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HERMESIA LEFEVRE, A RESURRECTED GENUS OF NEOTROPICAL EUMOLPINAE (COLEOPTERA : CHRYSOMELIDAE) PDF

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PROC. ENTOMOL. SOC. WASH. 97(1), 1995, pp. 35^5 HERMESIA LEFEVRE, A RESURRECTED GENUS OF NEOTROPICAL EUMOLPINAE (COLEOPI ERA: CHRYSOMELIDAE) R. Wills Flowers A&M Agricultural Research Programs, Florida University, Tallahassee, Florida 32307. Abstract.—Hermesia Lef^vre is reinstated as a valid genus name in the Neotropical Eumolpinac. The genus is redefined to include the species //. aurata (Olivier), //. cyanea Bowditch, and //. incnnis Bowditch. Characters for separation of Hermesia and Hylax Lefevre, and the disposition of other species formerly placed in Hermesia are given. Taxonomic characters found in the male endophallus are discussed and illustrated. Key Words: Chrysomelidae, Eumolpinae, Hermesia, Hylax, ovipositor, endophallus In the course of making an inventory of modified hind tibiae ofthe male //. auratus the Eumolpinae of Costa Rica and devel- (Fig. 1) were not spinose as in most Hylax oping a key to Central American genera of (Fig. 2), //. auratus was not otherwise sep- this taxonomically confusing subfamily, I arable from the rest of Hylax. Still later, repeatedly encounter instances where in- Bechyne (1955) reiterated his transfer of77. correct application of genus names causes auratus, placed Rhahdopterus violaceus Ja- confusion and needlessly complicates rec- coby as a subspecies o^Hylax auratus, and ognizingphylogenetically meaningful groups renamed Hylax violaceus(Jacoby)asHylax ofspecies. In the present paper I discuss the pseudoviolaceus. correctplacementofasmall butshowygroup While working with Costa Rican Eumol- ofeumolpines that are widespread in Neo- pinae I found many specimens ofa species tropical forests and are frequently found in lacking any modification ofthe hind tibiae general collections ofChrysomelidae. ofthe male but agreeing closely with Hylax Lefevre(1877)established thegenusHer- auratus in all other respects. This form mesiato include ColaspisaurataOlivierand proved to be Hermesia inermis Bowditch two new species, //. purpurea and //. ful- listed under Parachalcoplacis by Bechyne gicIicoHis, later adding //. janthina Lefevre (1953) in his catalog. On further study, it (1885). By the time ofBlackwclder's check- becameclearthat //. auratusand P. inermis list (1946), 12 additional species had been are congeneric, that both show substantial describedby Bowditch(1921),Jacoby(1882, differencesinstructureofthepronotum from 1900a, b, 1904), and Weisc (1921). In his all Hylax that I have been able to examine, catalogue of the Neotropical Eumolpinae, and that neither species is congeneric with Bechyne(1953)synonymi/edHermesiawith Chalcoplacis (= Parachalcoplacis as syn- Hylax Lefevre (1884) and transferred Her- onomi/cd by Monros and Bechyne 1956). mesia aurata (Ol.) and Hermesia cyanea I have examined a long scries of/^. inermis Bowditch to thisgenus. The remaining spe- in the collections ofthe National Biodiver- cies ofLeftvrc's Hermesia were transferred sity Institute ofCosta Rica, identified spec- to othergenera (sec second list below). Lat- imens o\'Hermesia and Hylax in the Bow- er, Bechyne (1954) stated that although the ditch Collection of the Museum of 36 PROCEEDINGSOFTHE ENTOMOLOGICALSOCIETY OFWASHINGTON \\v VOLUME 97, NUMBER 37 ComparativeZoology, Harvard University, endophallic lateral digits of Askevold, are and general collections of Eumolpinae at associated with complex basal supporting Cornell University and the California De- block(BSB)fromwhichalongflexiblescler- partment ofFood and Agriculture. As a re- ite(MEG) protrudes forward; the ELD'sare sult of these studies, I regard Hennesia as thus subapical and the endophallus contin- a valid genus, based on external characters ues distally beyond them, sometimes for a and supported by internal characters ofthe considerable distance. At the base of the male endophallus and the female oviposi- endophallus there are additional sclerites, tor. presumablyderived from thedorsalandba- solateral sac supporting sclerites of Aske- Endophallic structures used in vold. Among these is another pair of less this study sclerotized but movable basal lateral digits Terminology used for structures ofinter- (BLD) which apparently have no homolog nal female genitalia follow Askevold and in Donaciinae (Askevold, pers. com.). Be- Flowers (1994). There is no accepted ter- low the two sets of digits, there is often a minology for the structures ofthe male en- field of fine setae on the underside of the dophallus ofEumolpinae. Askevold (1988, endophallus (bb). 1990, 1991)hasstudied theDonaciinaeand Themaindifficulty inevertingeumolpine identified a system of sclerites ofthe male endophalliistryingtopullthiscompactmass endophallus. Ofthese, the endophallic lat- of sclerites, which is under tension when eraldigits(ELD), thebasal supportingblock retracted, out through the delicate mem- (BSB),themedianejaculatoryguide(MEG), braneous tube, and work it free without and the basal setal brush (bb) appear to be tearing the membrane. Failures are fre- analogous to structures visible in the en- quent. Fig. 8 shows the endophallus of H. dophalli ofa number ofEumolpinae I have aurata after an unusually successful prep- studied. I am provisionally adopting As- aration in which both sets of lateral digits MEG kevold's terminology in this paper to refer and the are everted. However, even to these structures, which are analogous in here over half the endophallus is still re- position andappearanceto those illustrated tracted. The ELD'sareshown inapicalview fortheDonaciinae.Themostimportantdif- in Figs. 10 and 14. Basal digits (BLD) are ferencebetween theDonaciinaeandthe Eu- blunt membraneous lobes with sclerotized molpinae is that in the Donaciinae the en- distal surfaces (Fig. 13). Distad ofthe basal dophallus is relatively short with the ELD's block, the endophallus is a simple mem- in the apical position. In most Neotropical branous tube with a small internal apical genera ofEumolpinae so far examined, the sclerite (Figs. 11, 16) that may be a guide endophallus is a very long tube that, when for the flagellum (not illustrated). retracted, is doubled back upon itself and Hermesia Lefevre 1877: clxxviii lies within the basal hood. There appear to resurrected name be two groups ofsclerites that deploy as the endophallus is everted. The apical lateral Type species: Colaspis aurata Olivier digits (ELD in Figs. 8-17), analogous to the 1808, designated by Bechyne (1950a). Figs. 1-7. External characterso{Hennesia and Hylax. 1, 2, posteriortibia ofmale (redrawn from Bechyne 1949). 1, Hermesiaaurata. 2, Hylax viani. 3, fore (left)and middle (right) tibiae and tarsi oiHermesiaaurata. 4, 5, prothorax oiHermesia aurata. 4, pronotum. 5, lateral view ofprothorax. 6, 7, prothorax oiHylaxsp. 6, pronotum. 7, lateral viewofprothorax. CXI = insertion offorecoxa. 38 PROCEEDINGSOFTHE ENTOMOLOGICALSOCIETY OF WASHINGTON — / ./ ^MEG 11 Figs.8-n. Hennesiaaurata,malegenitalia.8,lateralviewofmedianlobewithpartiallyevertedendophallus. 9, apex of median lobe. 10, apical view ofELD's. 11, apical sclerites ofendophallus. Abbreviations: AF = apodeme, bb = basal setal field, BH = basal hood, BLD = basal lateral digit, BS = basal spur, BSB = basal supportingblock, ELD = endophallic lateral digit, MEG = median ejaculatoryguide, SBF= subbasal fenestra. Body oblong; length 4.8-6.2 mm; head, Antennae with scape oval, pedicel sub- pronolum, elytra, underside, and legsbright globose, shorterthan scape,distinctlyshort- metallic green, gold-green, or cobalt blue. er than segment 3; scape and pedicel yel- Head with labrum apically emarginate, lowish to reddish brown ventrally, metallic with 2 submedian setae and short row of green or cobalt blue dorsally; segments 3-6 lateral setae along outer apical angle. Frons reddish brown to piceous, usually paler at and clypeus coarsely, sparsely punctate; extreme apex, remaining segments piceous; punctures on clypeus separated by distance all segments filiform, each slightly wider at equal to the diameter ofa puncture, and on apex, elongate (L/W seg. 7 = 3.5-4); seg- frons by distance greater than the diameter ments 3-6 with scattered adpressed setae, of a puncture; surface between punctures segments 7-1 densely pubescent, with 1 smooth or with widely scattered punctulae; whorl of long erect setae at apex of each antennal calli impunctate; genae with mi- segment from 3-10; segment 1 1 short, con- croreticulateareaanteriorlybetweeneyeand ical. base of mandible; frons with longitudinal Mouthparts reddish brown to piceous; impressed median line, deep between an- maxillary palpwith apical segment spindle- tennal calli, becoming obsolete toward ver- shaped. Mandibles short, broad, strongly tex. Eyes oval, weakly emarginate at anten- angulatelycurved,withoutersurfacesmooth nal insertion. and shiny; apical teeth broad, acute. VOLUME NUMBER 39 97, 1 BLD / 13 ELD K BSB '•vjr' 14 16 Figs. 12-16. Hermesia inermis, male genitalia. 12, lateral view ofmedian lobe with partially everted en- dophallus. 13, frontal view ofBLD's. 14, apical viewofELD's. 15, apex ofmedian lobe. 16, apical scleritesof endophallus. Abbreviations as in Figs. 8-11. Prothoraxdistinctly widerthan long, L/W erect yellowsetae, intercoxal process broad, = 0.48-0.61; disc ofpronotum evenly, fine- 1.1-1.6X diameterofprocoxa, widened be- ly punctate, punctures separated by 1-2 hind procoxae, longer than mesostemum, times the diameter of a puncture, surface subequal to metastemum. between punctures smooth, shining, with Mesostemum broad, subequal in width scattered punctulae. Apical and basal mar- to prostemum, strongly convex between ginal bead narrow; lateral margin evenly coxae, width 1.4x width ofmesocoxa, sur- rounded, with broad distinct flange, broad- face smooth, slightly wrinkled with sparse estat middle, andwith finetransversestriae short yellow setae. on dorsal surface offlange (Fig. 4). Anterior Metasternum smooth, slightly swollen and posterior angles distinct, outwardly di- anteriortometacoxae,withsparseshortyel- rected, formed from projections of lateral low setae; metepistemum gradually nar- flange, each angle with single longerect seta rowed posteriorly, surface alutaceous. in large puncture. Proepisternum with an- Legssparselycoveredwith shortprostrate terior margin straight, surface alutaceous to setae, all surfaces alutaceous. Femora wrinkled. Proepimeron coarsely punctate, strongly swollen in middle; tibiae gradually punctures separated by distancegreaterthan widening toward apex, multicarinate, the diameterofa puncture, surface smooth, slightly to moderately sulcate between ca- shiny. Prostemum weakly declivous ante- rinae, with setae linearly arranged in sulci rior to procoxae (Fig. 5), surface with long and increasing in length toward apex oftib- 40 PROCEEDINGSOFTHE ENTOMOLOGICAL SOCIETY OFWASHINGTON BLD ""^-x'---. VOLUME 97, NUMBER 41 '------>—SD 18 t2^ 23 20 19 Fig. 18-25. Femalecharactersof//erm^5/aai/ra/aand//v/a.vsp. 18-19, ovipositorof//crwraaawra/a: 18, apex, lateral view; 19, ventral view. 20, Hyla.xsp., base ofovipositortube(sternum VIII), ventral view. 21, 22, spermatheca: 21, Hermesia aurata; 22, Hyla.x sp. 23, pygidium ofHermesia aurata. 24, 25, female sternum VII:24,Hermesiaaurata;25,Hylaxsp.Abbreviations:ALA=apicolateralarmsofsternumVIII,AS=apodeme ofsternum VIII, B = baculum, CS = coxostyli, DS = dorsal sclerites ofsegment VIII, GC = gonocoxae, HS = hemistemitesofsegment XI, PP = paraprocts, SD = spermathecal duct, SG = spermathecalgland. 4 . 42 PROCEEDINGS OFTHE ENTOMOLOGICALSOCIETY OFWASHINGTON 8, 12),apexpointed(Figs. 9, 15); basal hood Hermesia inermis Bowditch 1921: 194 (BH)long,lightlysclerotized,apodemes(AP) (original description) distinctat lateral margins ofhood; subbasal Parachalcoplacis inermis; Bechyne fenestra (SBF) present; basal spurs (BS) 1953: 170 (catalogue) prominent; tegmen slender. Endophallus Rhabdopterus violaceus Jacoby 1882: elongate, with two pairsoflateral digits, the 151 (original description), new basal pair (BLD) simple, lobe shaped (Figs. synonymy 12, 13), the apical pair (ELD's) bifurcate, Hylaxauratusviolaceus; Bechyne 1953: articulating with a basal supporting block 165 (catalogue) (BSB); a ventral basal brush (bb) between The remaining species listed as Hermesia the two pairs oflateral digits; with thin me- in Blackwelder (1946) were placed by Be- dian ejaculatory guide (MEG) projecting chyne (1953) and Bechyne and Bechyne f8,or1w0a);rdwiwthhesnmaelnld,oipnhtaelrnlauls,icsoemvpelretxedsc(Feirgist.e (n1o96j1u)dgienmtehnetsfoaltlotwhiinsgtgiemneeraa.s Itocatnhemacokre- at tip ofendophallus (Figs. 11, 16); flagel- lum extremely long. rectness ofthese generic placements; I can Female genitalia.—Segments VIII-XI onlyconfirm thattheydonotbelonginHer- mesia as defined herein. forming elongate ovipositor (Fig. 19). Ster- num VIII with longrod-like basal apodeme Corysthea B2Ar. 1865: 336 (A8) and weakly sclerotized apicolateral gregalis (Weise) 1921: 49 arms (ALA) with several setae; dorsal scle- rw/^a (Weise) 1921: 49 rites (DS) weak, rod-like. Segment IX cov- Hermesilla Bechyne: 1954: 216 ered with minute setae in basal half; hem- fulgidicollis (Lefevre) 1877: clxxix istemites (HS) with long basal rods, poorly / lampros (Jacoby) 1900a: 352 sclerotized apically; paraprocts (PP) sepa- janthina (Lereyre) 1885: 39 rated into pair ofslender dorsal rods, api- 5/m/7/5 (Bowditch) 1921: 193 cally forming hood-like projection above Allocolaspis Bechyne: 1950b: 81 genital orifice (Fig. 18); baculum (B) dis- brunnea (Jacoby) 1900b: 489 tinct, apical, subequal in length to gono- confusa {Bo\^d\Xc\\) 1921: 194 coxae (GC). Gonocoxae narrow, elongate, 7^co/)y/ (Bowditch) 1921: 193 with longsetae in apical half; coxostyli (CS) Lyraletes Bechyne: 1952: 15 distinct,withseverallongapicalsetae. Sper- purpurea (Lefevre) 1877: clxxix matheca (Fig. 21) with fine annuli, nar- varicolor (Jacoby) 1904: 51 rowed at ramus; duct short, straight, trans- Ledesmodina Bechyne: 1951: 263 parent, with sclerotized outlet into bursa. erosula aenea (Jacoby) 1900a: 351 Key to Male Hermesia Species included Hermesia aurata (Olivier) (Female Hermesia cannot be distin- Colaspis aurata Olivier 1808: 882 guishedatpresentexceptbyassociationwith (original description) males.) Chalcophana nitidissimus Erichson 1 Innermarginofhindtibiastraight,lackingapi- 1847: 162 (original description); cal emarginate area H. inermis Bechyne 1953: 165 (catalogue) Innermarginofhindtibiaexpanded,withapi- Hylax auratus; Bechyne 1953: 165 cal emarginatearea lined with longsetae(Fig. 1) 2 (catalogue) 2. Hind tibia expanded at middle, apical emar- Hermesia cyanea Bowditch 1921: 193 ginateareaalmost one-halfthe length oftibia (original description) H. aurata VOLUME NUMBER 43 97, 1 Hind tibia expanded in apical third, emargin- anteriorly and its anterior margin meetsthe ateareanomorethanone-fourththelengthof gula well behind the mouthparts. This con- tibia H. cyanea dition is widespread in the Eumolpinae and can be seen in the familiarNorth American Discussion ColaspisandBrachypnoea(=Nodonota, see The single male of Rhabdopterus viola- Flowers et al. 1994) species. In Hylax, on ceus mentioned by Jacoby and deposited in the other hand, the anterior margin of the the Bowditch Collection isaHermesialack- prostemum is somewhat to distinctly con- ingtibial modifications, hencethis form be- cave for reception ofthe postgenal area of longs underH. inennis. Both H. aurataand the head. Viewed in profile (Fig. 7) the pro- H. inermisarefoundinbrightmetallicgreen stemum appears to meet the gula almost and cobalt blue color forms (all specimens perpendicularly. When the head is in the of//, cyanea I have seen are metallicgreen). restingposition, theprostemumrestsagainst Maleso{HermesiaI haveseenfromCentral thegulaclose to thebaseofthe mouthparts. America have all been //. inermis; this spe- This condition is less common in the Eu- cies also extends into Colombia (Bowditch molpinae but is found in other Neotropical Collection); H. aurata and //. cyanea are genera such as Eumolpus Weber and Le- apparently limited to South America. pronota Chapuis. Inthe field, thebrightlycoloredHermesia ThepronotallateralmarginalbeadofHy- species are likely to be confused only with lax is narrow, evenly rounded, lacks stria- members of a group of Colaspis that also tions and is slightly thickened as it passes are bright metallic green or cobalt blue (the around the apex of the apical angles (Fig. ""bridarollei"' group ofBechyne). These Co- 6). The anteriorapical anglesare very large, laspis, however, have clear yellow to rufo- directed forward and formed from the pro- testaceouslegs(inHermesiathelegsare me- notum itself Thisisquitedifferentfrom the tallic, always the same color as the elytra) widestriatebeadofHermesia(Fig. 4)which and much more densely and/or coarsely ends in a smallbeveled angle at the anterior punctate elytra and pronotum. Host plant apical angle. data for these species are (as usual for the The apical abdominal stemite ofthe fe- Eumolpinae) scarce but I have collected //. male is quite different between the two gen- aurata from leaves of Psychotria (Rubi- era for all specimens examined. In Her- aceae) in Rondonia, Brazil. mesia, the lateral margins are strongly Hermesia, as here delineated, is most crenulate and the apical notch is narrow, similarto Lyraletes Bechyne butcan bedis- with a median tooth (Fig. 24). In Hylax, tinguished by the following combination of this stemite is smooth on the margins and characters (based on specimens referable to theapex hasabroadbisinuateemargination Lyraletes in the Bowditch Collection): the (Fig. 25). lateral flange ofthe pronotum is distinctly There are some differences between the sinuateinLyraletes, evenlyroundedinHer- genitalia ofHermesia and Hylax (based on mesia; and theelytrao{Lyraletesare widest dissectionofaseriesof//, auratafrom Peru, in their apical third (in Hermesia the sides //. inermis from Panama and Costa Rica, are subparallel). and specimens ofHylaxnr. chalybaeus Lef Aside from differences in the hind tibiae from Costa Rica). In females, segment VIII of the males (discussed above), Hermesia ismoreextensivelysclerotizedinHylax, the can be clearly differentiated from Hylax on basal apodeme has a more marked apical the following characters. In Hermesia the expansion (Fig. 20); the spermatheca has a prostemum (Fig. 5) is gradually declivous different shape (Fig. 22), and the sperma- 44 PROCEEDINGSOFTHE ENTOMOLOGICALSOCIETY OFWASHINGTON — thecal duct is very long and convoluted. In . 1990. Reconstructioned phylogeny and re- males, thegeneral morphology oftheHylax classification ofthe genera ofDonaciinae (Cole- optera: Chrysomelidae). Quaestiones Entomolgi- endophallus is like that of Hermesia but —cae 26: 601-664. shape of both pairs of lateral digits (ELD 1991. Classification, reconstructed phyloge- and BLD) is different in the two genera. In ny,. and geographic history of the New World Hylax the basal setal brush is much more members ofPlateumaris Thomson, 1859 (Cole- extensive than in Hermesia, the setae are optera: Chrysomelidae: Donaciinae). Memoirs of theEntomological SocietyofCanada 157: 1-175. arranged indefinite rowsandthereare rows Askevold, I. S. and R. W. Rowers. 1994. Glyptos- of setae on the dorsal surface behind the celoides dentatus, a genus and species ofEumol- basal digits (Fig. 17). The median lobe apex pinae new to Chile (Coleoptera: Chrysomelidae). also differs slightly but this character varies Revista Chilenade Entomologia. (In press.) within genera and is not ofgeneric value. It Baly,J.S. 1865. Descriptionsofnewgeneraandspe- cies ofPhytophaga. Transactions ofthe Entomo- isdifficultto say ifdifferencesin thesestruc- logical Society ofLondon, series 3 2: 333-357. tures identified here are phylogenetically Bechyne,J. 1949. ListprovisoiredesEumolpidesde useful at thegenus level. They do, however, laRepubliqueArgentine[etobservationsdiverses weigh in favor ofrestoring Hermesia. sur les Eumolpides de TAmerique du Sud] (Co- leoptera, Chrysomeloidea). Acta Zoologica Lil- Acknowledgments loana 8: 457-535. This research was begun during a six- rA.mer19i5q0uae.duLeSsudgeentedruotCyepnetsredeasveeculmeosldpiiadgensosdees month sabbatical with the National Biodi- des formes nouvelles. (Coleoptera, Phytophaga, versity Institute of Costa Rica (INBio) in Chrysomeloidea). Mittilungen Miinchen Ento- 1991. I thank Dr. Rodrigo Gamez, MSc. mologische Gesellshaft 40: 264-292. Angel Solis and the INBio staff for their . 1950b. Notes sur les Eumolpidesde rAme- riqueduSud,(Col.Phytophaga).AnnalsandMag- many kindnesses during that time. I also azine ofNatural History. Series 12 3: 75-85. thank Dr. David G. Furth, Museum of . 1951. Liste provisoire des Eumolpides de Comparative Zoology (MCZ), Harvard Bolivieetobservationsdiversessurlesespecesde University, for his invaluable assistance in rAmerique du Sud. (Col. Phytophaga). Entomo- logischen Arbeiten aus dem Museum GeorgFrey providing specimens from the Bowditch 2: 227-352. Collection for this project. I also thank Dr. . 1952. DuxiemenotesurlesEumolpidesneo- Michael C. Thomas, Florida State Collec- tropicauxdescollectionsdeITnstitutroyaldesSci- tion of Arthropods (FSCA), Gainesville, encesnaturellesde Belgique(1). Institutroyaldes Florida, and Dr. Fredrick G. Andrews, Cal- SciencesnaturellesdeBelgique. Bulletin28: 1-20. ifornia Department of Food and Agricul- . 1953. Katalog der neotropischen Eumolpi- den (Coleoptera Phytophaga Chrysomeloidea). ture, for the loan ofadditional specimens; Entomologischen Arbeiten aus dem Museum and Dr. IngolfS. Askevold for useful com- Georg Frey 4: 26-304. mentson the manuscript. Thisresearchwas . 1954. LalistedeseumolpidesdeRioGrande funded in part by a grant (FLAX 91005) do Sul (Bresil)et observationsdiversessurleses- peces neotropicales. Arquivos do Museu Paran- and in part by a National Science Foun- aense 10: 141-230. dation Mid-Career Fellowship (BSR- . 1955. Reisedes Herm G. Frey in Siidamer- 9003898). ika: Eumolpidae (Col. Phytophaga). Entomolo- gischenArbeitenausdem Museum GeorgFrey6: Literature Cited 569-657. Bechyne,J. and B. Springlovade Bechyne. 1961. In- Askevold, I. S. 1988. ThegenusNeohaemoniaSzke- secta amapaensia: Chrysomeloidea (Col. Phy- kessy in North America (Coleoptera: Chrysomel- toph.). Studia Entomologica 4: 409^28. idae: Donaciinae): Systematics, reconstructed Blackwelder, R. E. 1946. Checklistofthecoleopter- phylogeny, and geographic history. Transactions ousinsectsofMexico, Central America, theWest oftheAmericanEntomologicalSociety 113: 361- Indies, and South America. Part4. United States 430. National Museum Bulletin 185: 551-763.

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