Revision of the genus Figura ( Coleoptera : Coccinellidae ) with descriptions of new species

The African genus Figura Ukrainsky, 2006 is revised. A detailed redescription of its morphology is presented. Three new species are described, F. bitalensis sp. n., F. lineata sp. n. and F. ruwenzorica sp. n. All species are illustrated. A key to all species and a map showing the distribution of the genus Figura are provided.


INTRODUCTION
The tribe Epilachnini was established by Mulsant (1846).Formerly it was treated as a separate subfamily Epilachninae within the Coccinellidae, divided into four tribes: Epilachnini, Madaini, Epivertini and Eremochilini.Recently Ślipiński (2007) and Seago et al. (2011) recognized Epilachninae as a tribe within a broadly defined subfamily Coccinellinae.
Epilachnini is a large group of herbivorous ladybird beetles including 25 genera (Jadwiszczak & Węgrzynowicz, 2003;Szawaryn, 2011;Szawaryn & Tomaszewska, 2013;Tomaszewska & Szawaryn, 2013) with a worldwide distribution.Because of its very uniform morphology, current classification of this tribe is unsatisfactory and most of the genera are in need of critical revision.
About 300 species of Epilachnini, grouped in 8 genera, are known from the Afrotropical region.Four of the genera were formerly classified in the tribe Madaini: Figura Ukrainsky, 2006, Megatela Weise, 1906, Merma Weise, 1898and Tropha Weise, 1900.All of them consist of only a few species.The genus Figura was established as Bambusicola by Fürsch (1986) (nec Bambusicola Gould, 1862;Aves) and replaced by Figura Ukrainsky, 2006.Until now the genus Figura contained four species with F. aberratica (Fürsch, 1975) the type species.During a recent examination of the material borrowed from several European museums, three new species were found and are described herein as F. bitalensis sp.n, F. lineata sp.n and F. ruwenzorica sp.n.
Entire beetles, where available, or at least their genitalia and mouth parts were dissected, cleared in 10% solution of KOH, rinsed in distilled water, transferred to glycerol and examined on slides.Illustrations were made from slide preparations using a camera lucida attached to Leica or Carl Zeiss Jenamed microscopes.Mouth parts were cleared in ethyl alcohol and transferred to adhesive tape for SEM photography.After examination the genitalia and mouth parts were transferred to micro-vials and pinned beneath the specimen.Measurements were made using an ocular micrometer attached to a dissecting microscope.The following terms are used: TL -total length, from apical margin of clypeus to apex of elytra; PL -pronotal length, from the middle of anterior margin to the middle of the base; PW -pronotal width at widest part; EL -elytral length along suture including scutellum; EW -elytral width across both elytra at the widest part.Habitus photographs were taken using a digital camera and enhanced using Helicon Focus software.Scanning electron micrographs (SEM) were taken using a HITACHI S-3400N in the Electron Microscopy Laboratory of the MZPW.
The beetle morphology follows Lawrence et al. (2011) including the use of Roman numerals for the abdominal segments; specific terminology used in Coccinellidae follows Ślipiński & Tomaszewska (2010).

Diagnosis.
Figura is an easily distinguishable genus within the African Epilachnini.The tarsal claws of its species have a characteristic shape with the inner teeth touching each other and forming a cordate pattern; a similar shape occurs only in the Neotropical genus Malata Gordon, 1975.Moreover, species of Figura do not possess tibial spurs.Also the structure of male and female genitalia is unique within Epilachnini.
Redescription.Length 2.8-4.1 mm.Body (Figs 106-114) oval, convex; dorsum pubescent.Pronotum yellow to reddish brown or black; elytra light brown, orange to red with black spots or black border, sometimes entirely black with red maculae.
Legs are moderately long and stout, with apices of mid and hind femora protruding slightly beyond outer margin of elytral epipleuron (Fig. 68).Trochanters are simple, on inner margin with shallow cavity for receiving tip of tibia (Figs 2,56,69,99).Femora are cylindrical with a weak groove along their inner sides for receiving tibiae; hind femora in males more swollen (Figs 1,5) than in females (Figs 6,22).Tibiae cylindrical and lacking apical spurs; tarsi pseudotrimerous; tarsal claw bifid, inner teeth touching or almost touching each other forming cordate pattern (Fig. 82).
Body (Figs 1, 106, 107) round-oval, convex; surfaces covered with short, sparse pubescence.Head, mouthparts, antennae, hypomeron, prosternum, mesoventrite, legs and abdomen yellow; metaventrite yellow to dark brown or black.Pronotum yellow to orange.Elytron orange usually with four black spots arranged as 1-2-1.First, basal spot touching elytral margin, not touching suture and scutellum (sometimes reduced); one spot in second row close to elytral suture (sometimes absent), second one near elytral margin (sometimes fused with posterior spot); posterior spot on apical part of elytron; all spots can vary in size.
Prosternum with anterior ridge incomplete medially (ridge not clearly visible), exceeding half of the length of procoxae; prosternal process (Fig. 2) rounded apically, about 1.5 times longer than wide, width about 2.3-2.4 times the length of prosternum in front of coxae at the narrowest point.Mesoventral process about as wide as mesocoxal width at the same position (Fig. 3); mesometaventral connection straight.Epipleuron (Fig. 1) about 2.3-2.5 times broader than metepisternum.
Male terminalia and genitalia.Apical margin of tergite X is truncate (Fig. 11).Viewed from the inside the penis guide is slightly wider medially; about as long as parameres (Figs 14,20).Tegminal strut about as long as tegmen.Viewed from the inside the penis is wider anteriorly and pointed apically (Figs 12,15).
Female genitalia as in Figs 18 and 21.-2.xi.1911-2.xi. , N. Ruwenzori, 6,000-8,500 ft. (1912-193)-193) Note.Paratypes of E. aberratica have a different colour pattern to the holotype and many different morphological characters such as structure of antennomeres, structure of prosternal process, or shape of the male genitalia.More-over, the holotype and paratypes were collected from opposite regions of the range of occurrence for the genus Figura.The specimens designated by Fürsch as paratypes of E. aberratica are therefore described below as a new species F. lineata.
Body (Figs 22,114) round-oval, convex; surfaces covered with short, sparse pubescence.Body yellowishbrown.Pronotum yellow-ochre; elytron yellow-ochre, each elytron with two macule, one large, oval brown in the middle of elytra, near elytral margin, second paler, oval c-shaped, near the first one but closer to suture.
front of coxae at the narrowest point.Mesoventral process about 1.35 times as wide as mesocoxal width at the same position (Fig. 24); meso-metaventral connection straight.Epipleuron (Fig. 22) about 2.1 times broader than metepisternum.
Male not examined.Female genitalia as in Figs 28 and 29.Note: Abdomen of the type specimen is partly destroyed, cut into three pieces.Also genitalia are partly destroyed and not all structures could be studied.Figura bitalensis sp.n. 111) Diagnosis.This species resembles F. ruwenzorica sp.n. and some forms of F. centralis (Fig. 110) in its colour pattern but it can easily be distinguished from them in having a sinuate meso-metaventral connection.It is also distinct from them in that antennomere 3 is about as long as antennomeres 4 and 5 combined whereas in F. ruwenzorica and F. centralis antennomere 3 is shorter than 4 and 5 combined.
Prosternum with anterior ridge complete medially, exceeding half the length of procoxae; prosternal process depressed medially (Fig. 32) and broadly rounded apically, with weak longitudinal carina on apical part, about 1.6 times longer than wide, width about 2.3 times the length of prosternum in front of coxae at the narrowest point.Mesoventral process about 1.1 times as wide as mesocoxal width at the same position (Fig. 33), with dense, deep punctures; meso-metaventral connection sinuate.Epipleuron (Fig. 31) about 2.5 times broader than metepisternum.

Figura centralis
Diagnosis.Specimens with typical colour pattern are easily recognizable by their red colour with large black spot on central part of elytra.Untypical form resembles F. bitalensis sp.n. but is distinguishable from this species in that its meso-metaventral connection is straight.Moreover antennomere 3 is shorter than antennomeres 4 and 5 combined, and antennomere 4 shorter than 5. Similar antenomere structure is present in F. ruwenzorica but F. centralis can be distinguished from this species as it lacks a longitudinal carina on apical part of prosternal process.
Body (Figs 43,109,110) round-oval, convex; surfaces covered with short, sparse pubescence.Head, mouthparts, antennae, hypomeron, prosternum, mesoventrite and legs ferruginous; metaventrite ferruginous or black; abdomen ferruginous sometimes with the median part of ventrite 1 dark brown or black.Pronotum red; elytron red with one large black spot on medially, touching suture; sometimes with one additional black, small spot on anterior part of elytron; sometimes median macula extends to lateral margins of elytra.
Female not examined.Figura lineata sp.n. 108) Diagnosis.F. lineata is easily distinguishable by its dorsal colour pattern, which is unique among Figura species; yellow pronotum with large black macula medially and pattern of yellow and black maculae on elytra as in Fig. 108.This species can also be distinguished from its congeners by the prosternal process creased laterally.
Body (Figs 55, 108) round-oval, convex; surfaces covered with short, sparse pubescence.Mouthparts, antennae, hypomeron, prosternum, scutellum and legs yellow.Epipleurom, meso and metaventrite black.Head black with clypeus and frons yellow.Abdomen with first two ventrites black and the rest yellow.Pronotum yellow with large, black macula on disc touching anterior and posterior margins, elytron with characteristic yellow and black maculae pattern as in Fig. 108.
Prosternum with anterior ridge complete medially, exceeding half the length of procoxae; prosternal process creased laterally, (Fig. 56) rounded apically, about 1.6 times longer than wide, width about 2.5 times the length of prosternum in front of coxae at the narrowest point.Mesoventral process about 1.1 times as wide as mesocoxal width at the same position (Fig. 57); mesometaventral connection straight.Epipleuron (Fig. 55) about 2.3 times broader than metepisternum.
Female not examined.Etymology.The name is derived from the characteristic elytral colour pattern with stripe-like black and yellow maculae.
Figura ruwenzorica sp.n. 113) Diagnosis.F. ruwenzorica is the largest species of the genus Figura.Its almost completely black elytra are characteristic.It is most simillar to F. bitalensis sp.n. in terms of body size and presence of weak longitudinal carina on apical part of prosternal process but is distinct in having a straight meso-metaventral connection.Moreover, antennomere 3 is shorter than antennomeres 4 and 5 combined, whereas in F. bitalensis it is same length.
Body (Figs 68,113) round-oval, convex; surfaces covered with short, sparse pubescence.Head, mouthparts, antennae, hypomera, prosternum, mesoventrite, scutellum, epipleura and legs yellow; metaventrite black; abdomen yellow with median parts of ventrites 1 and 2 dark brown to black.Pronotum yellowish-brown to reddish-brown; elytron black with narrow red lateral border, with two red maculae touching anterior margin, first on callus, second touching suture, apex of elytra red; in some specimens anterior and posterior markings reduced to narrow lines.
Prosternum with anterior ridge complete medially, exceeding half the length of procoxae; median part of prosternal process depressed (Fig. 69), polygonal apically, with weak longitudinal carina apically, about 1.6 times longer than wide, width about 2.1 times the length of prosternum in front of coxae at the narrowest point.Mesoventral process about 1.2 times as wide as mesocoxal width at the same position (Fig. 70), with sparse, small, shallow punctures; meso-metaventral connection straight.Epipleuron (Fig. 68) about 2.5 times broader than metepisternum.
Anterior ridge of prosternum complete medially, very short, not half the length of procoxae; apex of prosternal process (Fig. 98) polygonal, about 1.5 times longer than wide, width about 2.9 times the length of prosternum in front of coxae at the narrowest point.Mesoventral process about 1.1 times as wide as mesocoxal width at the same position (Fig. 99); meso-metaventral connection straight.Epipleuron (Fig. 97) about 1.8 times broader than metepisternum.
Abdominal postcoxal lines not studied because the holotype lacks ventrite 1.
Female not examined.Note: The genitalia of the holotype were partly destroyed; median part of the penis guide cut longitudinally (Fig. 101), tegminal strut not present.

DISCUSSION
Representatives of the genus Figura share some characters with other genera of Epilachnini: Lack of tibial spurs (similar to Afidentula Kapur, 1958, Megatela Weise, 1906, Merma Weise, 1898and Lalokia Szawaryn & Tomaszewska, 2013); bifid tarsal claw with inner teeth touching each other (similar to Malata); lack of apodeme in abdominal segment IX in males (similar to Epiverta Dieke, 1947, Megatela Weise, 1906and Subaffisa Bielawski, 1963).Unique morphological characters that distinguish the genus Figura include structure of labium with scale-like processes on dorsal surface; female genitalia lack sperm duct, spermatheca and accesory gland; coxites fused laterally with paraprocts and male genitalia with tegmen with asymmetrical median lobe.Two membranous sacs in female reproductive system that are attached to the base of the oviduct are regarded as a place for storing sperm instead of the spermatheca (Katakura, 1981).
All species that belong to the genus Figura inhabit the African region of Albertine Rift.In this region the species diversity is high and there is one of the largest numbers of endemics (Plumptrea et al., 2007) and possibly it is one of the top biodiversity hotspots in Africa (Myers et al., 2000).High biodiversity might explain the relatively high species richness within the genus Figura in this relatively small region.All specimens were found at high altitudes between 1,700 and 2,700 m.Several specimens have a label with annotation "Bambous" or "mont.avec bamb.".At this high altitude there are two types of ecosystems, mountain forests (1,500-2,500 m) and bamboo forests (2,500-3,000 m).It is possible that Figura species feed on bamboo leaves.