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ISSN 0968-0454 Bulletin of Museum The Natural History THE NATURAL MSBTORV ;f/_UM -7 JUN 2001 .0 Entomology Series VOLUME 70 NUMBER 28 JUNE 2001 1 TheBulletin ofTheNaturalHistoryMuseum (formerly: Bulletin oftheBritish Museum (NaturalHistory)), instituted in 1949, is issued in four scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology. The Entomology Series is produced underthe editorship ofthe Keeper ofEntomology: R.I. Vane-Wright EditorofBulletin: Gaden S. Robinson Papers in the Bulletin are primarily the results ofresearch carried out on the unique and ever-growing collections ofthe Museum, both by the scientific staffandby specialists from elsewhere who make use ofthe Museum's resources. Many ofthe papers are works ofreference that will remain indispensable for years to come. All papers submitted for publication are subjected to external peer review before acceptance. A volume contains about 192 pages, made up by two numbers, published in the Spring andAutumn. Subscriptions may be placed forone or more ofthe series on an annual basis. Individual numbers and back numbers can be purchased and a Bulletin catalogue, by series, is available. Orders and enquiries should be sent to: Intercept Ltd. RO. Box 716 Andover Hampshire SP10 1YG Telephone: (01264) 334748 Fax: (01264) 334058 Email: [email protected] Internet: http://www.intercept.co.uk Claims for non-receipt ofissues ofthe Bulletin will be metfree ofcharge ifreceived by the Publisher within 6 months for the UK, and 9 months forthe rest ofthe world. WorldListabbreviation: Bull. nat. Hist. Mus. Lond. (Ent.) © The Natural History Museum, 2001 Entomology Series ISSN 0968-0454 Vol. 70, No. 1, pp.1-502 The Natural History Museum Cromwell Road London SW7 5BD Issued 28 June 2001 TypesetbyAnnBuchan(Typesetters).Middlesex PrintedinGreatBritainbyHenryLingLtd,attheDorsetPress.Dorchester.Dorset - w Bull. nai. Hist. Mux. Loncl. (Ent.) 70(1): 1-502 Issued 28 June 2001 — A revision of the tribe Macariini Tr \L (Lepidoptera: Geometridae: Ennominae) of Africa, - JUN 7 : Madagascar and Arabia MARTIN KRUGER LepidopteraDepartment, TransvaalMuseum, N.F.I., P.O. Box413, Pretoria, 0001 SouthAfrica. CONTENTS Introduction 2 Methods 3 Acknowledgements 4 Morphology 4 Biology 9 Biogeography 1 2 Phylogeny 9 1 Classification 2I Diagnosis and description of tribe Macariini 21 Genera included in the present study and extralimital genera 22 Checklist 25 Key to genera 3 I Platypepla group of genera 32 Acanthovalva gen. n 33 Narraga Walker 39 Plateoplia Warren 40 Sphyrocosta gen. n 4 I Milocera Swinhoe 42 Platypepla Warren 6 I Other Macariini 70 Chelotephrina gen. n 7 I Tephrina Guenee 73 Isturgia Hubner 74 Itame Hubner 1 13 Boarmioides Lucas 6 I 1 Macaria Curtis 1 17 Chiasmia Hubner 20 I Species auctorum 271 Malgassothisa Herbulot 272 References 273 Tables 472 Appendix 480 Gazetteer 480 List of species and subspecies by countries 493 Index 500 SYNOPSIS. A revision of the Macariini (Lepidoptera: Geometridae: Ennominae) of Africa, Madagascar and Arabia, comprising a total of 270 species, is presented as part of an international effort to revise the macariine world fauna. Major changes to global macariine taxonomy are proposed at generic and tribal levels. Practically all accessible ©TheNaturalHistoryMuseum,2001 M. KRUGER primary types were examined. The Macariini of the study area comprise 14 genera, three ofwhich are described as new. Five ofthese, Itame Hubner, Tephrina Guenee. Boarmioides Lucas, Macaria Curtis and Narraga Walker, are confined to Palaearctic North Africa. The tribe and the genera are defined or redefined. Revision resulted in the description of 89 new species, one new subspecies and the establishment of 30 new specific and seven new generic synonymies. Milocera dubia (Prout), Chiasmia coronoleucas (Prout), C. ammodes (Prout), C. sareptanaria (Staudinger) and C. tenuiata (Staudinger) are raised to specific status. Lectotypes were designated for 60 taxa. A total of 178 new combinations is proposed; one species is removed from Isturgia Hubner. Nine species (Isturgia triseriata (Prout), Chiasmiafuscataria (Moschler), C. geminilinea (Prout), C. infabricata (Prout), C. nana (Warren), C. semicolor (Warren), C. suriens (Strand), C. threnopis (Fletcher) and C. unifilata (Warren)) are recorded from southern Africa for the first time. Keys to subtribes, genera and species, as well as illustrations ofadults and genitalia ofall available species are provided. The phylogenetic position of the enigmatic monotypic genus Malgassothisa Herbulot is discussed and it is concluded that Malgassothisa probably represents a highly derived lineage of uncertain affinities. The morphology of the early stages and adult moths was examined for deriving taxonomic characters. Complete or partial descriptions are made for the early stages of 14 species. Information on life cycle, larval foodplants and parasitoids was collated. Distribution maps were prepared for 115 of the 116 species recorded from southern Africa. The biogeography and habitat associations of Macariini are discussed. Many species-groups exhibit fairly strong associations with one of the major biomes. Extant distribution patterns are interpreted as the combined result of historical processes and ecological constraints. Hypotheses about the evolutionary history of Macariini are provided. Cladistic analyses were carried out to determine relationships between selected macariine genera on a world level and to elucidate the phylogeny of species-groups within the two largest Afrotropical genera, Chiasmia Hubner and Isturgia Hubner. A gazetteer and a list ofspecies and subspecies on a country basis complement the study. Keywords: Lepidoptera;Geometridae;Ennominae;Macariini;Afrotropical;Palaearctic; distribution;keys;biology. INTRODUCTION nomenclaturalandtaxonomicproblemsaspossiblein theAfrotropicalMacariini,thepresentworkalsoaims to contribute towards a better understanding of the Macariini are a tribe of ennomine Geometridae that tribal classification ofthe Ennominae. Although the occurthroughouttheworldbutarepoorlyrepresented exact position ofMacariini within the subfamily re- inAustralia(Nielsenetal,1996).Theadultmothsare mains unresolved, several likely sister-groups for predominantly nocturnal, with cryptically patterned Macariiniareproposed. wings. The larvae feed on a variety ofplants; in the Asworkprogressed,itbecameclearthatthecompo- Afrotropical region they are mostly associated with sitionofthetribewasmoreinclusivethantheexisting leguminoustrees,particularlyofthelargegenusAcacia. literatureindicated.Whilemembersoftypical'macariine Itwasintended,originally,torestrictthisstudytothe genera(suchasChiasmia,MacariaorSemiothisa)are MacariiniofsouthernAfrica, whichwerelastrevised easilyrecognizableassuchbyexternalfeatures,other by Janse (1932). However, it soon became apparent generaarelessobviouslymembersofthetribe,arecog- that many questions, particularly concerning the nitionproblemthatisaggravatedinsomeinstancesby phylogenyofMacariini,couldonlybeeffectivelyad- convergence in wing pattern both within and outside dressed in a wider geographical context. It was thus Macariini.Afurthertaxonomicproblemencounteredis decided to include the entire Afrotropical fauna; the thefactthatthetwosynapomorphiesdefiningthetribe relativelyfewspeciesfoundinPalaearcticAfrica,Ara- arenotalwaysbothpresent.Themajorityofmacariine biaandtheMiddleEastwerealsoincludedbecauseof generaweredescribedbeforetheimportanceofgenitalic theirfaunisticinterest.Thepresentcontributionshould structurewasrealizedearlyinthe20thcenturyandare be seen against the background of a current inter- thusbased,inadequately,onexternalcharactersalone. national effort to revise the Macariini on as wide a Even when considering genitalic characters as well, geographical scale as possible (Holloway, [1994]; difficulties were encountered in finding suitable Ferguson,inprep.;Scoble&Kriiger, inprep.). synapomorphiestodefinethevariousgeneraof'horned' Apart from resolving as many of the numerous andunhomedMacariini. REVISION OF THE TRIBE MACARIINI Anotheraimofthisstudywastoimproveourunder- synonym Xenoneura Warren, an invalid homonym of standingofthe phylogenyot'Macariini.An important Xenoneura Scudder [Neuroptera]) (Fletcher, 1979), resultwastheidentificationofagroupofgenerawhich Tephrinopsis Warren, Gonodela Boisduval and cladistic analysis suggests form a distinct clade, the Thamnonoma Lederer(ajuniorobjective synonymof Platypepla group of genera. While sharing several Grammatophora Stephens). Several ofthe macariine charactersfromthemacariine 'groundplan',members generaWarrenrecognizedweresynonymizedrecently ofthiscladeexhibitarangeofvariationinmalegenitalic with Godonela by Holloway ([1994]). These include structure that exceeds that ofthe species-rich genera Acadra Herrich-Schaffer (type species rectistriaria likeIsturgiaorChiasmiaandcontrastswiththeiruni- Herrich-Schaffer from SouthAfrica). EvarziaWalker formfacies.Withfewexceptions,thePlatypeplagroup (type species ozararia Walker from Borneo), and isconfinedtotheAfrotropicalregion. GubariaMoore(typespeciesfasciosariaHiibnerfrom Cladisticandbiogeographicalanalysisprovidesevi- India).Others,includingBulongaWalker,Chorodnodes denceforapost-GondwananoriginofMacariini inthe Warren, PetrodavaWalkerandNadagarodes Warren, Afrotropicalregion.Furthermore,partoftheevolution- fall outside the concept of Macariini adopted in this aryhistoryofmany lineages is reflected inthehabitat study. associationsofextant members,andnotably themore derivedspeciesarelikelytohaveariseninthesavanna ecosystemofAfrica. METHODS History of family-group names The majority of specimens examined for this study Thehistoryofmacariinefamily-groupnameswassum- camefromtheextensivecollectionofsouthernAfrican marizedbyHolloway(11994]).Thefirstnametoappear GeometridaehousedattheTransvaal Museum,Preto- intheliteratureseemstohavebeen'Macaridae'(Guenee, ria, South Africa (TM). Material was also examined f1858]);thisnamethereforetakespriority(asMacariini). fromtheinstitutionslistedbelow.Withtheexceptionof Hollowaynotedthemoreextensiveusageinthelitera- the Wiesbaden Museum and the Transvaal Museum tureofSemiothisini (basedon SemiothisinaeWarren. acronymsfollow Heppner& Lamas(1982). 1894). Not only is Semiothisini a later name, but its TheNaturalHistoryMuseum(formerlyBritishMu- usagewouldcauseconsiderableconfusionduetopast seum (Natural History)), London, U.K. (BMNH) misinterpretation of the type species of Semiothisa provided many types and otherspecimens. In Europe Hiibner - a name that has been variously applied to and the United States further material was examined unrelatedOldandNewWorldspecies.Fidoninaesensu fromthefollowinginstitutionsandprivatecollections: Packard(1876),comprisingtheMacaridae,Zerenidae UniversitetsZoologiskaInstitut,Lund,Sweden(UZIL); and Fidonidae of Guenee ([1858]), are based on Naturhistoriska Riksmuseet, Stockholm, Sweden FidonitesDuponchel(1844).F/Y/rw/c/Treitschke,ajun- (NRS);MagyarTermeszettudomanyiMu/.eum,Buda- ior objective synonym of Eurranthis Hiibner (type pest, Hungary (TMB); University Museum, Oxford, species Phalaena plummistraria Villers) (Fletcher, England(OMU);DeutschesEntomologischesInstitut, 1979),differs from Macariini (as Semiothisini) in the Eberswalde, Germany; Zoologisches Museum der sense of Forbes (1948) and Holloway (| 1994]). Humboldt Universitat, Berlin, Germany (ZMHB); McGuffin(1972)includedFemaldellaHulst(withthree Zoologisches Museum und Forschungsanstalt Senck- Nearctic species) in his concept of Semiothisini, so enberg,Frankfurt/M.,Germany(SMF);Zoologisches FernaldellinaeHulst(1896)isrelevanthere.Theclose Forschungsinstitut und Museum Alexander Koenig, relationship between this genus and the Palaearctic Bonn,Germany(MAKB); MuseumWiesbaden,Ger- genusNarragaWalkerwasalreadynotedbyMcGuffin many ZoologischeStaatssammlung,Munich.Germany : (loc. cit.); indeed, Femaldella was treatedas ajunior (ZSBS);NationaalNatuurhistorischMuseum,Leiden, synonym ofNarraga by Povolny & Moucha (1959) TheNetherlands(NNML);MuseumNationald'Histoire and Hodges etal. (1983), a decision followed in the Naturelle,Paris,France(MNHN);C. Herbulotcollec- presentwork. tion, Paris, France; Museum Royal de FAfrique MostoftheearlysystematicworkontheAfrotropical Centrale, Tervuren, Belgium (MRAC); Naturhistor- macariinefaunawasdonebyWarren(1894-1914).He ischesMuseum,Basel,Switzerland(NHMB);Museum definedwhathetermedSemiothisinaeentirelyonexter- d'Histoire Naturelle, Geneva, Switzerland (MHNG); nalcharactersandconsequentlyincludedsomegenera Zoological Museum 'La Specola', Florence, Italy that do not belong in Macariini. True Macariini in- (MZF); and National Museum of Natural History, cluded by Warren were: Semiothisa Hiibner, Washington,U.S.A.(NMNH).InAfrica,materialfrom Hyostomodes Warren, Acadra Herrich-Schaffer, the following collections was examined: National EvarziaWalker,TephrinaGuenee(whichhemisapplied MuseumsofKenya,Nairobi,Kenya(NMKE) National ; to horned forms), Oxymacaria Warren (with its Museum of Natural History, Bulawayo, Zimbabwe M. KRUGER (NMB);NationalCollectionofInsects,Pretoria,South Forthedescription ofwing venation, thegenerally Africa (NCI), South African Museum, Cape Town, accepted systemofComstock& Needham (1898/99) SouthAfrica(SAM);andtheprivatecollectionsofthe wasadopted. late Mr N.J. Duke, Mbabane, Swaziland (NJD); Dr D.M. Kroon, Sasolburg, South Africa (DMK); Mr Keys H.S. Staude, Krugersdorp, South Africa (HSS) and Mr. P.S. Roos,Turffontein, SouthAfrica(PSR). Keys are provided to all species and to the species- groupswithinMiloceraSwinhoe,IsturgiaHiibnerand Approximately 14,000adultspecimenswereexam- ined,includingmostprimarytypes.Fromthismaterial, Chiasmia Hiibner. These keys are mostly based on about900microscopic slides,mostlyofgenitalia,but malegenitaliastructure,complemented,whereapplica- ble, by characters provided by female genitalia and alsoofwingsandotherbodyparts,wereprepared. generalfaciesoftheadult.Sincemanycharacterssuch Preparation of genitalia slides followed the tech- as valve shape and wing markings occur more than niques outlinedin Robinson (1976). Specimens were stainedinaweakalcoholicsolutionofmercurochrome once,andapparentlyindependentlyofeachother,itis andslidemountedinEuparal. imperativetouseallinformationprovidedinacouplet todeterminecertainspeciesaccurately. IllustrationswerepreparedusingaBritexprojection microscopeandaWILDdissection microscope (type Citation of specimen labels 181300)withattachedcameralucida. Eggspreviouslystoredin70%ethanolwereprepared Labels ofall types are cited verbatim, unless a very for scanning electron microscopy by fixing in 2.5% largenumberofspecimenswasinvolved(whensome glutaraldehyde,rinsinginphosphatebufferandfixingin repetitiveinformationwasomittedorreferenceismade 0.25%aqueousosmiumtetroxideaccordingtothemethod totheoriginaldescription).Incaseswherebothsidesof descibed in Coetzee & van der Merwe (1994). SEM alabel contain text, this is indicatedby a"/".Texton micrographsweretakenwithscanningelectronmicro- differentlabels is separatedbya ';'.Namesofcollec- scopesofthetypesJEOL840andHitachiS-510. tors unaccompanied by iegit' or a similar suffix are giveninroundbrackets,evenifthesedidnotappearon Gazetteer theoriginallabel.Lastly,wheremissingpiecesoftext Duetoinadequatelabellingofspecimens,considerable had to be inferred, these addenda are given in square brackets. difficultieswereencounteredduringpreparationofthe distribution mapsandcompilationofthegazetteer. In additiontoseveralatlases,anumberofothergazetteers was consulted (Anonymous, 1950-1972; Leistner & Acknowledgements. Theauthorwouldliketoacknowl- & edge the following persons for the loan of material and Morris, 1976; Crawford-Cabral Mesquitela, 1989; help in various other ways: M.J. Scoble, D. Carter, K. Viette, 1991).Evenso,manylocalitiesremainobscure. Buckmaster, M. Honey, L. Pitkin, G. Martin and M. Sincecompletionofthedistributionmaps,provincial Parsons (London), R. Danielsson (Lund). B. Gustafsson boundariesinSouthAfricahavebeenredrawn,increas- (Stockholm), G.C. McGavin and I. Lansbury (Oxford), R. ing the number of provinces from four (Transvaal, Gaedike (Eberswalde). W. Mey and B. Krutzsch (Berlin), Orange Free State, Natal and Cape Province) to nine H. Schroder (Frankfurt/M.), D. Stiining (Bonn), M. (Gauteng, Northern Province, North-West, Geisthardt (Wiesbaden), A. Hausmann (Munich). R. de Mpumalanga, Free State, KwaZulu-Natal, Northern Jong (Leiden), J. Minet (Paris), U. Dall'Asta (Tervuren), Cape,EasternCapeandWesternCape). IntheGazet- M.Brancucci(Basel),D.Burckhardt(Geneva),L.Bartolozzi teer, South African localities appear under the new (Florence), D.C. Ferguson (Washington). M. Mungai provincial names; throughout the descriptive section (Nairobi), R. Sithole (Bulawayo), R. Oberprieler (Pretoria, thenewnamesappearinsquarebrackets. now Canberra), M. Cochrane (Cape Town), D.M. Kroon (Sasolburg) and the late N.J. Duke (Mbabane). Measurements and terminology Forewinglengthwasmeasured,tothenearestmillime- MORPHOLOGY tre, as the distancebetween wingbase and apex. The rangesgivenarethoseofthesmallestandlargestspeci- menintheseriesexamined. Early stages (Figs 1-6) Unless indicated otherwise, measurements ofeggs andlarvaearebasedonarepresentativesampleoffive Littlepublishedinformationexistsontheearlystages specimens. ofMacariini.MostHolarcticspecieshavebeenreared, The terminology for genitalia is based on Klots but the available information exists in the form of (1970).Afewadditionaltermsarealsoused;theseare isolateddescriptionsofmaturelarvae,mostlyofspecies explainedinFigs 19and20. of Chiasmia and Isturgia. McGuffin (1972) detailed I. REVISION OF THE TRIBE MACARIINI the information available on the Canadian fauna and plete. The chaetotaxy is characteristically geometrid: Forster & Wohlfahrt (1981) on the fauna of central presenceofafourthSV setaonA6;LI andL2widely Europe. Practically nothing is known about the early separated on A1-8; seta L3 more anterior in position stagesofspeciesbelongingtothePlatypeplagroupof comparedwithotherfamilies(Stehr, 1987).Secondary genera in the study area, although a few isolated de- setaeareabsent. scriptionsexistofspeciesofNarragaandAcauthovalva Head. The larval head is hypognathous; the fromthePalaearcticandNearcticregions. frontoclypeus extends dorsally to nearly halfthe dis- Egg (Fig. 1 a-f) tance to the epicranial notch. The cranium is heavily sclerotizedandoftendarklypigmented. Sixstemmata TheeggofMacariini,likethatofotherGeometridae,is occur on the side ofthe head. Changes take place in ofthe 'flat type', in which the longitudinal axis runs craniumshapebetweeninstars,buttheseareminorand horizontaltothesurfaceonwhichitisdeposited.Eggs confined totheheadcapsule becoming more strongly ofAcauthovalva inconspicuaria(Hiibner)andoffour convexinlaterstages.McGuffin(1972)providedillus- andeightspeciesofIsturgiaandChiasmiarespectively trationsdI larvalheadsofseveralspecies.Thechaetotaxy were available for study. The eggs of most species of the cranium of the first and final instar larva of examined are elliptical and more or less oblong. The Chiasmiaprocidata is as follows (setal nomenclature rosetta-shaped micropyle is illustrated in Fig. Id.The afterStehr, 1987): eggofC. subcurvaria(Fig. lc,d),however,differs in TactileSetae beingbarrel-shaped.Thechorionhasareticulate,penta- (i) Anteriorgroup. A1-A3 on parietal lobe. Inadult orhexagonalstructureinalleggsexamined,thoughthe larva Al, the most ventral seta, some distance from pirntoecnisdiattyaapnardtocfoatrhseecnheosrsioonfisrsetmiocoutlhat.iAonerovapryyl;esinmaCy. sAtleminmafr3o,ntaboofvestaentmemnana]2;bdaosres:amlesdeitaanAs3etainA2froanbtovoef beinconspicuousorraisedandprominentlydeveloped. stemma PoreAadorsadofA2. WiththeexceptionofC. subcurvaria. morphologyof I. theAfrotropicalspeciesexaminedfallswithintherange (ii) Stemmatalgroup. S1 close tostemma4: S2 near illustratedbyMcGuffin(1972)andSalkeld(19X3). In stemma I;S3behindstemma6.nearcentreofstemmatal size,eggsexamined ranged between 0.56 x 0.41 mm semicircle.PoreSaclosetostemma4;poreSbabsent. and0.77 x0.57mm. Usuallyeggsarepaletomedium (Hi) Substemmatal group. SSI between stemma 5 greenwhen laid,turningreddishordarkening priorto andmandible;SS2nearstemma5;SS3somedistance from stemma5 towards maxilla. PoresSSatoSSc not hatching. located. Larva (Figs 2-5) (iv) Frontalseta. One pairofsetae (Fl) in centreof frons.PoreFamesadofFI,situatedbelowsetaeinfirst Platypeplagroupofgenera.Although being typically instarandslightlyabovesetaeinfinal instar. geometrid inhavingthe numberofprolegsreducedto (v) Adfrontalgroup. AF1 setaeventradofAF2;both two pairs - one on A6 and the other on A10, the groups situated between ecdysial line and lateral brownishlarvaofPlateopliaacrobeliadiffersfromthe adfrontalsuture.PoreAFaaboutmidwaybetweenAF generalizedgeometridtypebyhavingastronglyswol- andAF2. len thorax, not unlike the larva of Macaria abydata (vi) Clypeal group. As illustrated, with C2 Guenee (see PI. 19, Fig. 3 in Holloway, 1993). When ventrolateradofC1.Therearenoporesassociatedwith disturbed, itdrops totheground and rolls itselfintoa thisgroup. ball, resembling a fecal pellet (Staude,pers. cotnm.). (vii) Lateralseta. On sideofhead above stemma 1 Theearlystagesofothermembersofthegroupremain PoreLanearandslightlybelowLI butpoorlydefined unknown. and inconspicuous due to heavy pigmentation ofcra- OtherMacariini. LarvaeareknownofIsturgiaand nium. Chiasmia species only. The southernAfrican species (viii) Posteriodorsalgroup. P2closetotheendofthe examined are mostly either smooth or possess only epicranial stem; PI some distance further down. PI small tubercles, usually on segments A4 and A5; somewhat longer than P2 (but not much longer as occasionally tubercles are more numerous, as in C. statedinStehr, 1987: 301). PoresPaandPbdorsadof turbulentata.Colouration ismostlycrypticintonesof andclosetobaseofPI andP2respectively. green and brown (Figs 2. 4), although the larvae of somespeciesarebrightlycoloured(e.g., C. streniata. Proprioceptorsetae Fig. 5). Thefollowing linesoccur: dorsal (ormiddor- (i) Dorsalgroup. Setae very small. MD1 and MD2 sal),addorsal,subdorsal,lateral,subventral,adventral between Pb and vertex; MD3 dorsad of MD2 (not and ventral (or midventral). In the species studied, illustrated). Pore MDa between MD2 and MD3 (not usuallynotallofthesearepresent.Withtheexception illustrated).Setaeandporenotdiscernibleinfirstinstar ofthe lateral line, lines are often weak and/orincom- craniaatamagnificationof400x. 6 M. KRUGER (ii) Genalgroup.Thesesmallsetaearesituatedatthe L2 andL3 onT2-3; L3 and SV3, SV4 onAl; SV on lower,rearportionofthehead(Stehr, 1987)butcould A2-5; L3 onA6; SV3 onA7 andA8;andL2onA10. notbelocatedintheslidepreparationsmade. NochangesinthechaetotaxyofsegmentA9occursin eitherspecies.Thepatternofchaetotaxydescribedfor/. Thoraxandabdomen. Firstand/orfinal-instarlarvae deerrariaandC. interruptalargelycorrespondstothe of11 speciesofChiasmiaandthreespeciesofIsturgia generalizedpatternprovidedforCanadianMacariiniby wereavailableforexamination.ThesetaeofLI larvae are somewhat trumpet-shaped and grooved, with a McGuffin(1972).Morespeciesneedtobeexaminedto assessthetaxonomicsignificanceoftheobserveddif- sweirtrhatetdheritmi.pItnelramtienraitnisntgarsintheaysbmeacllomevesaitctleen.uatTehde, ferences,suchastheapparentabsenceofV1onsegments chaetotaxy ofLI and L4 larvae ofIsturgiadeerraria Tl andT2in/. deerraria. andChiasmiainterruptais: Pupa(Fig.6) Isturgiadeerraria: Firstinstar. Tl: XD1,XD2;Dl,D2;SD1,SD2;LI, With the exception of single exuviae of Platypepla L2; SV1, SV2.T2-3: Dl, SD1, SD2; LI, SV1.Al-5 macilentaandP. griseobrunnea (broken) (Platypepla Dl, D2; SD1; LI, L2; SV1, SV2; VI. A6: Dl, D2 groupofgenera),onlypupaeoftwospeciesofIsturgia SD1; LI, L2; subventral group of3 setae. A7-8: Dl andsevenspeciesofChiasmiawereavailableforstudy. D2; SD1; LI, L2; SV1; VI. A9: Dl, D2; SD1; LI All areobtectandfairly uniforminshape andcolour, SV1; VI. A10: D2; SD1, SD2; CD1, CD2; LI, L3 beingmoderatelystoutandspindle-shaped,mediumto CP1,CP2.Inthisinstar,noventralsetawasdetectedon darkbrownandslightlyglossy.Thepupaearesparsely segmentsTl andT2. setose;thesetaeareshortandconfinedtothedorsumof Fourth instar. As first instar, but with the follow- the whole body and the ventral side ofthe abdomen. ingchanges:Tl:VI present.T2-3:D2,L2,L3andVI Thecremasteris welldevelopedandin Macariini not present.A1: SV2reduced;L3;SV3,SV4present.A2- belonging tothe Platypepla grouptypically bifurcate 5: SV4present.A6: L3 present; subventral groupof5 with truncated or pointed tips frequently ofdiffering setae.A7-8:SV3present.A10:Dlpresent;L2present. length.Bycontrast,thecremasterofPlatypeplaisrhom- Crotchetsinamesoseries. boid,withtwothinfilamentouslateralprocesses.The cuticleofthebaseofthecremasterfrequentlyexhibitsa Chiasmiainterrupta: numberoffine,convolutedridges.Whendisturbed,the Firstinstar. Tl:D1,D2;SD1,SD2;L1,L2;SV1 and pupa is capable ofmoving its abdomen in a rotating SV2 arising from pinaculum; VI; additional ventral manner.Pupaeofcertainspecieshavebeenfiguredby seta (?). T2-3: Dl, D2; SD1, SD2; LI; SV1; VI; McGuffin (1972). additional ventral seta(?).Al: Dl, D2; SD1; LI, L2; Forbes (1948),buildingonearlierworkby Mosher SV1;V1.A2-5:D1,D2;SD1;L1,L2;SV1,SV2;V1. (1916),usedcremastershapetodefineennominetribes. A6:Dl,D2;SD1;L1,L2;subventralgroupof2setae. He distinguished between a Group A, in which the A7-8: Dl, D2; SD1; LI, L2; SV1; VI. A9: Dl, D2; cremasterendsintwospines,andaGroupB,inwhich SD1; LI; SV1; VI. A10: Dl. D2; SD1, SD2; CD1, it ends in eight hooked setae, placing Abraxini and CD2;L1,L3;CP1,CP2. Macariini(asSemiothisini)inGroupA.Abraxinihave Fourth instar. As first instar, butwith the follow- been proposed as the sister taxon of Macariini ing changes: Tl: XD1, XD2 present; SV1, SV2 (McGuffin, 1987;seealsosectiononPhylogeny).The separated;additionalventralseta(?)reduced.T2-3:L2, verydifferentrhomboidcremasterofPlatypeplamay L3present;additionalventralseta(?)reduced.A1:L2, well constitute an apomorphy notonly forthe genus, L3present;SV3,SV4present.A2-5:L3present;SV3, but possibly for the whole Platypepla group. It is SV4 present. A6: L3 present; subventral group of4 particularlyimportant,therefore,thatmorepupalmate- setae.A7: SV3present.A8: SV3present;VI reduced. rialofthisgroupisstudied. A10: CD3 present; L2 present. Crotchets unbroken Forbes(1948)notedtwofurtherpupalcharactersas (i.e.,ofequallengthandarrangedinonegroup). importantforthehigherclassificationofGeometridae. Thefollowingtrendsinmacariinechaetotaxycanbe The first is the development ofthe 'suture' (Forbes's recognized: the setaeofthedorsal (Dl, D2) and sub- term) separating segments A9 and A10. In many dorsal(SD1,SD2)groups,aswellasthefirsttwosetae Ennominae,althoughnottheeasternNearcticMacariini, of the lateral (LI, L2) and subventral (SV1, SV2) this 'suture' is strongly developed, forming a dorsal groupsarepresentthroughoutthedevelopmentofboth groove (a scalloped ornotched posterioredge) and a ChiasmiaandIsturgiaandsubjecttoverylittlevaria- lateral groove (atriangularorobliquedepression). In tion. Most variation is observed in the presence of the Afrotropical pupae, and in exuviae of Macariini additional lateral and subventral setae in the mature examined, I found this suture to be unmodified as in larva.ChiasmiainterruptaandIsturgiadeerrariaboth Nearctic membersofthetribe. Secondly, segmentA5 acquirethefollowingsetaeduringlarvaldevelopment: may develop what Forbes (1948) termeda spiracular REVISION OF THE TRIBE MACARIINI furrow (a shallow, usually unsculptured and shining illary palpi are very small, one-segmented and com- depressioncrossingorlyingimmediatelyinfrontofthe pletely hidden beneath scales. The proboscis is well spiracle) or a flange-plate (i.e., a sharp vertical ridge developedandnotscaledatitsbase. lying well in frontofthe spiracle, preceded by adeep groove, which isfrequentlysculpturedatthe bottom). Thorax While a tlange plate is absent in the macariine pupae studied, confirming Forbes's findings, the develop- The thorax of most species is slender as in many mentofaspiracularfurrow is apparently inconsistent geometridsthoughitmayberobustasinspeciesofthe within Macariini. It is absent in Platypepla, weakly Isturgia supergressa-gcoup. developed in Chiasmia furcata, C. interrupta, C. Legs. Themacariinelegistypicallygeometrid,be- simplicilinea, C. trizonaria and C. turbulentata and inglongandslenderwithacoveringofsmoothscales. well developed in Isturgia deerraria, I. spissata and Thetarsiarealsolong,andthetibialspurarrangement Chiasmiasubcurvaria. is 0-2-4. Modifications occur only in the male hind tibiaofmanyspeciesofIsturgiaandC/uasmiawhereit Adult - comments on skeletal may become dilated and/or develop a lateral groove morphology whichmaybelinedwithappressed,finehairsorbeara hair-pencil. Head (Figs7,8) relWatiinvgest.o boTdhyeswiziengwshiolfe tmhoosste oMfacsaormiienimeamrbeebrrsoaodf ThevestitureofthecraniuminMacariini iscomposed thePlatypeplagroupofgenera,suchasPlatypepla. are ofelongated,recumbentscalescoveringthevertexand narrower.The apex ofthe fore wing may be rounded, the antennal scapus. Scales on the vertex frequently pointed or scalloped. The termen ofthe hind wing is differ in colour from other areas ofthe head and are slightlyattenuatedinmanyspecies,givingthemaweakly often paler. The frons is usually smooth, much less 'tailed' appearance, but isalso frequently roundedor. denselyscaledandmoreorlessconvexinmostspecies. morerarely,crenulated.Thewingsofmostspeciesare C It bears a conspicuous horseshoe-shaped prominence smooth-scaled; in C. nubilata and extrusilinea, a intwospeciesoftheChiasmianubilata-groupandthe patch of raised scales indicates the position of the twoprobablymostprimitivemembersofChiasmia, C. (absent) fovea.The fovea, ifpresent, is formedby the calvifrons and C. puerilis (Fig. 8), while a simpler, sigmoid base of AI and demarcated dorsally by a scaledconeispresentinseveral morederivedspecies- recurved process ofCuA. It usually contains a corru- groups of the same genus. These structures thus gatedmembraneofuncertainfunction.Thedistribution probablyaroseindependentlyinseveralmacariineline- ofthisstructureexhibitsnoobvioustaxonomicpattern. ages. It is present in most Macariini, although in some in- Thechaetosemataarenotoftheshapetypicallyoccur- stancesincompletelyformed. ringinmacrolepidoptera(i.e.,adiscretearticularpatch ThewingpatternofmembersofthePlatypeplagroup with bristles or narrow scales situated behind each ofgeneraisrathersimple.Usuallyitconsistsatmostof antennalbasis)butweredescribedbyForbes(1948)as basal,medianandpostmedianlinesonforewinganda 'transverselyelongate' (Fig.7).Thisconditionconsti- postmedianonlyonhindwingonafairlyuniformand tutes a synapomorphy of Macariini, although it was frequentlyochreousbackground.OtherMacariinidis- apparently secondarily lost in some derived taxa. in- play,however,agreatvarietyofpatterns.Theunderside cluding the Oriental genera Iridoplecta Warrren and pattern may consist of the same elements as on the Lampadopteryx Warren. Transversely elongated upperside and simply be more or less intense, or its chaetosemata also occur outside the Macariini in composition maybeentirelydifferent. InCostaRican ScardamiaGuenee(Scardamiinae)(Holloway,[1994]). species of Chiasmia and Semiothisa, Scoble (pers. Thestructureoftheantennaevariesconsiderablyin comm.)foundthepatternontheundersidetobemuch the two subtribes, and they may be plumose (i.e., stronger, though otherwise similar to the upperside. bipectinatewithverylonganddelicaterami),bipectinate Thevariouswingpatternelementsreferredtointhetext withpectinationsofvaryinglengthandthickness,but areillustratedinFig.9.Discalspotsarepresentinmost shorter than in the plumose type, serrate, or ciliate. macariines,althoughtheymaybeinconspicuous. While many ofthe derived speciesofChiasmia have Venation throughoutthetribe isfairlyconstantand ciliateantennae,thereareexceptionsandmaleantennal ofthe normal ennominetype, showingareductionof structuremayvaryevenwithinspecies-groups(e.g.,C. M2 in the hind wing (Figs 10-18). The observed procidata-andfurcata-group). variationconcernsmostlyScandRintheforewingand Thethicklyscaledlabialpalpiareprominent,varying the development of vein 2A in the hind wing. As inlengthbetweenoneandtwotimesdiameterofeyes. already pointed out by Capps (1943), differences in Inthemajorityofspeciestheyareobliquelyporrectand ennomine venationdonot accurately reflectrelation- three-segmentedwiththethirdsegmentsmall.Themax- shipsasvariationoccursevenbetweencloselyrelated M. KRUGER groups. This observation was confirmed by Janse sclerotizedarmwhichcurvesoverthetympanumandis (1932) whoillustrated someofthe variationfound in notfoundoutsidetheGeometridae,inlsturgiapulinda SouthAfricanspeciesofChiasmia(citedasSemiothisa) (Walker); thisextension is alsopresentin theAfrican and lsturgia (cited as Tephrina). The venation ofthe species of the disputaria-group. No variation of the differentgeneraisdescribedindetailinthesystematic ansabetweenthePlatypeplagroupofgeneraandother section. Macariiniwasfound.Thetympaniclacinia,asclerotized Generally,SctendstoanastomosewithRl orR1+2 plateperpendiculartotheansa,usuallysemicircularin intheforewing;theanastomosismaybeshortorlong. Ennominaeandthoughttoprotectthetympanum(Cook The radial veins are variously stalked: combinations &Scoble, 1992),wasfoundinmanyspeciesoflsturgia includeR2-5stalkedwithR2andR3beingcoincident; and Chiasmia,butnotamongthePlatypeplagroupof R2-^l stalked with R3 and R4 being coincident; and genera. R3-5stalked,thestalkusuallyarisingfromnearendof cell. Veins 1A and 2A may be separate foruptoone- Genitalia thirdofthelengthof1A+2Abeforetheyfuse,ormay Theventralaspectofthemaleandfemalegenitaliaofa anastomosefortheentirelength. Inthehindwing,Sc+R1 andRsareapproximatedor representativeofthePlatypeplagroupofgeneraanda meetforashortdistancebutdonotanastomose.Vein 'horned'representativeofthetribeispresentedinFigs 2Aisabsentoronlyindicatedbyafold.Wherepresent, 19and20respectively. itdoesnotreachthewingmargin(Figs 10-18). Male (Figs 19, 500-744) Abdomen Theuncusisalwayswelldevelopedandmaybeattenu- ated (e.g., Fig. 538), triangular (e.g., Fig. 507) or Coremataarenotevidentinthegroup;however,males dome-shaped(e.g..Fig.594).Whiletheattenuatecon- of some species of Milocera {Platypepla group of ditionisconfinedtothePlatypeplagroupofgenera,a genera) possess groups ofelongated, hair-like scales dome-shaped uncus, with or without enlarged setal oneithersideoftheoctavals.Theirfunctionisnotclear, 'horns', is typical of many other Macariini. I call a andanassociationwithscentdistribution,probablyfor modificationofthelattertype'mitre-shaped*(e.g.,Fig. thepurposeofcourtship,isspeculative.Malesofsev- 575); this type is characteristic for the lsturgia eral generaofMacariini frequently have atransverse preshitaria-gwup. comb of setae on sternum A3. Several species of A gnathos may be present but has apparently been Milocerashowagroupofenlargedscalesinthisposi- secondarily lost several times within the Platypepla tion, which maybe theunmodifiedhomologueofthe group. Where present it may be cingulate (i.e., sup- setalcombfoundinhighermacariines. portedbyfairlystrongarms)andthenhavethemedial Octavals(paired,hairyorscobinatecaudalprocesses element either upright or drooping (e.g., Figs 507, of the posterior margin of segment A8 of the male) 557), or alternatively be deeply emarginate with thin occur widely throughout the tribe. Theirpresence or arms(e.g..Fig.594).Asadeeplyemarginategnathosis absence,andparticularlytheirshape,arefrequentlyof typicalforhighermacariinessuchasChiasmia,Icon- diagnostic value at species and species-group level. siderittobetheapomorphiccondition. ComparedwithotherMacariini,theoctavalsofmem- The valve structure ofAfrotropical Macariini, and bers ofthe Platypepla group are simpler in structure particularlythePlatypeplagroup,showsaconsiderable and weakly sclerotized. Most are either broadly w- degreeofvariation(Figs500-544(Platypeplagroupof shaped or consist of a narrow, sclerotized lip, as in genera); Figs 545-744 (other Macariini)). In the species most species of Platypepla or Plateoplia Platypeplagroup,costaandsacculusareusuallymore acrobelia(Wallengren). completelyseparatedthaninotherMacariini.However, Wherepresent,theoctavalsofMacariiniaresmallto thevalvae in sometaxabelongingtothe lattergroup, verylargeandusuallyarcuateorfurcate,withrounded suchasOxymacariaandIridoplecta,inwhichsacculus orpointedtips.Theymaybeshallowordeeplyexcised, and/orcostahavebeensecondarilyreducedtoanarrow and in some speciesbearafringeoffairly longhairs. sclerotizedbar, may at first appeartobe closertothe Octavals areabsentinbothspeciesofChelotephrina, platypepline condition. The valvula (i.e., the central themembersofthelsturgiasupergressa-groupprevi- regionofthevalva,lyingbetweencostaandsacculus) ously placed in Enconista, as well as some other isnotmodifiedinthePlatypeplagroup,andisoccasion- species-groupsoflsturgia,andhavebeensecondarily allydevelopedasasmall 'hump'inotherMacariini. lostin somespeciesofChiasmia. geoTmheetrtiydmipnansatrlucotrurgean(se.go.,fKMeancnaerlii&niEagrgeertsy,pi1c9a3l3l;y Female (Figs 20, 745-968) Cook & Scoble, 1992). Cook & Scoble (1992) first ThestructureofthefemalegenitaliainthePlatypepla recorded an asymmetrical extension of the ansa, a groupofgeneradiffersfromthatofotherMacariini.In

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