Bulletin of the Natural History Museum Entomology

508 Pages·2001·32.3 MB·English

by  Natural History Museum (London)

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
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© 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
-
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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