Natural history of the flea beetle genus Arrhenocoela Foudras (Coleoptera, Chrysomelidae, Alticinae)

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1 Italian Journal of Zoology ISSN: (Print) (Online) Journal homepage: Natural history of the flea beetle genus Arrhenocoela Foudras (Coleoptera, Chrysomelidae, Alticinae) Maurizio Biondi & Giuseppina de Nardis To cite this article: Maurizio Biondi & Giuseppina de Nardis (2002) Natural history of the flea beetle genus Arrhenocoela Foudras (Coleoptera, Chrysomelidae, Alticinae), Italian Journal of Zoology, 69:1, 83-93, DOI: / To link to this article: Copyright Taylor and Francis Group, LLC Published online: 28 Jan Submit your article to this journal Article views: 107 View related articles Full Terms & Conditions of access and use can be found at Download by: [ ] Date: 17 November 2017, At: 20:46

2 ({$% Ital.J. Zool., 6ft (2002) Downloaded by [ ] at 20:46 17 November 2017 Natural history of the flea beetle genus Arrhenocoela Foudras (Coleoptera, Chrysomelidae, Alticinae) MAURIZIO BIONDI GIUSEPPINA DE NARDIS Dipartimento di Scienze Ambientali, University of L'Aquila, via Vetoio, Coppito (AQ) (Italy) ABSTRACT Some taxonomical, biological and ecological notes for Arrhenocoela lineata are reported, and the systematic position of the genus Arrhenocoela is discussed. The complete descriptions of the egg and the first-instar larva are also given. Line drawings and scanning electron micrographs of particular morphological aspects for both the adult and the larva are also provided. INTRODUCTION The genus Arrhenocoela was described by Foudras (I860), who separated it from the genus Crepidodera Chevrolat, 1836 (sensu Chevrolat, 1836) "par la disposition du corselet qui n'offre pas de depression transversale, mais en sillon profond et limit e de chaque cote par une fossette qui n'atteint pas le bord posterieur... La forme du dernier segment de l'abdomen du male est tres-remarquable ainsi que celle de l'hemicycle et de l'edeage". This flea beetle genus includes the only species A. lineata (Rossi, 1790) occurring mainly in the coastal areas of the Mediterranean Basin. Our study regards many aspects of this genus, such as its systematic position, new ecological and biological information and a complete description of the egg and the first-instar larva. MATERIALS AND METHODS Adult specimens of A. lineata living on Erica arborea (Ericaceae) were collected in September 1998 in Lido di Ostia (Rome), 0 m a.s.l. (Latium, Italy). For the rearing, preservation, and slide preparation procedures used in this study we refer to LeSage (1984) and Biondi & de Nardis (1998). The material was preserved in 70% ethyl alcohol. Adults and larvae were dissected using a stereoscopic microscope. For a detailed morphological study of the anatomically minute structures, the larvae were placed in Faure liquid, mounted on slides, and observed with a transmitted-light microscope (Leitz, Ortholux 2: obj. 10, 25 / 1.25) or, after preparation through 'critical point drier' and 'sputtering technique', analysed with the scanning electronic microscope (SEM, Philips XL30 CP). Abbreviations used: ab, abdominal segment (I to X); am, articulating membrane; ch, chorion; cl, clypeus; els, clypeolabral suture; Cui a, first subbranch of the cubitus vein; cu la -cu lb, crossvein between Cu la and Cu Ib ; Cu lb, second subbranch of the cubitus vein; lcuc, first cubital cell; ex, coxa; ds, digitiform seta; eb, eggburster; ech, epichorion; eph, epipharynx; fe, femur; fls, filiform seta; fr, frons; ga, galea; hy, hypopharynx; i, insertion of seta; lc, lacinia; leps, long capitate setae; Ig, ligula; lb, long sensillum basiconicum; lp, labial palpus; lr, labrum; M3, third branch of the media vein; mnd, mandible; msx, mesothorax; mtb, microtubercle; mtx, metathorax; mxp, maxillary palpus; oc, ocellus; Pcu, post-cubitus vein; pda, peduncolate seta; pe, peritreme; pf, palpifer; pic, placoid sensillum; pm, prementum; po, pygopodium; psm, postmentum; pv, pulvillus; px, prothorax; py, pygidium; r-m, crossvein between medial and radial veins; sa, sensory appendage; sb, short sensillum basiconicum; scps, short capitate seta; spo, spiracular opening; ss, styloconic sensillum; sst, sensory structure; st, stipes; ti, tibia; tr, trocanter; trn, trocantin; ts, tarsungulus. KEY WORDS: Coleoptera Chrysomelidae - Arrhenocoela lineata - Larval morphology - Biology - Taxonomy. AKNOWLEDGEMENTS We thank Dr. Maria Giammatteo for her technical assistance with the scanning electron microscope (SEM). This research was supported by a grant from "Ministero dell'universita e della Ricerca Tecnologica e Scientifica" (CLUSTER-11). (Received 2 July Accepted 9 October 2001) TAXONOMIC ACCOUNTS Arrhenocoela lineata (Rossi, 1790) (Figs 1-43) Morphological description of the egg (Figs 19-21) Size: length = 1.12 ± 0.18 mm; width = 0.60 ± 0.05 mm (n = 40). Shape: subcylindrical. Colour: orange yel-

3 Downloaded by [ ] at 20:46 17 November Figs Arrhenocoela lineata, adult. 1 - Head, dorsal view (73x). 2 - Ditto, ventral view (67x). 3 - Pronotum (66x). 4 - Scutellum (478x). 5 - Prosternum (64x). 6 - Mesonotum (60x). 7 - Mesosternum (53x). 8 - Pygidium O" (73x). low when laid; later, brown yellow, or reddish yellow. Surface: epichorion rugose, densely covered with small protuberances; chorion translucid, without evident reticulation. Head (Figs 23-27, 42E-I) M. BIONDI, G. DE NARDIS Hypognathous, subglobose, strongly sclerified (Fig. 23). Cephalic sutures complete and clearly distinct; endocarina straight, black, posteriorly fused with the coronal suture; coronal suture short and thin along the epicranial suture inner margin. Frontal sutures Y-shaped, pale. Epistomal suture well sclerified forming with the endocarina a unique blackish T-shaped suture. Antennae (Fig. 24) one-segmented, supported by a well-developed translucid articulation membrane. Antennomere cylindrical, with strongly sclerified annular ring with five sensorial pores (two dorsal and three ventral); upper side membranose, with a big conical sensory papilla basally slightly raised, with nine basiconic sensilla different in shape and size; three long basiconic sensilla (cf. Ritcey & Mclver, 1990; Bartlet et al., 1999) of which two longer, more thin, distally pointed, slightly curved; one peg-like sensilla, finger-shaped; six short basiconic sensilla of which three longer, cone-shaped, raised from bulbous base, and three very small, dentiform. Morphological description of the first-instar larva The larva of A. lineata was first described by Perris (1873, 1876). For those times, the description supplied by the French author, based on mature larvae ["Long. 6 mill." (Perris, 1876: 198)], can be considered a good one. However, it is scarcely useful because practically lacking in precise references to the chaetotaxy and the distribution of tubercles. We therefore believe it necessary to report a complete and up-to-date re-description of the first-instar larva of this species. Size: body length = mm; pronotal width = mm (n = 30). Habitus: subcylindrical, thickset, gradually narrowed from the head to the pygidium (Fig. 22). Colour: body lemon yellow with blackish cephalic capsule, pronotum, tubercles, legs and pygidium. Distribution and nomenclature of the tubercles is reported in Table II. Figs Arrhenocoela lineata, adult. 9 - Front leg d" (35x) Tarsal claw of the front leg (282x) Metafemoral spring, anterior and posterior view (151x) Dorsal view of the median lobe of the aedeagus (54x) Ditto, ventral view of the distal part (203x) Ditto, dorsal view of the distal part (200x) Ditto, lateral view (54x).

4 THE GENUS ARRHENOCOELA 85 Downloaded by [ ] at 20:46 17 November 2017 Fig Arrhenocoela Hneata. A, spermatheca, dorsal view. B, ditto, inner lateral view. C, ovopositor styli, ventral view. D, ditto, dorsal view. D Epicranium posteriorly with four pairs of minute trichoid sensilla and two pairs of sensorial pores; dorsally with four pairs of filiform setae, one pair of small setae, two pairs of sensorial pores and one pair of cupuliform ocelli in postero-lateral position to the antennae; lateroventrally with four pairs of filiform setae and one pair of sensorial pores; posterior epicranial margin arcuate. Frons wide, subpentagonal with six filiform setae, of which the central ones much shorter and thin, and two sensorial pores in proximal position. Clypeus (Fig. 42H) transverse, trapezoidal, laterally rounded especially at the basal half. Anteclypeus sclerified, medially weakly incised, with eight trichoid sensilla; postclypeus membranose with distal margin straight. Labrum (Fig. 42H) strongly sclerified, subtriangular, with four filiform setae and two sensorial pores; distal margin medially weakly incised. Epipharynx (Fig. 421) rectangular, in the middle densely covered by microtricha (styloconic sensilla sensu Bartlet et al., 1999); dorsal side laterally with five pairs of robust pedunculate setae, weakly uncinate; distal margin dorsally with two pairs of filiform setae; ventral membranose area with eight groups of campaniform sensilla, symmetrically arranged laterally to the ventral sagittal axis. Mandibles (Fig 42F, G) strongly sclerified, reddish brown, ventrally concave and dorsally with two setae and two sensorial pores, five-dentate; tooth I reduced, very short; tooth III serrate; tooth V shorter than IV. Mola not present. No ialine process along the inner basal margin. Stipes subcylindrical, thickset, partially strongly sclerified, with one pair of sensorial pores and six pairs of setae different in shape and in size. Maxillary articulation membrane translucid. Cardo triangular, strongly sclerified with two lateral filiform setae. Palpiferi ellyptical, with two long setae. Maxillary palpi (Figs 26-27, 42E) three-segmented; segments 1-2 subcylindrical, segment 3 elongate, stumped, with the apical part membranose, with ten small cone-shaped sensorial structures: nine arranged in a circle and one central (similar to the short basiconic sensilla III type of Ritcey & Mclver, 1990, but not arising from a raised bulbous base). Basal segment with two sensorial pores; middle segment with one sensorial pore and two setae; apical segment with one large placoid sensillum, one lateral external long basiconic sensillum and one lateral inner small seta. Galea (Fig. 42E) lobate, ventrally supported by a subrectangucula-cuib Fig Arrhenocoela Hneata. A, hind wing. B-D, spiculum ventrale with 8 th abdominal segment, dorsal, lateral and.ventral view.

5 Downloaded by [ ] at 20:46 17 November M. BIONDI, G. DE NARDIS Figs Arrhenocoela lineata. Egg: 19 - Egg with the exit hole (h) of the new-born larva ditto, detail Epichorion. First-instar larva: 22 - Habitus, lateral view Head, lateral view Left antenna, dorsal view Ocellus Maxillae and labium, ventro-lateral view.

6 THE GENUS ARRHENOCOELA 87 TABLE I - Morphological differences between the genera Arrhenocoela and Altica. Genus Arrhenocoela Genus Altica ADULT - Procoxal cavities closed (Fig. 5) - Hind wing venation with an evident Cu la, linked to the Cuj h by a short crossvein (cu la -cu lb ) (Galerucinae type) (Fig. 18A) - The Pcu vein directly associated with the Cu lb to form the lcuc; - Pcu disappears after associating with the Cu lb (Fig. 18A) - Elytral punctation arranged in regular rows - Metafemoral spring attributable to the Blepharida Morpho-Group (Figs 11-12) (cf. Furth, 1985) ADULT - Procoxal cavities partially open. Hind wing venation without Cu la (Alticinae type) - The Pcu vein completely fused with the Culb to become indistinguishable from the latter - Elytral punctation confused - Metafemoral spring attributable to the Altica Morpho-Group (cf. Furth, 1985) Downloaded by [ ] at 20:46 17 November 2017 FlRST-INSTAR LARVA - Antennae 1-segmented (Fig. 24) - Tubercles Dai e Dpi separated (cf. Table II) - Tubercles Dm of the abdomen present (cf. Table II) - Tubercles Dpe with 1 long seta (cf. Table II) - Annuliform tubercles with accessorial rooms (Figs 41C-E) lar sclerite with seven setae along the apical margin and two basiconic sensilla, peg-shaped, in the middle area. Lacinia membranose, with robust setae gathered behind the galea. Prementum membranose, supported by a narrow semicircular sclerite bearing two filiform setae and two short basiconic sensilla. Ligula narrow and convex, with five pairs of small setae. Labial palpi twosegmented; segment 1 subcylindrical with one sensorial pore; segments 2 with one placoid sensillum, one long basiconic sensillum and six-seven apical cone-shaped sensorial structures similarly to segment 3 of maxillary palpi. Postmentum short, sclerified with four filiform setae and four sensorial pores. Hypopharynx with rugose surface, covered with sensorial laminate structures similar to an indented fan. Thorax (Figs 28-34, 41C, 41F, 41H, 42A-D) FlRST-INSTAR LARVA - Antennae 2-segmented - Tubercles Dai e Dpi united - Tubercles Dm of the abdomen absent - Tubercles Dpe with 2 long setae - Annuliform spiracles simple Segments subcylindrical. Prothorax slightly larger. Pronotum ellyptical with nine pairs of setae different in length, seven pairs of minute trichoid sensilla and twothree pairs of sensorial pores. Median ecdysial line pale dividing the pronotal plate into two symmetrical parts, each with two groups of setae, one anterior constituted by six filiform setae apically pointed and another by three long club-shaped setae. Mesonotum and metanotum with similar distribution to tubercles. Ruptor ovi blackish, dentiform, in central position near the capitate seta of tubercle Dlpi (Fig. 30) (cf. Table II). Mesothoracic spiracle annuliform with darker circular peritreme (Figs 29, 41C). Legs (Figs 28, 42A-D) robust, moderately elongate. Metathoracic legs larger and longer. Legs partially membranose, six-segmented with trochantin, coxa, trochanter, femur, tibia, tarsungulus and pulvillus. Coxa circular, normally with four robust filiform setae distinctly pointed and ten minute spiniform sensilla. Trochanter strongly sclerified, narrow, subcircular, with seven-eight sensorial pores; membranose area between trochanter and femur with four robust filiform setae of which the ventral one very long and pointed. Femur strongly sclerified with five setae of which four along the distal margin; membranose ventral area between femur and tibia with two long and robust pointed setae; tibia longer than femur, with five setae and one sensorial pore in dorsal-apical position. Tarsungulus (Fig. 32) strongly curved and uncinate, with one long and weakly arcuate basiconic sensillum. Abdomen (Figs 35-40, 41D-E, 41G, 43A-C) Segments I-VII similar in chaetotaxy and tubercle disposition but different in size; dorsally with two main rows of tubercles Dm (cf. Table II). Intersegmental articulation membrane constituted by many transverse and little deep folds and densely covered by subtriangular and imbricate microtubercles, apically posteriorly bent. Segment VIII with posterior dorsal tubercles fused along a median line (cf. Table II); spiracles constituted by an atrium with circular transversal section generally surrounded by nine accessorial chambers ('air tubes', cf. Booth et al., 1990) regularly distributed along the perimeter (annular multiforus spiracle) (Fig 41D, E). Pygidium (Fig. 43A) elliptical, well sclerified in the dorsal central area; dorsal setae capitate of which eight very long, placed along the external margin, and two proximal very short; ventral side more membranose with a subrectangular tubercle supplied with four long filiform setae and two minute trichoid sensilla. Pygopodium (Fig. 43B) cylindrical, partially membranose, with a ventral sclerified area with two filiform setae, six microsetae and four trichoid sensilla.

7 Downloaded by [ ] at 20:46 17 November M. BIONDI, G. DE NARDIS Figs Arrhenocoela lineata, first-instar larva Maxillary and labial palpi, detail ventro-lateral view, legs of different size Mesothoracic spiracle Left egg-burster, dorsal view Right metathoracic leg, front view Metathoracic tarsungulus, front view tarsungulus and pulvillus of the metathoracic leg, lateral view Pulvillus of the metathoracic leg, posterior view.

8 Downloaded by [ ] at 20:46 17 November 2017 THE GENUS ARRHENOCOEIA 89 Dm Figs Arrhenocoela lineata, first-instar larva Terminal part of the abdomen, lateral view Abdominal segment V, setae on the tubercles Dai and Dm, lateral view Abdominal segment VI, capitate seta on the tubercle Dae Abdominal segment V, capitate seta in transversal section Abdominal segment VI, digitiform seta on the tubercle Dm Abdominal segments VII, VIII and : IX (pygidium), dorsal view (for abbreviations of the tubercles see Table II). DISCUSSION Particularly interesting is the shape of the larval dorsal setae in A. lineata obtained with SEM. They show, from the thorax to the pygidium, a gradual increase of the capitate shape with distal part distinctly bulbous (Figs 35-38). Similar setae were described by Boving (1929) for some species of Galerucinae. This author distinguished 'clavate' and 'capitate' setae according to a narrower or larger apical part. However, the morphology of most of the dorsal setae in A. lineata is very particular. They show a cylindrical shape in almost all their length, while the apical part is similar to a small membranose bulb, translucid and weakly sclerified. In tran-

9 90 M. BIONDI, G. DE NARDIS Downloaded by [ ] at 20:46 17 November 2017 TABLE II - Distribution of the tubercles in the first-instar larva A. lineata. Regions of the body: 1, prothorax; 2, mesothorax; 3, metathorax; I- X, abdominal segments; DR, dorsal region; DLR, dorso-lateral region; EPR, epipleural region; PR, pleural region; SR, sternal region. Tubercle nomenclature and abbreviations: D-DL-EPa, dorsal, dorso-lateral and epipleural anterior fused; Da, dorsal anterior (Dai fused on mesothorax); Dae, dorsal antero-exterior; Dai, dorsal antero-interior; DLae, dorso-lateral antero-exterior; DLe, dorso-lateral exterior; DLp, dorso-lateral posterior; DLp-S, dorso-lateral posterior and spiracular fused; Dm, dorsal median (scutal tubercle); Dp, dorsal posterior (dorsal postero-interior and dorsal posteroexterior fused on VIII); Dpi, dorsal postero-interior; Dpe, dorsal postero-exterior; Dpe-DLpi, dorsal postero-exterior and dorso-lateral postero-interior fused); EP, epipleural; EPa, epipleural anterior; EPp, epipleural posterior; ES, eusternal; ES-SS, eusternal and sternellar fused; P, pleural; PST, presternal, S, spiracular; SS, sternellar; T, trocantin. sversal section, these setae show centrally a longitudinal channel running from the base to the bulb. Probably, the bulb contains urticating liquid; the surface of the bulb seems smooth and without pores. The proximal setae of the pronotal plate and the setae on the tubercles Dm are more properly clavate-digitiform. of The SEM morphological study also allowed us to clarify the morphology of the pulvillus and the tarsungulus (Figs 32-34). The pulvillus is constituted by a flexible, wide and pleated membrane, inserted between the posterior side of the tarsungulus and the tibia; in dorsal view, it appears as a hemispheric hood protecting the tarsungulus below. The pulvillus is asymmetrically inserted on the distal part of the legs. Therefore, the tarsungulus is anteriorly completely uncovered consenting the larva to catch at the plant without impediments from the protection membrane. This species generally lives in the coastal areas of the Mediterranean Basin (cf. Gruev & Doberl, 1997); sometimes it can penetrate inner regions following the maquis vegetation. It is associated with Ericaceae; the adults mate from September to November; the larval development is in spring; the pupation is in the soil. Females lay their eggs on plants of Erica arborea and E. scoparia, mainly in the bifurcations of young terminal branches or on the leaves. The eggs, always singly laid, are almost completely covered by vegetal fragments assuming a cryptic aspect. At the beginning, the coloration of the egg is orange yellow then, when it becomes ripe, reddish brown. The larva comes out from the egg through the part of the chorion in contact with the plant, using a small hole made with the mandibles in sub-apical position. In laboratory, the incubation period is very long, about 2.5 months. Very probably this species spends the winter as an egg. The first-instar larva shows an aposematic lemon yellow coloration, with cephalic capsule, pygidium and tubercles smooth black, which contrasts strongly with the green leaves, the reddish brown branches, and the white {E. arhored) or greenish flowers (E. scoparia) of Erica. From laboratory observations, it emerged that the larva penetrates into flowers through a small circular hole made in the corolla and feeds on covering tissue of the ovary and the immature ovules (Fig 41A, B). No first-instar larva was observed to feed on foliar parenchyma or floral corolla. The systematic position of the genus Arrhenocoela Some authors have considered Arrhenocoela closely related to Neocrepidodera Heikertinger, 1911 [= Asiorestia Jacobson, 1925 (cf. Konstantinov & Vandeberg, 1996); = Crepidodera Chevrolat, 1836 sensu Foudras, I860], mainly for its having an evident pronotal transverse antebasal sulcus laterally delimited and the elytral punctation arranged in regular rows. Heikertinger (1950) also included Arrhenocoela within his "Crep/rforfera-Verwandschaft", taxonomical group comprehending some Holarctic and Oriental genera characterised by: antennae 11-segmented; pronotum normally with a transverse antebasal sulcus laterally delimited; elytral punctation arranged in regular rows; procoxal cavities generally posteriorly closed; tibiae unmodified.

10 THE GENUS ARRHENOCOELA 91 Downloaded by [ ] at 20:46 17 November 2017 More recently, other authors (cf. Doguet, 1994) have considered Arrhenocoela closely related to the genus Altica Geoffroy, This hypothetical affinity is based on the following characteristics: larvae ectophagous in both genera; similar shape of the median lobe of aedeagus and, especially, of the spermatheca; host plants represented exclusively (.Arrhenocoela) or partially (Allied) by Ericaceae. On the basis of some characters of Galerucine type hindwing venation (Fig. 18A), such as: the presence of Cu la (associated with Cu lb ); the presence of the crossvein cu la -cu lb ; the Pcu directly associated with Cu lb to form the lcuc (cf. Suzuki, 1994), the genus Arrhenocoela can be considered a taxon of ancient origin. However, on the basis of some morphological characters mainly regarding the particular shape of the median lobe of aedeagus (Figs 13-16) and abdomen in male (Fig. Fig Arrhenocoela lineata, first-instar larva. A, first-instar larva in the flower of Erica arborea. B, first-instar feeding in the ovary. C, mesothoracic spiracle. D, abdominal spiracle, segments I-VII. E, abdominal spiracle, segment VIII. F, meso- and metathorax, dorsal view. G, abdominal segments VI-VIII, dorsal view. H, left side of the prothorax, dorsal view. 8), spermatheca (Fig. 18A, B), ovopositor styli (Fig. 17C, D) and spiculum ventrale (Figs 18A, D) in females, this Mediterranean genus shows evolutionary novelties that make more problematic the interpretation of its systematic position within the Western Palearctic flea beetle fauna. Our comparative morphological study revealed that Arrhenocoela occupies a rather isolated position, with respect to the other flea beetle genera occurring in the western Palaearctic Region. In our opinion, the affinities of Arrhenocoela must be looked for in some genera from the Oriental Region (see below). With regard to the relationship between Arrhenocoela and Neocrepidodera, these two flea beetle genera show a different metafemoral spring morphology. In fact, Neocrepidodera has a metafemoral spring with an evident 'recurve flange', a characteristic not present in Arrheno- Fig Arrhenocoela lineata, firstinstar larva. A-D, mesothoracic leg, posterior, front, dorsal and ventral view. E, right maxilla, inner lateral view. F, G, right mandible, dorsal and ventral view. H, clypeus and labrum, dorsal view; I, epipharynx, inner view.

11 92 M. BIONDI, G. DE NARDIS Downloaded by [ ] at 20:46 17 November 2017 Fig Arrhenocoela lineata, flrst-instar larva. A, abdominal segments VIII and IX (pygidium), dorsal view. B, pygopodium, ventral view. C, abdominal segments VIII, IX and X, latero-ventral coela (Figs 11-12) (cf. Furth, 1985). Moreover, with respect to Arrhenocoela, the genus Neocrepidodera shows: the longitudinal furrows on pronotum long and reaching the pronotal base; the frontal tubercles not distinctly delimited and the hindwing venation more simplified with absence of Cu la. Among the genera near Neocrepidodera, the nervature Cu la is only sometimes very weakly visible in Crepidodera Chevrolat, 1836 (= Chalcoides Foudras, I860) (Jolivet, 1959; personal data). Some characters present in Arrhenocoela, such as the similar hindwing venation (Fig. 18A), the transverse antebasal pronotal sulcus laterally delimited (Fig. 3), the procoxal cavities posteriorly closed (Fig. 5) and the elytral punctation arranged in regular rows, are shared with the genus Afrorestia Bechyne, 1959 occurring in the Afrotropical Region. However, also in this case Afrorestia has a different metafemoral spring with an evident 'recurve flange' (pers. data). Concerning the supposed affinity of Arrhenocoela to Altica, we think that many significant differences existing between these two genera both in the adult and in the larva (see Table I) allow them to be considered phylogenetically distant and the shared ecological similarities such as convergences, to be evaluated. In our opinion, Arrhenocoela is more closely related to the Xuthea Baly, 1865, flea beetle genus widespread with 10 species mainly in the Oriental Region and associated with Urticaceae (Sen Gupta & Basu, 1977; Scherer, 1983; Medvedev, 1997). This affinity is based on the following shared morphological characters: procoxal cavities posteriorly closed (Fig. 5); tarsal claws appendiculate (Fig. 10); anterior intercoxal process apically clearly widened (Fig. 5); pronotal sulcus laterally delimited (by short longitudinal furrows not reaching the pronotal base in Arrhenocoela; by long longitudinal furrows reaching the pronotal base in Xuthea; pronotal base sinous (Fig. 3); frontal tubercles subtriangular, well delimited (Fig. 1); maxillary palpi with segment 4 elongate and the segment 3 not enlarged, about of the same length as segment 2 (Fig. 2); elytral punctation arranged in regular rows; metafemoral spring of the 'Blepharida morphological-group' (Furth & Suzuki, 1998) (Figs 11-12); similar hindwing venation, with evident Cu la and crossvein cu la -cujb (Fig. 18A). Unfortunately, no information about the morphology of the larval stages of the genus Xuthea is yet available. REFERENCES Bartlet E., Romani R., Williams I. H., Isidore N., Functional anatomy of sensory structures of the antennae of Psylliodes chrysocephata L. (Coleoptera: Chrysomelidae). Int. J. Insect Morphol. Embryol., 28: Biondi M., De Nardis G., Descrizione della larva di primo stadio di Longitarsus minimus e L. pratensis (Coleoptera, Chrysomelidae). Fragm. Entomol., 20: Booth R. G., Cox M. L., Madge R. B., Guides to insects of importance to man, 3. Coleoptera. C. A. B. International, Wallingford, Oxon, 384 pp. Boving A. G., Beetle larvae of the subfamily Galerucinae. Proc. U. S. natl. Mus., 75: Chevrolat L., In: P. E. Dejean (ed.), Catalogue des coléoptères de la collection de M. le comte Dejean, 2nd ed. Paris, Livr. 5, 443 pp. Doguet S., Faune de France, 80. Coléoptères, Chrysomelidae. Volume 2: Alticinae. Fédération Française des Sociétés de Sciences naturelles, Paris, 681 pp. Foudras C., Altisides. In: E. Mulsant (ed.), Histoire naturelle des coléoptères de France. Magnin, Blanchard et C ie, Paris, 384 pp. Furth D. G., Relationships of Palearctic and Neartic genera of Alticinae. Entomography, 3: Furth D. G., Suzuki K., Studies of Oriental and Australian Alticinae genera based on the comparative morphology of the metafemoral spring, genitalia, and hind wing venation. In: M. Biondi, M. Daccordi & D. G. Furth (eds), Proceedings of the Fourth international Symposium on the Chrysomelidae, Proceedings of XX I.C.E. Firenze, Museo Regionale di Scienze Naturali, Torino, pp Gruev B., Döberl M., General distribution of the flea beetles in the Palaearctic subregion (Coleoptera, Chrysomelidae: Alticinae). Scopolia, 37: Heikertinger F., Bestimmungstabellen europaischer Käfer. LXXXII. Fam. Chrysomelidae. 5. Subfam. Halticinae. 2. Gatt. Crepidodera-Verwzndschaft weitesten Sinnes. Koleopterol. Rundsch., 31: Jolivet P., Recherches sur 1'aile des Chrysomeloidea (Coleoptera). Deuxième Partie. Mém. Inst. R. Sci. nat. Belg. Sér. 2, 58: 1-152; pis. XXI-XL.

12 Downloaded by [ ] at 20:46 17 November 2017 THE GENUS ARRHENOCOELA 93 Konstantinov A. S., Vandeberg N. J., Handbook of Palearctic flea bettles (Coleoptera: Chrysomelidae: Alticinae). Contributions on Entomology, International, Associated Publishers, 1: LeSage L., Immature stages of Canadian Neochlamisus Karren (Coleoptera: Chrysomelidae). Can. Entomol., 116: Medvedev L. N., A new subgenus and species of Xuthea Baly from Burma (Coleoptera Chrysomelidae, Alticinae). Atti Soc. Ital. Sci. nat. Mus. civ. Stor. nat. Milano, 137: Perris E., Résultats de quelques promenades entomologiques. Ann. Soc. Entomol. France, 3' Perris E., Nouvelles promenades entomologiques. Ann. Soc. Entomol. Fr., 6: Ritcey G. M, Mclver S. B., External morphology of antennal sensilla of four species of adult flea beetles (Coleoptera: Chrysomelidae: Alticinae). Int. J. Insect Morphol. Embryol., 19: Scherer G., Beitrag zur Gattung Xuthea Baly (Coleoptera- Chrysomelidae-Alticinae). Entomol. Arb. Mus. G. Frey Tutzing bei Munch., 31-32: Sen Gupta T., Basu C. R., A revision of Xuthea (Coleoptera: Chrysomelidae: Alticinae) from India with descriptions of two new species. Orient. Insects, 11: Suzuki K., Hindwing venation in the family Chrysomelidae (Coleoptera). In: P. Jolivet, M. L. Cox & E. Petitpierre (eds), Novel aspects of the biology of the Chrysomelidae. Kluwer Academic Publishers, Dordrecht, pp