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

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1 Ital J Zool, (2002) = Natural history of the flea beetle genus Arrhenocoela Foudras (Coleoptera, Chrysomelidae, Alticinae) MAURI ZIO BIONDI GIUSEPPINA DE NARDIS Dipartimento di Scienze Ambientali, Uni ve rsity of L'Aquila, via \Tetoio, Coppito (AQ) (Ira l)') INTRODUCTION The genus Arrhenocoela was described by Foudras (1860), who separated it from the genus CrePidodera Chevrolat, 1836 (sensu Chevrolat, 1836) "par la disposition du corselet qui n'offre pas de dépression transversale, mais en sillon profond et limité de chaque cote par une fossette qui n'atteint pas le bord postérieur... La forme du dernie r segment de l'abdome n du male est très-remarquable ainsi que celle de l'hémicycle et de l'édeage". This tlea beetle genus includes the onl)' species A. lineata CRossi, 1790) occurring mainly in the coastal areas of the Mediterranean Basin. Our stud)' regards many aspects of this genus, such as its systematic position, new ecological and biologica l information and a complete description of the egg and the fjrst-instar larva. MATERIALS AND METHODS ABSTRACT Some taxonomical, biological and ecological notes for Arrbenocoela lineala are reponed, and the systematic position of the genus Arrbenocoela is cliscussed. The complete clescriptions of the egg a nel the first-instar larva are also given. Line elrawings and scanning electron micrographs of particul,lr morphological aspects for both the aclult ancl the larva are also provided. Adult specimens of A. linea/a living on Erica arborea (Ericaceae) were collected in September 1998 in Lido cli Ostia (Rome), O m a.s.l. (Latium, Ital )'} For the rearing, preservation, ancl slide pre paration proceclures usecl in this stucl ), we refer to LeSage (1984) ancl Biondi & cl e Na rclis (998). The materia l was prese rvecl in 70% eth )'1 alcohol. Adults and larvae were clissected using a stereoscop ic microscope. For a delaile cl morphological stucly of the anatomically minute strucrures, the Iarvae were placed in Faure liq uicl, mountecl o n slides, and observed w ith a transmitted-light microscope (Leitz, Ortholux 2: obj. lo, 25 / 1.25) or, after preparation through 'criticai point drier' ancl 'sputtering technique ', analysed witb the scanning electronic microscope (SEM, Philips XL30 CP). Abbreviations usecl: ab, abdominal segment CI to X); am, articulating membrane; ch, chorion; cl, clypeus; cls, clypeolabral suture; CUla ' first subbranch of the cu b itus vein; cul a-cu lb' crossvein between Cu la ancl Cu 11>; CUl b' seconcl subbranch o f the cubitus vein; l Cuc, first cubital celi ; cx, coxa; cls, digitifo rm seta; eb, eggburster; ech, epicho ri o n; eph, epipharynx; fe, femur; fl s, filiform seta; fr, frons; ga, galea; hy, bypopharyfl'(; i, inseltion of seta ; le, lacinia; leps, lo ng ca pitate seme; 19, ligula; lb, long sensillum basiconicum; lp, labial palpus; Ir, la brum; M3, third branch of the med ia vein; mncl, mandible; msx, mesotborax; mtb, microtubercle; nux, metathorax; mxp, maxillary palpus; oc, ocellus; Pcu, p ost-cubitus vein: pcla, peelunco late seta; p e, peritreme; pf, palpifer; pie, placoid sensillum; pm, prementum; po, p)'gopodium; psm, postmentum; pv, pulvillus: px, prothorax; py, p)'gicliul11; r-m, crossvein between meclial ancl radiai veins; sa, sensory appenclage: sb. short sensillum basiconi cum: scps, sho rt capitate seta; spo, spiracular opening; 5S, st)'loconic sensillum; sst, sensory structure; st, slipes: ti, tibia ; ti', trocante!': trn, trocantin; ts. ta rsungulus. KEY \XfORDS: Coleoptera Chrysomeliclae - Arrbenocoela linea/a - Larval morphology - Biolog)' - Taxonomy. TAt'{ONOMIC ACCOUNTS AKNOWLEDGEMENTS We thank Dr. Maria Giammalteo for he r rec hnical assistance with the scanning electron microscope (SEIvI) This research was supportecl by a grant from "Ministero clell'università e clella Ri cerca Tecnologica e Scientifica" (CLUSTER-J1 ). (Rece-ived 2/uly Accepted 9 Oclober 2001) A rrhenocoela lineata (Ross i, 1790) (Figs 1-43) MOIphological description ojthe egg (Figs 19-21) Size: le ngth = 1.12 ± 0.18 mm; width = 0.60 ± 005 mm (n = 40) Shape: subcylindrical. Colo ur: orange yel

2 84 M. mondi, G. DE NA RDIS Head (Figs 23-27, 42E-I) Hypognathous, subglobose, srro ngly sclerified (Fig. 23). Cephalic su tures compiere and c1early disrincr; endocarina straighr, black, posterio rly fused with the coronai suture; coronai suture shorr and thin along the epicranial suture inne r margino Frontal sutures Y-shapecl, pale. Epistomal suture well sclerified forming with the endocarina a unique blackish T-shapeel suture. Antennae (Fig. 24) one-segmented, supporred by a well-developed translucid articulation membrane. Anrennomere cylinelrical, wirh strongly sclerified annular ring with five sensorial pores (two dorsal and three ventral); uppe r siele 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 & McIver, 1990; Bartler et al, 1999) of which rwo longer, more rhin, distall y poinred, slightly curved; one peg-like sensilla, finge r-shapeel; six sho rt basiconic sensi Ila of which three lo nger, cone-shaped, ra ised fro m bul bous base, and three very small, denriform. Figs Anhenocoela lineata, adult. l - Head, dorsa l view (73x) 2 - Ditto, ventral view (67x). 3 - Pronotum C66x). 4 - SC litellum C 478x) 5 - Prosternllm (64x). 6 - Mesonotum (60x). 7 - Mesosternum C53x). 8 - Pygidillm d' C73x). low when laid; later, brown yellow, or reddish yellow. Surface: epichorion mgose, densely covered with sma ll proruberances; chorion translucid, without evide nr reticulation. MOlphoLogical description ol the first-instar Larva The larva of A. Lineata was first described by Perris 0873, 1876) For those times, the description supplied by the French autho r, based on mature larva e ["Long. 6 mili. " (Penis, 1876: 198)), can be considereel a goocl one. However, it is scarcely useful because practically lacking in precise references to the chaetotaxy and the distribu tion of tubercl es. We therefore believe it necessary [O report a complete and up-ro-elate re-description of the first -instar larva of this species. Size: body length = ± O15 mm; pronotal width = 0.37 ± 0.03 mm (n = 30) Habitus: subcylindrical, thickset, gradually narrowed fro m the head to the pygidium (Fig. 22) Colour body lemon yel low with blackis h cephalic capsule, pronotum, tubercles, legs and pygidium. Distribution anel nomenclature of rhe tubercles is reporred in Table II. Figs Arrhenocoela linea/a, ad llit. 9 - Front leg d' C35x). lo - Tarsal c1a w of the fro nt leg C282x) Metafe moral spring, anterior and posterior view (J51x) Dorsal view of the median lobe of the aedeagus C54x) Ditto, ventra l view of the distai part C203x) Ditto, dorsal view of the distai part C200x) 16 - Dino, lateral view C54x)

3 THE GENUS ARRHENOCOELA 85 I I I ~"! ht, #jlji ' B Epicranium posteriorly with four pairs of minute trichoid sensilla anel two pairs of sensorial pores; elorsally with four pairs of filiform setae, one pair of small setae, two pairs of sensorial pores anel 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 wiele, subpentagonal with six filiform setae, of which the centrai ones much shorter anel thin, and two sensorial pores in proximal position. Clypeus (Fig. 42H) transverse, trapezoielal, laterally rounded especially at the basai half. Anteclypeus sclerifteel, mediauy weakly inciseel, with eight trichoid sensilla; postclypeus membranose with distai margin straight. Labrum (Fig, 42H) strongly sclerified, subtriangular, with four filiform setae and two sensorial pores; distai margin meelially weakly incised. Epiphaly~,{ (Fig. 420 rectangu[ar, in the middle elensely covered by microtricha (styloconic sensilia sensu Bartlet et al., 1999); dorsal side laterally with five pairs of robust pedunculate setae, weakly uncinate; elistal margin dorsally with two pairs of filiform setae; ventral membranose area with eight groups bf campaniform sensilla, symmetrically arrangeel laterally to the ventral sagittal axis. Mandibles (Fig 42F, G) strongly sclerifieel, reeldish brown, ventrally concave and elorsally with two setae ancl two sensorial pores, five-dentate; tooth I reclucecl, vely short; tooth III serrate; tooth V shorter than IV Mola not present. No ialine process along the inner basai margino Stipes subcylindrical, thickset, partially strongly sclerified, with one pair of sensoriai pores and six pairs of setae elifferent in shape and in size, MaxilJary articulation membrane translucido 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 smajl cone-shaped sensorial structures: nine arrangecl in a circle and one centrai (similar to the short basiconic sensilla III type of Ritcey & McIver, 1990, but not arising from a raised bulbous base). BasaI segment with two sensoriai pores; middle segment with one sensorial pore and two setae; apical segment with one large placoid sensillum, one lateral external long basiconic sensillum anel one lateral inner small seta. Galea (Fig. 42E) lobate, ventrally supported by a subrectangu- Culb Fig Arrhenocoela lineata. A, spermatheca, dorsal view. B, ditto, inner laleral view. C, ovopositor slyli, ventral view. D, ditto, dorsal view. / lcuc,-""\7 ". ",, Cula cula-culb, 08mm ~ t-. l1 / li ~ B c D Fig Arrhenocoela lineata. A, hind wing B-O, spiculum ventrale with 8 th abdominal segment, clorsal, lateral and,ventral view. A

4 86 M. BIO DI, G. D E NARDIS Figs Arrbenocoela linea/a. Egg: 19 - Egg with the exit ha le (h) of the new-born larva ditto, delai! Epichorion. First-instar larva: 22 - HabiLus, later:ti view I1ead, laleral view L\: ft antenna, dorsal view Ocellus Maxillae and labium, ventro-ia teral view

5 THE GENUS ARRHENOCOB.A 87 T ABLE I - M01phological differences between the genera Arrhenocoela and Altica. Genus Arrhenocoela Genus Altica ADULT - Procoxal cavities closecl (Fig. 5) - Hind wing venation with an evident CUla' linkecl to the CU1b by a short crossvein (CLllo-CU Il,) (Galerucinae type) (Fig. 18A) - The Pcu ve in directly associated with the CUlb to form the 1Cuc; - Pcu disa ppears after associating with the CUlb (Fig. 18A) - Elytral punctation arranged in regular rows - Metafemoral spring attributable to the Blepharida Morpho-GroliP (Figs 11-12) (cf. Furth, 1985) FIRST-INSTAR LARVA - Ante nnae l -segme nted (Fig. 24) - Tllberc1es Dai e Dpi separated (cf. Table 11) - Tllbercles Dm of the abdomen present (cf. TabJe II) - Tllbercles Dpe with 1 long seta (cf. Table II) - Annlllifo rm tllbe rcles witll accessorial rooms (Figs 41C-E) ADULT - Procoxal cavities partially open _ Hind wing venation without CU l a (A lticinae type) - The Pcu vein completely fll sed w ith the Cll1b to become indistinguishable from the latter - Elytral punctatiou confused - Metafemoral spring attributable [O che Altica Morpho-Group (cf Furth, 1985) FIRST-INSTAR larva - Anteunae 2-segmemed - Tubercles Dai e Dpi united - Tllbercles Dm of che abclomen absent - Tubercles Ope with 2 long setae - Annllliform spiracles simple lar sclerite with seven setae along the apical margin anel two basiconic sensilla, peg-shaped, in the middle area. Lacinia membranose, with robust setae gathered behind the galea. Prementum me mbranose, supported by a narrow semicircul ar sclerite bearing two filiform setae anel two short basiconic sensilla. Ligula narrow and convex, with five pairs of small setae. Labial palpi twosegmented; segment 1 subcylindrica l with one sensorial pore; segments 2 w ith o ne placoid sensillum, one long basiconic sensillum and six-seven apical cone-shapecl sensorial stru ctures si mil arly to segment 3 of maxillary palpi. Postmemum shorr, sclerified with four fijiform setae and four sensorial pores. Hypopharynx with rugose surface, covered with sensorial laminate structures similar to an indented fan. 7horax (Figs 28-34, 41C, 41 F, 41H, 42A-D) Segments subcylindrica l. Prothorax slightly larger. Pronotum ejlyptical with nine pairs of setae different in length, seven pairs of minute trichoid sensiua and twothree pairs of sensorial pores. Median ecdysial line pale dividing the pronotal plate imo two symmetrical parts, each with two groups of setae, one anterior constitutecl by six filiform setae apically poimed and another by three long club-shaped setae. Mesonotllm and metanotum with similar distribution to tllbercles. Ruptor ovi blackish, dentiform, in centrai 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, mode rately elongate. Metathoracic legs larger and longer. Legs partially membranose, six-segmented with trochantin, coxa, trochanter, l'emur, 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 seta e of w hich the ventrai one very long and pointed. Femur strongly sclerified with five setae of which four along th e distai margin; membranose ventral area between femur and ti bia with two long anel robust pointed setae; tibia lo nger than l'emur, with five setae and one sensorial pore in elorsal-apical position. Tarsungullis (Fig. 32) strongly curveel anel uncinate, with one long and weakly arcuate basiconic sensillum. Abdomen CFigs 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 constitutecl by many transverse and tiule cleep folds and densely covered by subtriangular and imbricate microtubercles, apically posteriorly bent. Segment VIII with posterior dorsal tu bercles fllsed along a median line (cf. Table II) ; spiracles constituted by an atriul11 with circlllar transversal section generally surrounded by nine accessoria I chambers C'air tubes', cf. Booth et al, 1990) regularly distributed along the perimete I' Cannular multiforus spiracle) (Fig 41D, E) Pygidium (Fig. 43A) ejliptical, well sclerified in the dorsal centrai 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 filil'orm setae, si.,'\( microsetae and four trichoid sensilla.

6 88 M. BIONDI, G. DE NARDIS Figs Arrhenocoela linea/a, first-instar larva Maxi llary and labial palpi, detail ventro-lateral view, legs of diffe rent size Mesothoracic spiracle Lefr egg-burster, dorsal view Right meta thoracic leg, front view Metathoracic tarsungulus, front view tarsungu lus and pulvillus of the metathoracic leg, lateral view 34 - Pulvillus of the metatboracic Jeg, posterior view.

7 THE GENUS ARRHENOCOELA 89 Figs Arrbenocoela lineata, first-ìl1star larva TerminaI part of tl1e abdomen, latera l view Abclominal segmenr V, setae on the tubercles Dai ancl Dm, lareral view Abclominal segmenr VI, capitate sera on the tubercle Dae Abclominal segment V, capitare seta in transversal secrion Abclominal segment VI, cligiriform se ra on the tubercle D m Abdominal segments VII, VIII and IX (pygiclium), dorsa l view (for abbreviations of the tubercles see Table II). DISCUSSION for some species of Galerucinae. This author distinguished 'clava te' and 'capitate' setae according to a nar Particu!arly interesting is the shape of the larva l dorsal rower or larger apica l part. However, the morphology setae in A. lineetta obtainecl with SEM. They show, from of most of the dorsal setae in A. lineata is very particllthe thorax to the pygidium, a gradua! increase of the lar. They show a cylindrical shape in almost a 11 their capitate shape with distai part distinctly bulbous CFigs length, while the apical part is similar to a small mem 35-38). Similar setae were described by Boving (1929) branose bulb, translucid and weakly sclerified. In tran

8 ~ ~ 90 M. BlONDI, G. DE NARDIS,O..' Q C) CQiD c "O DI.nl! 0 0 l)l,p. " :OQ C) C) ~: O 6 '.0 OC)c> O O oc)o _: 0 O C) C) d O O O oc)o!oo C) C) O.0 2: :. O OC)c> O O oc)o :0 6 C) C) c 5,o > : O _! O, Q C) >, O c)c> O O OC)O -fo. 6~ C) ;;:. O b ', 0 OC)c> O O OC)O O.C) O 0 0.L' ;; ~ 000 >< X TABLE II - Distribution 01 tbe tubercles in tbe lirst-instar larva 01 The SEM morphological study also allowed us to cla A. lineata. rify th e morphology of the pulvillus and the tarsungulus OR SR (Figs 32-34). The pulvillus is constituted by a flexible, DLR ( A ì DLR EPR PR ( A ì wide and pleated membrane, inserted between the poste rior side of the tarsungulus and the ti bia; in dorsal C---=", )0" view, it appears as a he mispheric hood protecting the O PST.~ tarsungulus below. The pulvillus is asymmetricajly inserted on the distai part of the legs. Therefore, the tar Coxa <El S OPST o sungulus is anteriorly completely uncovered consenting (J~ ~.~. Opc ~ EPa Q:3 0 Dpi OLpi OEPp ~ the larva to catch at the plant without impediments O from the protection membrane. OPST (JO~ B This species generally lives in the coastal areas of O Mediterranean Basin (cf. Gruev & D6 berl, 1997); sometimes it ca n penetrate inner regions fo llowing the maquis vegetation. It is associated with Ericaceae; the adults mate fro m September to November; the la rval development is in spring; th e p upation is in the soi\. Females lay their eggs o n p lants of Erica a rborea and E. scoparia, mainly in the bifurcations of young terminai branches or on the leaves. The eggs, always singly laid, are almost completely co vered by vegetai fragments assuming a cryptic aspect. At the beginning, the coloration of the egg is orange yellow then, when it becomes l'ipe, reddish brown. The larva comes out from the egg througb the part of the chorion in contact with m C)c> O O OC)O the plant, using a small hole made with the mandibles C) do in sub-apical positio n. In laboratory, the incubatio n period is very lo ng, about 2.5 mo nths. Very probably this species spends the w inter as an egg. The first-instar larva shows an aposematic lemon yellow coloration, with cephalic capsule, pygid ium and tubercles smooth black, which contrasts strongly w ith the green leaves, the reddish brown branches, and the white C>~z~ (E. arborea) or greenish flowers (E. scoparia) of Erica. From laboratoly observations, it emerged that the larva penetrates into flowers through a small circular hole C ",","m~ made in the corolla and feeds on covering tissue of the ovary and the immature ovules (Fig 41A, B). No first-instar larva was o bserv.ed to feed on foliar parenchyma or fl oral coroll a. Regions of tbe body: 1, protborax; 2, mesorborax; 3, rnetathorax; I X, abdominal segments; DR, dorsal region; DLR, dorso-iateral region; EPR, epipleural region; PR, pleural region; SR, sternal region. Tubercle nomencl ature ancl abbreviations: D-DL-EPa, dorsal, cl orso-iate ral ancl epipleurai ante rior fused; Da, dol'sal anterior (Dai fused on rnesotborax); Oae, dorsal antero-exterior; Da i, dorsal antero-interior; OLae, dorso-iateral antero-exte rio r; OLe, clorso-iateral exterior; OLp, clo rso-iare ral posterio r; OLp-S, clorso-iateral posterior anel spiracular fu sed; Dm, dorsal median (scural tubercle); Op, dorsal posterio r (clorsal postero-inte rio r and dorsal poste roexterior fu sed on VIU); Dpi, clorsal postero-interior; Ope, dorsal postero-exte rior; Dpe-DLpi, clorsal postero-exterior and clorso-lateral postero-interior fused); EP, epip leural; EPa, epiple ural anterior; EPp, e pipleural posterior; ES, eusternal; ES-SS, eusternal anel sternell ar fusecl ; P, pleural; PST, presternal, S, spiracular; SS, sternellar: T, trocantin. sversal sectio n, these setae show centrally a longitudinal channel ru nning from the base to the bulbo Probably, the bulb contains urticating liquid; the surface of the bulb seems smooth and without po res. Tbe proximal setae of the pronotal plate and the setae on the tu bercles Dm are more properly c1avate-digitiform. The systematic position al the genus Arrhenocoela Some authors have considered Arrhenocoela closety related to NeocrePidodera He ikertinger, 1911 (= Asiorestia Jacobson, 1925 (cf. Ko nsta ntinov & Vandeberg, 1996); = CrePidodera Chevrolat, 1836 sensu Foudras, 1860], mainly for its having an evide nt pronotal transverse antebasal sul cus laterally delimited and the elytrai punctation arranged in regul ar rows. Heikertinger (1950) also includecl Arrhenocoela within his "CrePidodera-Ve rwanclschaft", taxono mical grou p comprehencling some Holarctic and Orientai genera characterised by: antennae l1-segmented; pronotum normally with a transverse antebasal sulcus laterally clelimited; elytral punctation arranged in regular rows; procoxal cavities generally posteriorly closed; tibiae unmodified.

9 THE GENUS ARRHENOCOELA 91 l' b.'1 dioll, ~D 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. More recently, other authors (cf. Doguet, 1994) ha ve consiclerecl Arrbenocoela closely relatecl to the genus Altica Geoffroy, This hypothetical affinity is basecl on the following characteristics: larvae ectophagous in both genera; similar shape of the meclian lobe of aecleagus ancl, especially, of the spermatheca; host plants representecl exclusively (Arrbenocoela) or partially (Altica) by Ericaceae. On the basis of some characters of Galerucine type hinclwing venation (Fig. 18A), such as: the presence of CUla (associateci with CUlb); the presence of the crossvein cula-culb; the Pcu clirectly associatecl with CUlb to form the 1Cuc (cf. Suzuki, 1994), the genus Arrhenocoela can be consiclerecl a taxon of ancient origino However, on the basis of some morphological characters mainly regarcling the particujar shape of the meclian 10 be of aedeagl.ls (Figs 13-16) and abclomen in male (Fig. 8), spermatheca (Fig. 18A, B), ovopositor styli (Fig. 17C, D) ancl spiculum ventraje (Figs 18A, D) in femajes, this Mecliterranean genus shows evolutionary novelties that make more problematic the interpretation of its systematic position within the Western PaJearctic flea beetle fauna. Our comparative morphojogical stucly revealecl that Arrbenocoela occupies a rather isojatecl position, with respect to the other flea beetle genera occurring in the western Palaearctic Region. In our opinion, the affinities of Arrbenocoela must be Jookecl for in some genera from the Orientai Region (see below). With regard to the relationship between An-benocoela ancl Neocrepidodera, these two flea beetje genera show a different metafemoraj spring morphology. In fact, NeocrePidodera has a metafemoraj spring with an evident 'recurve flange', a characteristic not present in Arrbeno rç:;~ '/, tl' I A Y PV ",L:::.J\~~~~. f!',bcd O.l rn m D 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 ancl.. G labrum, c10rsal view; I, epipharynx, H inner view.

10 92 M. BIONDI, G. DE NARDIS l! o; In our opinion, Arrhenocoela is more closely related to the Xuthea Baly, 1865, flea beetle genus widespread with lo species mainly in the Orientai Regio n ane! associated with Urticaceae (Se n Gupta & Basu, 1977; Scherer, 1983; Mee!vee!ev, 1997) This affinity is based on the following sharee! morphological characters: procoxal cavities posteriorly closed (Fig. 5); tarsal claws a ppencliculate (Fig. lo); 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 longituclinal furrows reaching the pronotal base in Xuthea; pronotal base sinous (Fig. 3); frontal tuberc1es subrriangular, well delimited (Fig. I); maxillary palpi with segment 4 elongate and the segment 3 not enlargee!, about of the same length as segment 2 (Fig. 2); e lytral puncta ti o n arranged in regular rows; metafemoral spring of the 'Btepharida morphologica l-group' (Fu rth & Suzuki, 1998) (Figs 11-12); similar hindwing venation, with evident Cu la and crossve in cula-culb (Fig. 18A). Unfortunate ly, no info rmation abollt the morphology of the larval stages of the genus Xuthea is yet available. REFERENCES B Fig Arrhenocoela lineata, first-instar larva. A, abclo minal segments VIII ancl IX (pygicliu m), clorsal view. B, pygopoclium, ventrai view. C, abclominal segmems VIIl, IX,lncl X, latero-vemral view. coeta (Figs 11-12) (cf. Furth, 1985) Moreover, with respect to A rrhenocoela, 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]a- Among the genera near Neo crepidode ra, the nerva ture CU 1a is only sometimes very weakly visible in CrePidodera Chevrolat, 1836 (= Chalcoides Foudras, 1860) Oolivet, 1959; personal data). Some characters present in Arrhenocoela, such as the similar hine!wing venation (Fig. 18A), the transverse antebasal prono tal sulcus late rally delimited (Fig. 3), the procoxal caviries posteriorly closed (Fig. 5) and the elytrai punctation arrangee! in regular rows, are shared with the genus Afrorestia Bechyné, 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 ane! in the larva (see Table I) allow them to be considerecl pbylogenetically distant and the sharee! ecologica I similarities su cb as convergences, to be evaluated. Banlet E, Romani R.. Williams I. H., Is icloro N., Functional a natomy of sensory structures o f the amennae of Psy//iodes chrysocephala L. (Coleoplera: Chrysomeliclae ). 1m. ). Insect Morphol. Embryo l., 28: Bio ncli M., De Narclis G., Descrizione clella larva cii primo staclio cii Longltarsus minimus e L pratensis (Coleoptera, Chrysomeliclae). Fragm. Ento mo l., 20: Booth R. G., Cox M L., Maclge R. B., Guicles to insects of importance to ma n, 3. Coleoptera. C. A. B. Imernational, Wallingforcl, Oxon, 384 pp. Boving A. G, Beelle larvae of lhe subfamily Galerucime. Proe. U S. nati. Mus., 75: 1-49 Chevrolat L., In: P E. Dejean (ecl ), Ca talogue des coléoptères cle la collection cle M. le comte Dejean, 2ncl ecl. Paris, Livr. 5, 443 pp. Doguet S., Faune cle France, 80. Coléoptères, Chrysomeliclae. Volume 2: Alticinae. Féclération Française cles Sociétés cle Scie nces naturelles, Paris, 681 pp. Fouclras c., Altisicles. In: E. Mulsant (ecl.), Histoire naturelle cles coléoptères cle France. Magnin, Blancha rcl e t (ie, Paris, 384 pp. Furth D. G., Relationships of Palearcti c ancj Neartic genera of Alticinae. Entomography, 3: Furth D. G., Suzuki K, Stuclies o f Orientai anel Au str:dian Alticinae genera basecl o n the comparative morpho logy of the me tafemoral spring, genitalia, ancl hincl wing venatio n. In: M. Bi o ndi, M. Da ccorcli & D. G. Furth (ecls), Proceeclings of the FOLlIth inte rnarional Symposium o n the Chrysomeliclae, Proceeclings of XX l. C.E. Firenze, ] 996. Museo Regionale cii Scienze Naturali, Torino, pp Gruev B., Dbberl M., ] Generai distributio n of the flea beetles in the Palaearctic subregion CCo leoptera, Chrysome licl ae: Alticinae). Scopolia, 37: f-1 e ikertinger F, Bestimmungstabellen europaischer Kiifer. LXXXII. Fam. Chrysomeliclae. 5. Subfam. Halticinae. 2. Gatr. Crepidodera-Verwanclscha ft weitesten Sin nes. Koleoptero l Runclsch., 31: Jolivet P., Recherches s ur l'aile des Chrysomeloicl ea (Coleoptera). Deuxième Partie. Mé m. lnst. R. Sci. nar. Belg. Sér. 2, 58: 1-] 52; pls. XXI-XL

11 THE GENUS ARlU-IENOCOELA 93 Konstantinov A. S., Vandeberg N. J, Hanclbook of Palearc tic flea bettles (Coleoptera: Chrysomeliclae: Al ticinae). Contribu lions o n Entomo logy, Inte rnational, Associate ci Publishers, 1: LeSage L., Immature stages of Ca naclian Neoch lamisus Karren (Coleopte ra : Chrysomeliclae). Can. Entomo!., 11 6: Medveclev L. N., A new subgenus <l nd species of Xu/hea Bai)' from Burma (Coleoptera Chrysomelidae, Alticinae). Atti Soc. ltal Sci. nat. Mus. civ. Stor. nat. Milano, 137: Pe rris E., Résllltats de qllelqlles pro rnenades entomologiqlles. Ann Soc. Entorno!. France, 3: Perris E., NOllvelles pro menades entomologiqlles. Ann. Soc. Entomo!. Fr., 6: Ritcey G. M., McJver S. B., External rnorphology of antennal sensilla of fom species of adult flea beetles (Coleoptera: Chr)'so rnelidae : Alticinae). Ior. J lnsect Mo rphol. Embryol., 19: Scherer G., Beitrag Wl' Cattung X ulhea Baly (Coleoptera Chrysomelid <le-alticinae). Eoto mo l. Arb. Mus. G. Fley Tu tzing bei Mlinch., Seo Cupta T., Basll C. R., A revisio n of Xu /hea (Coleoptera: Chr)'sornelicl ae: Alticioae) from India w ith descliptions of two new species. Orient. Insects, 11 : Suzuki K., Hindwing venatio n in the family Chrysomelidae (Coleoptera). In: P. Jolivet, M. L. Cox & E, Petltpierre (eds), Novel aspects of the biology of the Chlysomelidae. KllI wer Academic Publishers, Dordrecht, pp

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