Genus-group names of Afrotropical flea beetles ( Coleoptera : Chrysomelidae : Alticinae ) : Annotated catalogue and biogeographical notes

This paper consists of an up to date annotated catalogue of the Afrotropical genera of Alticinae (Chrysomelidae), with biogeographical notes on the flea beetle fauna occurring in Sub-Saharan Africa and Madagascar. The following new synonymies are proposed: Eugonotes Jacoby, 1897 (a subgenus of Sanckia Duvivier, 1891) = Brancucciella Medvedev, 1995 syn. n.; Amphimela Chapuis, 1875 = Dibolosoma Jacoby, 1897 syn. n.; Amphimela Chapuis, 1875 = Halticova Fairmaire, 1898 syn. n.; Podagrica Chevrolat, 1837 = Podagrixena Bechyné, 1968 syn. n.; Aphthona Chevrolat, 1837 = Pseudeugonotes Jacoby, 1899 syn. n.; Nisotra Baly, 1864 = Pseudonisotra Bechyné, 1968 syn. n. The following new combinations are proposed: Afrorestia sjostedti (Weise, 1910) comb. n. (from Crepidodera); Bechuana natalensis (Jacoby, 1906) comb. n. (from Ochrosis); Sesquiphaera natalensis (Jacoby, 1906) comb. n. (from Sphaeroderma). The genus Hildenbrandtina Weise, 1910 is trasferred from Galerucinae to Alticinae. New distributional data for many genera 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 & Verma, 2002).The relationship between two of its subfamilies, the monophyletic Alticinae (commonly termed flea beetles) and closely related Galerucinae is an area of active research on Chrysomelidae phylogeny (Duckett et al., 2004;Gómez-Zurita et al., 2007).In this paper, the Alticinae and Galerucinae are considered to be separate subfamilies because of the metafemoral spring in Alticinae and specific structures of the spermatheca, median lobe of aedeagus and hind wing venation (cf.Furth & Suzuki, 1994, 1998).The Alticinae includes 4,000 to 8,000 species grouped in approximately 500 genera that are distributed worldwide, although most occur in the tropical regions of South America, Africa and Asia (Konstantinov & Vandenberg, 1996;Santiago-Blay, 2004).They are highly specialized phytophagous insects, of which the adults and larvae feed on stems, leaves and roots of most of the higher plant families (Konstantinov & Vandenberg, 1996).
Details of the composition of the Afrotropical flea beetle fauna are incomplete, which is highlighted by the discrepancy in the numbers of morphogenera and morphospecies in African entomological material preserved in public and private collections and those that have been officially described.Current scientific literature includes approximately 300 research papers dedicated in whole or part to Sub-Saharan and Madagascan Chrysomelidae Alticinae, including publications on taxonomy, faunistics and ecology.The chronological trend in the numbers of these publications in time is summarized in Fig. 1, begin-ning as early as 1830, although the first significant contribution on the Afrotropical (including Madagascar) flea beetle fauna was by the English coleopterist, Joseph Sugar Baly (1816Baly ( -1890)).Subsequently, in the twenty years following Baly's death (1890Baly's death ( -1910) ) there were three important entomologists working on these beetles: Léon Fairmaire (1820Fairmaire ( -1906)), a French specialist on Coleoptera and Hemiptera; Julius Weise (1884Weise ( -1925)), a German coleopterist that, despite his short life, published a large number of scientific papers, not only on Chrysomelidae but also on Coccinellidae, Curculionoidea, etc.;andMartin Jacoby (1842-1907), a German musician and coleopterist who published 150 articles on leaf beetles after moving to London.
A decrease in the number of publications on Afrotropical flea beetle fauna followed, until the revival in 1930-1940 initiated by the English coleopterist Gilbert Ernest Bryant (1878Bryant ( -1965) ) and the French chrysomelid specialist Victor Laboissière (1875Laboissière ( -1942)).The Alticinae specialists Jan Bechyné (1920Bechyné ( -1973) ) and Gerhard Scherer then published many monographs (see References) on the flea beetle fauna of Sub-Saharan Africa and, to a less extent, Madagascar, describing many new genera and species from 1950-1970.More recently, contributions on the Afrotropical flea beetle fauna were published by Gerhard Scherer, Maurizio Biondi, Paola D'Alessandro, Manfred Döberl and Serge Doguet (see References).

MATERIAL AND METHODS
The catalogue is arranged alphabetically by genus.Names that are in square brackets refer to synonymies, genera incorrectly reported in the Afrotropical region or, in some cases, to Aphthona Chevrolat, 1837 =Pseudeugonotes Jacoby, 1899 syn.n.

Notes.
The three species wrongly attributed to this Neotropical genus were transferred by Bechyné (1959c) to the genera Physomandroya Bechyné and Hemipyxis Chevrolat.

Distribution. Republic of South Africa (Free State and
KwaZulu-Natal).Ecology.The only species in this genus lives in alpine meadows (over 2,000 m a.s.l.) on the Drakensberg and was collected on inflorescences of Asteraceae, such as Senecio and Helichrysum (Biondi & D'Alessandro, 2003).Notes.One species.
Distribution.Afrotropical region (excluding Madagascar) and Arabian Peninsula(?).Ecology.Polyphagous genus associated with several plant families (cf.Jolivet & Hawkeswood, 1995;pers. data).Notes.About one hundred and fifty species in Sub-Saharan Africa.According to Biondi & D'Alessandro (2003), Gabonia is closely related to Luperomorpha Weise, 1887, a genus prevalent and widespread in the Oriental and Australian regions, and is probably a synonym.In fact, many species currently attributed to Gabonia do not show any significant differences from species of Luperomorpha.The diagnostic character reported by Scherer & Boppré (1997) for separating these two genera is the apical spur on the hind tibiae, long and straight in Gabonia and very short in Luperomorpha, but this character is not always reliable.In view of the high variability shown by the genus Gabonia and the need to consider various other genera, synonymy with Luperomorpha may be confirmed in the future by a detailed and careful comparative study of this complicated African genus.
Notes.This genus, with about ten species in Madagascar, is here transferred from Galerucinae to Alticinae because it has a metafemoral spring very similar to that described for the Indo-Malayan genus Mandarella Duvivier (1892: 433) (cf.Furth & Suzuki, 1994).
Notes.About twenty species in Sub-Saharan Africa.

DISCUSSION
Based on this catalogue, of the 101 flea beetle genera, 84 occur in Sub-Saharan Africa (CAF: 64; EAF; 52; SAF: 52; WAF: 40), 37 in Madagascar, 8 in the Mascarene Islands and 5 in the Seychelles Islands (Table 1, Fig. 2).However, these numbers are provisional as information concerning the Afrotropical flea beetle fauna is limited, particularly for Madagascar.This preliminary overview indicates that this fauna is clearly separated from those of other zoogeographical regions (Fig. 6).The geographic distribution of Afrotropical flea beetle genera has distinct Malagasy and Sub-Saharan African components (Fig. 6).
The percentage of genera occurring in both the Afrotropical and another zoogeographical region is 29.0%, with the cosmopolitan component significant and well represented [7.0% of the total (101)] (Fig. 4).Of those shared genera, the Afrotropical region shares the highest percentage with the Indo-Malayan region (31.0%) (Fig. 5).In this case, the presence of some genera in both regions may be due to a possible Gondwanian origin, such as Sanckia, which mainly occurs in Madagascar although species are found in the Sub-Saharan Africa and southern part of the Indo-Malayan region; Torodera, occurs in Sub-Saharan Africa and the Indo-Malayan region and is absent from Madagascar; Amphimela, Nisotra, and Paradibolia, occur in the Afrotropical, Indo-Malayan and the Australian regions; and Bikasha, occurs both on the Seychelles Islands and peninsula of Vietnam.Other genera, such as Hemipyxis, Luperomorpha, Lypnea, Manobia, Philopona and Trachytetra occur not only in the Afrotropical (including Madagascar although infrequently), Indo-Malayan and Australian regions, but also in the eastern part of the Palaearctic region.26.0% of the shared flea beetle genera occur in the Palaearctic region (Fig. 5), including the unique Pan-African flea beetle genus, Angulaphthona, which occurs in Mediterranean Africa, Sub-Saharan Africa and Madagascar (cf.Biondi & D'Alessandro, 2006).A slightly lower percentage of genera (21.0%) occur in both the Afrotropical and Australian regions, although all can also be found throughout the Indo-Malayan region.
As expected, a low percentage of genera occurs in both the Afrotropical and Nearctic regions (10.0%), and in Afrotropical and Neotropical regions (12.0%).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 & 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 the Gondwana.
Other likely Gondwanian elements in the Afrotropical flea beetle fauna are: 1. Zomba, this unique Afrotropical genus belonging to the tribe Monoplatini which mainly occurs in the Neotropical region with a few species in the Nearctic region.The genus Opisthopygme belonging to this tribe of flea beetles is also present in Australia (see above).

Fig. 2 .
Fig. 2. Number of flea beetle genera occuring in the various areas of the Afrotropical region (see Material and methods for abbreviations) (modified from Graf & Cummings, 2007).

Fig. 4 .
Fig. 4. Distribution of Afrotropical flea beetle genera in the different zoogeographical regions (see Material and methods for abbreviations).

Fig. 5 .
Fig. 5. Percentage of flea beetle genera co-occurring in the Afrotropical and another zoogeographical region (see Material and methods for abbreviations).

Fig. 3 .
Fig. 3. Number of flea beetle genera endemic to the various areas of the Afrotropical region (see Material and methods for abbreviations).

Fig. 6 .
Fig. 6.Faunistic similarities of the different areas of the Afrotropical region with those of other zoogeographical regions [Coincidence index and WPGMA clustering method (Weighted Pair Group Method using Arithmetic averaging) (cf.Biondi, 2006)] (see Material and methods for abbreviations). .