera with the Afrotropical region (Fig. 380), including the unique Pan-African flea beetle genus, Angulaphthona, which occurs in Mediterranean Africa, Sub-Saharan Africa and Madagascar (cf. Biondi and D'Alessandro 2006). A significantly lower percentage of genera (20.0%) occur in both the Afrotropical and Australian regions, although all of these can also be found throughout the Oriental region. As expected, the lowest percentage of genera occur in both the Afrotropical/Nearctic regions (10.0%) and Afrotropical/Neotropical regions (10.0%). All genera common to the Afrotropical, Nearctic and Neotropical regions are also found in all other zoogeographical regions with the exception of the genus Terpnochlorus, which only occurs in the Afrotropical region, Venezuela and Mexico (cf. Furth and Suzuki 1994). Moreover, the possible synonymy between the genera Abrarius from Madagascar and Gioia from South America (see above), if confirmed, could indicate an interesting zoogeographical connection among the ancient regions of Gondwana (Biondi and D'Alessandro 2010a).
As reported in Biondi and D'Alessandro (2010a), other likely Gondwanian elements in the Afrotropical flea beetle fauna are: The unique Afrotropical genus Zomba belonging to the tribe Monoplatini, which mainly occurs in the Neotropical region with a few species found in the Nearctic region. The genus Opisthopygme, also from the Monoplatini, is also present in Australia (see above).
There are 39 flea beetle genera known from Madagascar, 13 of which are endemic. One of them, Metroserrapha Bechyné, also occurs in the Mascarene Islands 2012, in press). Some of these Madagascan genera, such as Neodera, Physomandroya, Pseudadorium, Pseudophygasia, and Xanthophysca, show clear African affinities, but Antanemora, Ntaolaltica, and Metroserrapha are more closely related to Oriental genera (Biondi and D'Alessandro 2012, in press). The remaining endemic Malagasy genera, Anaxerta, Diphaulacosoma, Halticotropis, and Hildebrandtina, are probably all very ancient. Establishing their affinities with certainty, whether African or Oriental, is very difficult using only a comparative morphological approach.