Contribution towards the study of the Pyralinae (Pyralidae): Historical review, morphology, and nomenclature

10 Pages·1999·4.9 MB·English

by  M Alma Solis

Journal
of
The Lepidopterists' Society
Volume 53 1999 Number 1
journaloftheLepidopterists'Society
53(1),1999,1-10
CONTRIBUTION TOWARDS THE STUDYOFTHE PYRALINAE (PYRALIDAE):
HISTORICAL REVIEW, MORPHOLOGY, AND NOMENCLATURE
M. Alma Sous
SystematicEntomologyLaboratory,PSI,AgriculturalResearchService, U.S.DepartmentofAgriculture,
c/oNat. Mus. Nat. Hist., MRC 168,Washington,DC20560-0168USA
AND
Michael Shaffer
EntomologyDepartment,TheNaturalHistoryMuseum,Cromwell Road, LondonSW75BDEngland
ABSTRACT. ThemonophylyofthePyralinaeandthetwotribes, PyraliniandEndotrichini,isreviewedbasedonananalysisofpreviously
usedmorphologicalcharactersoftheadultandlarva. Characterspreviouslyusedtodefinethesegroups areplesiomorphic(i.e.,theyarenot
valid)orhighlyhomoplastic(i.e.,theyarenotreliable)tosupportthetaxaasmonophyletic,orboth.Wedescribethemalegenitaliaandpresent
characterstosupportthe monophylyofthe Endotrichini,butthePyraliniislikelyaparaphyletictaxon. Larvalcharactersdidnotprovideevi-
dencetosupportorrejectmonophylyforeithergroup.Basedonmalegenitalicmorphologywereassigngenera,andmakeadditionsorchanges
withinthesetaxainrecentlypublishedchecklists.IntheNeotropicalfauna:PerforadixSeinistransferredtothePyraustinaeandisanewsyn-
onymofSufetulaWalker;alectotypeisdesignatedforPerforadixsacchariSein;MicronixAmselistransferredfromthePyralinaetotheCram-
binae;andMicromastraSchausandTahogaDyar,revisedstatus,remaininthePyralinae. Inaddition,SufetulapygmaeaHampson,presentlyin
theCrambidae,istransferredtotheNoctuidae:Nolapygmaea Hampson(Nolinae),newcombination. IntheAustralianfaunaMacnaWalker
istransferredfromthePyralinaetotheChrysauginae. AlistofthesubfamiliesandtribesofthePyralidaeworldwideandofthespeciesofthe
PyralinioftheWesternHemisphereareincluded.
Additionalkeywords: Endotrichini,Pyralini, Neotropics,Australia,larvalmorphology.
Within the Pyraloidea, the Pyralinae are a large onlyin Asia andAfrica. The Pyralini include 118 gen-
group of about 900 species that are more diverse in era,withthevast majorityofthe species distributedin
AfricaandAsiathanintheWestern Hemisphere. This Africa and Asia, although some occurworldwide. The
subfamilyincludes theworldwide stored-productpest two tribes have been defined bytwo states ofahind-
species Pyralisfarinalis Linnaeus, also known as the wingvenational character (Endotrichini = Rs anasto-
meal moth. Acomplete studytoinvestigatethe mono- mosedwith Sc+Rj; Pyralini = Rsnotanastomosedwith
phyly of the Pyralinae has never been conducted. Sc+Rx) andtheyhave been shiftedbetween tribal and
However, Solis and Mitter(1992)proposedacharacter subfamilial rank based on the importance placed on
to define the Pyralinae and hypothesized it to be the this characterbyvarious authors (e.g., Ragonot 1891,
sister group to the phycitine + epipaschiine clade Hampson 1896, Whalley 1961, Minet 1982). We dis-
(Table 1). In this paperwe integrate previous findings pute thevalidityofthe use ofthe hindwingvenational
in the Pyralinae and our observations to facilitate fu- character at suprageneric levels. We also explore the
ture studies onthese moths. literature and investigate the morphology of larval
Presently, there are two tribes in the Pyralinae, the stage as anindependentcharacterset.
Endotrichini and Pyralini (Table 1). The Endotrichini Recent publication oftwo checklists (Shaffer et al.
includes 7genera, Endotricha Zellerbeingthe largest 1996, Shaffer & Solis 1995) of the Pyralini and En-
genuswithabout70species. Basedonourmorpholog- dotrichini of Australia and the Neotropics, and the
icalandlabeldataobservations,thetribeisdistributed previous publication ofthe checklist ofthe Pyraloidea
Journalofthe Lepidopterists' Society
Table 1. Higher classification ofthe Pyralidae; current tribal Adult and larval characters ofthe other subfamilies
names in use, although most tribes have not been shown to be ofthePyralidae, Chrysauginae, Galleriinae, Phycitinae,
monophyletie.
Epipaschiinae, were used for outgroup comparison
PyralidaeLatreille purposesbasedonaphylogeneticanalysisbySolis and
ChrysauginaeLederer Mitter (1992) (Table 1).
GalleriinaeZeller
Calleriini Historical ReviewandAnalysis of
Megarthridiini
Previously Used Characters
Tirathabini
Cacotherapiini Meyrick (1890) first brought the character of the
Pyralinae Latreille veins 7 [=Rs] and8 [=Sc+Rj inthehindwingtoatten-
Pyralini
Endotrichini tion, and since Ragonot (1891) the Pyralini and En-
EpipaschiinaeMeyrick dotrichini have been separated and definedprimarily
PhycitinaeZeller
by two character states of the hindwing venation: in
Cryptoblabini
Phycitini the Pyralini Rs and Sc+Rj approach each other (Fig.
Cabniini 5), butdonotanastomose; inthe Endotrichinithetwo
Anerastiini
veins anastomose for at least part oftheir length (Fig.
Peoriini
6). Weproposethatthis characteris notreliableinthe
separation or definition oftaxa at suprageneric levels
of North America north of Mexico (Munroe 1983) in the Pyralinae and in the following historical review
have laid the groundwork for studies on the systemat- we use italics to drawattention to these two character
ics ofthe Pyraloidea. A large number oftaxon trans- states in descriptions.
fers, and even misplaced taxa between superfamilies, Meyrick (1890) included Endotricha in the Pyrali-
have been documented in the recently published nae. He included the Pyralinae and Epipaschiinae in
checklists. We herein explain how assignments in re- the Pyralididae anddidnotrecognizethemasseparate
cent Australian (Shaffer et al. 1996) and Neotropical subfamilies. Meyrick defined (and spelled) the Pyrali-
(Shaffer & Solis 1995) checklists were made based on didae as follows:
our observations on male genitalic morphology and "Ocelli present, often concealed by scales. Tongue well-devel-
larvalmorphology. Wealsolistcorrections tothetribal oped, orsometimesobsolete. Maxillarypalpiwell-developed, or
and subfamilial headings of the recently published sracruerleylyrufduirmceanttearayt.baFsoer,eswoimnegtsiwmietshsviemipnle1,us4uaalnldys5hocrltolsyeolyroabp--
checklists, and list additions or changes made since proximatedatbaseoroftenstalked,7and8outof9. Hindwings
theirpublication. withoutdefinedpectenofhairsonlowermarginofcell,veins4
and5closelyapproximatedatbaseorfrom apointorstalked, 7
Materials and Methods [=Rs]outof6nearoriginorrarelyseparatebutcloselyapproxi-
mated, free or sometimes anastomosing with 8 [^Sc+Rj"
The collections at The Natural History Museum (Meyrick1890:433) [italicsours].
(BMNH), London, England; the National Museumof He used head and wing venational characters occur-
Natural History (USNM), Washington, D.C., USA; ringin othergroups (i.e., plesiomorphiccharacters) to
the Cornell University Collection (CU), Ithaca, New define the Pyralididae. The presence ofthe ocelli and
York, USA; andZoologische Staatssammlung (ZSBS), maxilliarypalpi areplesiomorphic characters, thepro-
Munich, Germanywere studiedtodeterminetaxanot boscis is secondarily lost, the forewing venation is
includedin recentlypublishedchecklists. Type speci- highly variable at lower taxonomic levels (and hence
menswereexaminedanddissectedwhennecessary. If notusedbymostlaterworkers), andthelackofahind-
the type specimen could not be located, the original wingpectenisplesiomorphic.
descriptions and genitalic illustrations were used to Ragonot (1891) was the first to separate the Pyrali-
place the species generically. Genitalia slides ofnon- nae and Endotrichinaebasedontheveins 7 [=Rs] and
typespecimenswereprepared, studied, andcompared 8 [=Sc+Rj ofthe hindwingin akey: "Nervures 7 et 8
whentypespecimenswerenotavailable, orwhentype soudees aux inferieures, tres rarement separees"
specimenswere notin suitable condition forstudy. ["Veins 7 and 8 fused in the hindwings"] keyed to the
Larvae from alcohol collections ofthe USNM and ChrysauginaeandEndotrichiinaeand"Nervures 7et8
BMNH ofEndotrichaflammealis (Denis & Schiffer- separees" ["Veins 7and8separate"] keyedtothePyra-
muller), Pyralisfarinalis Linnaeus, Aglossa caprealis lidinae (Ragonot 1891:446).
Hiibner, and Herculia psammioxantha Dyarwere ex- Hampson (1896) included only the Phycitinae,
aminedwithastereomicroscopetoverifytheliterature Chrysauginae,Epipaschiinae,Endotrichiinae,andPyral-
onlarval morphology. inae in his concept ofthe Pyralidae and used some of
.
Volume53, Number 1
uncus
uncus arm
tegumen
gnathos
valva
Fig. 1. PyralisfarinalisLinnaeus,typespecies;characteristicmalegenitaliaofPyralini. Fig.2. PyralisfarinalisLinnaeus,aedeagus.
the same plesiomorphic characters as Meyrick (1890). Hindwingwithmediannervurenon-pectinate;vein 7[=Rs]anasto-
But he groupedthe Epipaschiinae, Endotrichinae and mosingwith8"[italicsours].
Pyralinaebasedonthefollowingtwocharacters: The Endotrichini ofWhalley (1961) was described as
"The three subfamilies ofthe Pyralidae, the Epipaschiinae, En- follows:
dotrichinae, and Pyralinae, of which a classification is here at- "Proboscis well developed, maxillary palps present. Chaetosema
tempted,allbelongtothegroupofPyralidaewhichhavethemedian present. Forewingwithvein R.stalkedwith R andR Hindwing
nervure ofthe hindwing non-pectinate on upperside, and vein 7 4 3.
with medianveinnon-pectinate.VeinRsanastomosingwith Sc+R"
[=Rs] oftheforewing stalked with 8 [=Sc+Rj]" (Hampson 1896)
[italicsours].
[italicsours].
The lack ofa hindwing pecten is plesiomorphic, and Munroe and Shaffer (1980) revisedthree large gen-
he usedthe same character (stalkedveins 7 [=Rs] and era in the Pyralini (Pyralinae). Their definition ofthe
8 [=Sc+Rj) of Ragonot to define the Endotrichinae Pyralinae is basically a combination of Hampsons
andChrysauginae. (1896) definition ofthe Endotrichinae and Pyralinae
The relationship anddefinitionofthe endotrichines from a key with Whalleys (1961) rank oftribes. The
and pyralines was not addressed again until Whalley PyralinaeofHampson (1896)was describedas follows:
(1961), who did not provide characters to define the "Proboscisusuallywell-developed;maxillarypalpipresentandusually
Pyralinae or the Pyralini. To define the Endotrichini filiform. Forewingwithvein 7 stalkedwith 8,9. Hindwingwith the
he used the same plesiomorphic characters used by mediannervurenon-pectinate;vein8[=Sc+R,]/re<?"[italicsours].
Hampson (1896), with the exception ofthe presence The Pyralinae ofMunroe and Shaffer (1980) was de-
ofthe chaetosema, but this state is plesiomorphic as scribedas follows:
well. The Endotrichinae ofHampson (1896) was de- "ThethreegeneracannowbedefinedasbelongingtothePyralinae
scribed as follows: fromthefollowingcharacters:chaetosemapresent;maxillarypalpus
present;probosciswelldeveloped;forewingwithR5stalkedwithR4
"FPorroebwoisncigswwietlhlvdeeivnel7opsetadl;kemdaxwiiltlhary8,p9alp(i7 aprbesseenntt;inbuHielnddesclaesnidesr). andR3;hindwingwithRsnotanastomosedwith Se+flj( Pyralini),or
Journalofthe Lepidopterists' Society
uncus
uncus arm
tegumen
gnathos
transtilla
va
saccular process
Fig. 3. Enchtrichaflammealis (Denis & Schiffermiiller),typespeciescharacteristicmalegenitaliaofEndotrichini. Fig.4. Endotricha
flamrnealis (Denis&Schiffermiiller),aedeagus.
Rsanastomosedwith Sc+R (Endotrichini);medianveinnon-pecti- been observed in representatives of the New World
l
nated"[italicsours]. Cacotherapiini (Galleriinae) and some genera in the
Whalley (1961) recognized the problem with the Crambinae (e.g., Pseudoschoenobius Fernald). Based
definitionofthehindwingcharacterthatseparatedthe on our observations ofthe distribution of this hind-
two groups: "In several cases they have been said to wing characterin other groups within the Pyraloidea,
anastomose where, as close examination shows, they distributionofthehindwingcharacterwithinthePyral-
merely run very close together (e.g., Rostripalpus inae,andlackofconcurrencewiththecharactersofthe
Hampson)." The lackofanastomosis ofRs and Sc+Rj malegenitalia,weproposethatthehindwingvenational
varies in other groups within the Pyraloidea besides characterisnotreliableintheseparationordefinitionof
the Pyralini, andithasbeendocumentedashighlyho- taxaatsupragenericlevelsinthe Pyralinae.
moplasious at the generic level among the genera of Minet(1982, 1985)wasthefirsttomaintainthatthe
the Pococera complex of the Epipaschiinae (Solis Pyralinae were paraphyletic because characters used
1993) and at the species level (Shaffer & Solis 1994). bypast workers were plesiomorphic. He stated: 'Les
Other groups where the majority ofthe taxa lack the Pyralinae semblentparaphyletiques parrapport ades
anastomosis ofRs and Sc+R butwhere there are ex- taxa tels que les Endotrichinae, les Chrysauginae ou
1
amples where the two veins barely anastomose have les Epipaschiinae (dontils ne differentqueparunen-
Volume 53, Number 1
Results
Adult genitalic morphology. Previous authors
(Whalley1961, 1963, Munroe& Shaffer 1980) didnot
use genitalic characters to define the Pyralinae,
Pyralini, or Endotrichini, although theyused genitalic
morphologyat the species level fortheir studies. Solis
and Mitter (1992) proposed a characterofthe female
genitalia (corpusbursaebarelyextendingcephaladbe-
yond segment 7) to support the monophylyofthe Py-
ralinae.This study,however,wasbasedonasmallsam-
ple size, acharacterthatremains untested.
Pyralini (Figs. 1, 2, 7):
Description: Male: Uncus samewidth throughout orless nar-
rowthanthebase,flatorspatulate,ventrallywithspineclustersab-
sent;uncusarmslaterallynotlargeandearlike;downcurvedgnathos
witharms stronglydeveloped,withwell-developedmedial,narrow
spiketerminatinginasmalldorsallycurvedhook;tegumenstrongly
sclerotized; vinculum well developed; juxta simple, rarely heavily
sclerotized,spinycatena(baso-medialportionofanellus)presentor
absentorlaterallysclerotized,andheavilyspined,sometimesanellus
reflexed with heavy sclerotization; transtilla absent or, ifpresent,
membranous, rarelywell developed andheavily sclerotized; valva
variableinshape,samewidthtoapexormorenarrowdistally,basal
andcostal process absentorpresent, ifpresentwell developedor
not,withoutsaccularprocess,ventralsurfaceofvalvabearinghair-
likesetaenotarrangedinradiatingrows,costalsetaeabsent;vesica
of aedeagus with or without clusters of spinelike cornuti, vesica
Fig. 5. Wingvenation(idealized); arrowindicateslackofanas- sometimesspined, reflexedwithheavysclerotization,orwithbroad
tomosis. Fig.6. Wingvenation(idealized);arrowindicatesanas- bandsofsclerotizationitsentirelength. Female: Segment8andas-
tomosis. sociatedmembraneseithershort,orlongandextensible;apophyses
anterioresandposterioreslong,stoutorslender;ostialendofductus
bursae membranous, with small, well-sclerotized to large, heavily
sclerotizedcompactpouchespresentorabsent;ductusbursaelong
semble de caracteres plesiomorphes: palpes maxil- andnarrowwithareasofminutespinesimmediatelybelowantrum
laires bien developpes, ailes anterieures sans ecailles orothersclerotizedareas;corpusbursaelarge,signumvariable,ab-
sent,orifpresentfromscobinatepatches,usuallywithinsinglelarge
herissees, etc.)," ["The Pyralinae appear to be para- area,tolongandspinelike.
phyletic in comparison with taxa such as the En-
dotrichinae, theChrysauginaeorthe Epipaschiinae (in Endotrichini (Figs. 3, 4, 8):
that they share a group ofplesiomorphic characters: Diagnosis: Uncusbroadestatapex; uncus arms laterallylarge,
maxillary palpi well developed, forewings without earlike;gnathosmediallybroad,spatulate,platelike.
Description: Male: Uncusbroadestatapex,ventrallywithspine
raisedscales, etc.)."] buthe retainedpyralines anden- clusterspresentorabsent (uncusprocess ofWhalley); uncus arms
dotrichines at the subfamilylevel. Whalley (1963), in laterally large, earlike [socii ofWhalley; socii, according to Klots
his study ofEndotricha, found that the retention of (1956) arepairedprocessesoneithersideofthebaseoftheuncus;
these structures arenotsocii, but the mostlateralelements ofthe
Ragonot's conceptofthe Endotrichinae as asubfamily uncus arms ofthe Pyralidae (Solis & Mitter 1992)]; downcurved
was notwarrantedandproposedthe Endotrichinae as gnathosarmsstronglyorweaklydeveloped,usuallywithawell-de-
a tribe ofthe Pyralinae. He did not offer a reason or velopedmedial,broad,spatulate,andupturnedcentralplate;weakly
sclerotizedtegumen;vinculumwelldeveloped;juxtasimple,some-
characters to support this conclusion. Solis and Mitter
times with spiny manica; transtillapresent, usuallyheavily sclero-
(1992) agreed with Minet that previous characters tized;valvausuallysamewidthtoapex, maybearbasalprocessand
usedto define thetwotaxawere plesiomorphic states, saccularprocess;ventralsurfacebearinghairlikesetaeinrowspoint-
ing toward base ofvalva; prominent, reflexed, sometimes spear-
buttheytreatedtheendotrichines asatribewithinthe shaped costal setae may be present arising from costa near apex:
Pyralinae accordingto Whalley (1961) because Minet aedeaguswithvesicabearingsticklikeorclublikecornutusvaryingin
(1985), inhis studyofthe tympanal organs, presented shapeandlength.Femalegenitalia: Segment8andassociatedmem-
branes long and extensible; apophyses anteriores and posteriores
no apomorphies for the Pyralinae, Pyralini, or En- longandslender;ostialendofductusbursaeminutelyspined,with-
dotrichini. Solis and Mitter (1992) proposedonechar- out pouches; antrum sclerotized; ductus bursae short, minutely
acterofthefemalegenitaliaas asynapomorphyforthe spined, or very long and membranous; corpus bursae large with
signumscobinate.
Pyralinae, but proposed none for the Pyralini or En- Althoughwecanprovide synapomorphiesinstatesoftheuncus
dotrichini. andgnathos (seediagnosisabove) forthe Endotrichini in the male
Journalofthe Lepidopterists' Society
Fig.7. PyralisfarinalisLinnaeus,typespecies;characteristicfemalegenitaliaofPyralini. Fig.8. Endotrichaflammealis(Denis&Schif-
fermiiller),typespecies;characteristicfemalegenitaliaofEndotrichini.
Volume 53, Number 1
genitalia,wewere unable to find synapomorphies forthe Pyralini mastra and Taboga remain in the Pyralinae. Table 2is
wherethestatesareeithersharedwiththeEndotrichiniorwithre- acomplete fist ofthe Pyralinae (Pyralini) ofthe West-
vlaartieadbsluebfaatmliolwieesr.tTahxeongoemniitcalleivceclhsaorracptleerssioomfotrhpehiPcy,rail.ie.n,afeoaurnedeiitnhaelrl ern Hemisphere (Munroe 1983, Shaffer& Solis 1995).
relatedsubfamilies.WeprovideadescriptionofthegenitaliaforEn- Sein (1930) placed Perforadix sacchari Sein, com-
dotrichiniandPyralinibecauseaconceptbasedonthemorphology monlyknown as the Sugarcane root caterpillar, in the
ofthemalegenitaliawasusedtoassigntaxaintheEndotrichiniand
PyraliniintheAustralianandNeotropicalchecklists. Endotrichinae. This species is a major pest of sugar
Larval morphology. Comparison of the caterpillars of En- caneinPuerto Rico andnearbyislands. Sein (1930) il-
dotrichaflammealis with Pyralisfarinalis, Aglossa caprealis, and lustratedthe morphologyofall fife stages in great de-
Herculia psammioxantha (with caterpillars ofothersubfamilies of
the Pyralidaeasoutgroups)didnotresultinanyapomorphiechar- tail and describedits biologyand methods ofcontrol.
acters to supportthe monophylyofthe Pyralinae, Pyralini, orEn- The author failed to designate types or even fist type
dotrichini. Historically,the Hasenfuss(1960)conceptofthePyrali- specimens, butwefoundseven specimens eachwith a
nae consisted ofpresent-day galleriines, pyralines, and phycitines
(hedidnotincludechrysaugines orepipaschiines inhis study); he smalllabel"PR./Sein"andaredlabel"Cotype/Cornell
consideredEndotrichaasapyraline. U. No. 6087" at Cornell University. According to Sein
W
ThelarvaeofEndotrichahaveapinaculumringonSD1ofA9,a (1930), T M. Forbes, who was at Cornell Univer-
synapomorphy forthe Pyralidae (the plesiomorphic state, the ab-
sence ofthe pinaculum ring on SD1 ofA9, occurs in the Cram- sityatthetime, identifiedthematerialandpresumably
bidae). Insum,wefoundthatE.flammealis larvaelacktheunique he also labelled the material as cotypes. We designate
characters assigned to other subfamilies and have the same ple- one specimen (male) as the lectotype and the other 6
siomorphicsetalcharacterstatesassignedtothelarvaeofthePyral-
inae. The Epipaschiinae andPyralinaeboth lackapinaculum ring specimens as paralectotypes (material in poor condi-
onanyothersegmentotherthanA9(incomparisontothepresence tion, abdomens are missing), and they are labelled as
ofapinaculumringonT2ofthePhycitinae,T3oftheChrysauginae, such in the collection at Cornell University. We stud-
andAl ofthe Galleriinae;presenceineachsegmentisthederived
state, although thepinaculum has been secondarilylostin several iedadditional material collectedby Sein, identifiedby
generaand/orspeciesofeachsubfamily). BasedonworkbyHasen- H. G. Dyar, as stated by Sein (1930), and dissections
fuss(1960) andAllyson (1977)the EpipaschiinaeandPyralinaeare by Carl Heinrich at the USNM and found that Per-
separatedfromeachotherbythedistancebetweentheventralsetae
onA7andA9. IntheEpipaschiinaethetwoventralsetaearecloser foradix is a synonym of Sufetula Walker, new syn-
togetheronA7thanthoseonA9andinthePyralinaethetwoven- onymy, in the Pyraustinae. We discoveredthat based
tralsetaeonA7andA9areequidistant(theplesiomorphiccondition onthe morphologyofthe tympanal organs (i.e., cram-
sharedbyothersubfamiliesofthePyralidae).
bid "open" tympanal organs with apraecinctorium) it
Taxonomic placement ofgenera. Recently, gen- belongs in the Crambidae. Based on the external and
erafromsoutheastAsiaandAustraliapreviouslyplaced genitalicmorphologyaftercomparisonwithotherspe-
in the Endotrichinaewith anastomosed Rs and Sc+Rj ciesinthe genus,includingthetypespecies,itbelongs
in the hindwing, but with genitalic characteristics of in the genus Sufetela Walker. It is interesting to note
the Pyralini were transferred to the Pyralini in the that P. sacchari was originally identified for Sein by
Australian checklist (Shaffer et al. 1996) based on the H. G. Dyar as Sufetula grumalis Schaus, a species
genitalia morphology. Those genera transferred from presently placed in Sufetula (Munroe, 1995:76). We
the Endotrichinae to the Pyralinae were based on the also examined another species, Sufetula pygmaea
genitalia morphology: Gauna Walker, Curena Walker, Hampson, and found it does not belong in the Pyra-
Arescoptera Walker, Scenedra Meyrick, Tanyethira loidea, butin the Noctuidae (Nolinae): Nola pygmaea
Turner, ScenidiopsisTurner,PerisseretmaWarren, and (Hampson), new combination.
Ferula Mabille. Amsel (1956) describedMicronix nivalis in the En-
Accordingtothe definitionbasedongenitalicmor- dotrichinae. Nothing is known about the biology of
phology given above, there are no known species of thisVenezuelan species. Wewere unable to locatethe
Endotrichini in the Western Hemisphere, but four type, a male, but Amselprovided aphotograph ofthe
genera,Perforadix Sein,MicronixAmsel, Micromastra adult andpoorline drawings ofthe male genitaliaand
Schaus, andTaboga Dyarhavebeenhistoricallyplaced wings. Thehoodeduncus andthecostalprocess ofthe
within the Endotrichinae due to the anastomosing of male genitalia characteristic ofcrambines areveryev-
Rs and Sc+R in the hindwing. Perforadix, Micronix, ident in the illustration; therefore, we transfer this
1
and Micromastra were inadvertently excluded from monotypic genus tothe Crambinae. Althoughwe can-
the Neotropical Pyraloidea checklist (Shaffer & Solis notdetermineitsplacementwithintheCrambinae,we
1995). Taboga was includedin the Neotropical check- suggestthatitbelongsin thetribe Crambini.
list, but needed to have its position in the Pyralinae Schaus (1940) placed Micromastra isoldalis in the
verified. We found that Perforadix belongs in the Endotrichinae. Nothing is known about the biologyof
Pyraustinae and Micronix belongs in the Crambinae, thisPuertoRicanspecies. Dyar(1914)describedTaboga
bothherebytransferred, and, ofthe four, onlyMicro- inis in the Endotrichinae. The type series is from
Journalofthe Lepidopterists' Society
Table2. Pyralini(Pyralidae:Pyralinae)oftheWestern Hemisphere
AglossaLatreille, [1796] HypsopygiaHubner, [1825]
EuclitaHubner, [1825] costalis(Fabricius, 1775)(Phalaena)
AgriopeRagonot, 1894 fimbrialis([Denis&Schiffermuller], l775)(Pyralis)
acallalisDyar, 1908 purpurana (Thunberg, 1784)(Tortrnf)
baba Dyar, 1914 hyllalis (Walker, l859)(Pyralis)
cacamica (Dyar, 1913) (Pyralis) MapetaWalker, 1863
caprealis(Hubner, [1800-09])(Pyralis) Homalochroa Lederer, 1863
capreolatusHaworth, 1809 cynosura Druce, 1895
cuprealisHubner, [1825],missp. omphephora Dyar, 1914
aenalis(Costa, 1836)(Pyralis) schausi Druce, 1895
domalisGuenee, 1854 xanthomelasWalker, 1863
incultella (Walker, [1866])(Acrobasis) aestivalis(Lederer, l863)(Homalochroa)
enthealis(Hulst, 1886)(Tetralopha) divisa (Boisduval, 1870)(Pyralopsis)
cuprialisHeinrich, 1931, missp. MicromastraSchaus, 1940
costiferalis (Walker, 1886) (Pyralis) isoldalviSchaus, 1940
costigeralis(Walker,[1865](Pyralis),preocc.(Walker, 1862) NeodavisiaBames& McDunnough, 1914
cuprina(Zeller, 1872)(Pyralis) Davisia Barnes&McDunnough, 1913,preocc(Del
disciferalis(Dyar, 1908) (Pyralis) guercio, 1909[Hemiptera])
electalisHulst, 1866 melusinaFerguson, Blanchard,&Knudson, 1984
furva Heinrich, 1931 singularis(Barnes& McDunnough, 1913)(Davisia)
gigantalis Barnes&Benjamin, 1925 OcrasaWalker, [1866]
oculalisHampson, 1906 Parasopia Moschler, 1890
pinguinalis (Linnaeus, l758)(Pyralis) nostralis (Guenee, 1854)(Pyralis)
marmorella (Geoffroy, 1785)(Tinea) helenensis(Wollaston, l879)(Pyralis)
marmoratella (Villers, 1789)(Tmea) tenuis (Butler, l880)(Pyralis)
pinguiculatus (Haworth, 1809)(Crambus) dissimilalis(Moschler, 1890)(Para.sopia)
guicciardiiConstantinio, 1922 sordidalis (Barnes&McDunnough, l9l3)(Herculia)
ArispeRagonot, 1891 psammioxantha(Dyar, \917)(Herculia)
UscodysDyar, 1909 venezuelensis(Amsel, 1956)(Herculia)
cestalis (Hulst, l886)(Anerastia) tripartitalis(Herrich-Schaffer, 1871)(Asopia)
atalis(Dyar, lQ08)(Uscodys) PseudasopiaGrote, 1873
concretalisRagonot, 1891 cohortalis(Grote, 1878)(Asopza)
ovalisRagonot, 1891 florencealis(Blackmore, 1920)(//ercu/ja)
CatocrocisRagonot, 1891 intermedialis (Walker, 1862)(Pyralis)
CatacrocisRagonot, 1892,missp. sodalis(Walker, 1869)(Pt/rafts)
lithosialisRagonot, 1891 squamealisGrote, 1873
DolichomiaRagonot, 1891 phoezalis (Dyar, 1908)(Herculia)
amoenalis (Moschler, 1882) (Asopia) PyralisLinnaeus, 1758
isidora(Meyrick, l936)(Pyralis) AletesRafinesque, 1815,nom.nud.
binodulalis(Zeller, l872)(Asopia) Ceropsina Rafineques, 1815,nom.nud.
craspedalis(Hampson, 1906)(Tegulifera) SpyrellaRafinesque, 1815,repl. name
datames(Druce, l900)(Pyralis) AsopiaTreitschke, 1828
decetialis (Druce, l900)(Pyralis) SacatiaWalker, 1863
graafialis(Snellen, 1875)(Asopia) EutrichodesWarren, 1891
impurpuratalis (Dognin, 1910)(Pi/rafo) farinalisLinnaeus, 1758
nigrapuncta(Kaye, 1901)(P!/raZis) domesticalis(Zeller, 1847)(Asopw)
olinalis (Guenee, l854)(Pyralis) fraterna Butler, 1879
trentonalis(Lederer, 1863)(Asopia) nianihotalis.- Matsumura, 1900(notGuenee, 1854)
hinwnialis(Zeller, 1872)(Asopia) meridionalis Schmidt, 1934
infimbrialis(Dyar, 1908)(Herculia) orientalisAmsel, 1961
phanerostola (Hampson, \9\l)(Paractenia) nianihotalisGuenee, 1854
planalis(Grote, 1880)(Asopi<z) vetusalisWalker, [1859]
enniculalis (Hulst, 1886)(Asopia) gerontesalisWalker, [1859]
occidentalis(Hulst, 1886)(Asopia) laudatella (Walker, l863)(Sacatia)
plumbeoprunalis(Hampson, !9l7)(Herculia) despectalisWalker, [1866]
resectalis(Lederer, 1863)(Asopw) miseralisWalker, [1866]
thymetusalis(Walker, 1859)(Botys)Newcombination achatina Butler, 1877
devialis(Grote, l875)(Asopia) haematinalis(Saalrmiller, 1880)(Asopia)
vernaculalis(Berg, 1874)(Asopia) gerontialis(Meyrick, 1888)(Asopia),emend.
HerculiaWalker, 1859 centripunctalis(Gaede, \916)(Endotricha)
BuzalaWalker, 1863 pupalisStrand, 1919
CisseWalker, 1863 compsobathraMeyrick, 1932
BejudaWalker, [1866] TabogaDyar, 1914
BleoneRagonot, 1890 inis Dyar, 1914
HerculeaAmsel, 1956[index],missp.
tabidalis (Warren, \89\)(Pyralis)
Volume 53, Number 1
Panama. Studyofthe genitaliaofthe type series atthe ioral characters, although caution is suggested con-
USNMofbothofthesespeciesconfirmtheirplacement cerning the latter due to the convergent nature ofbe-
withinthe Pyralini, andnotinthe Endotrichini. havioralcharacters. Aphylogeneticanalysis ofthe gen-
In the Australian checklist (Shaffer et al. 1996:173) eraofthePyraliniwouldbethefirstattempttotestthe
the headings of the Endotrichini and Pyralini were paraphylyofthe Pyraliniwith respect to the Endotri-
mislabeled and difficult to change atproofstage. The chini, with the possibility that results may also invali-
Endotrichinae shouldhavebeen titledthe Endotrich- date the traditional tribal concept. Such a study may
ini andplaced underthe heading ofthe Pyralinae. In also provide characters to support the monophyly of
addition, the genus Macna Walker was inadvertently the Pyralinae.
includedinthe Pyralinae (Pyralini), butitshouldhave
Acknowledgments
been placed in the Chrysauginae. In the Neotropical
checklist (Shaffer& Solis 1995:80) the Pyralini should IthaWcea,tNhaenwkfYoorrkt,heiUrShAo,spiftoarlihteyl:pE.whRi.leHosetbuedkyein,gCtohreneSleliUnnimvaetresriitayl,
havebeenincludedasasubheadingunderPyralinaeto andA.Hausmann,ZoologischeStaatssammlung, Munich,Germany,
indicate the tribal placement ofthe genera found in whilestudyingtheAmselmaterial.WealsothankLindaLawrence,
theWestern Hemisphere. scientific illustrator, Systematic Entomology Laboratory, U.S. De-
partmentofAgriculture,forthefigures.Thefollowingprovidedre-
views or helpful comments: Vitor O. Becker, Marianne Horak,
Discussion DouglasFerguson,JamesPakaluk,andDavidSmith.
The Pyraloidea, one ofthe larger superfamilies of Literature Cited
the Lepidoptera, has over 15,000 described species, Allyson,S. 1977. ThestudyofsomeNorthAmericanlarvaeofthe
yet much remains to be done in taxonomy, and, more genus Tetralopha Zeller (Lepidoptera: Pyralidae: Epipaschi-
so, with the phylogenetic relationships. A taxonomic Amseiln,aeH)..CGa.n.1E9n56t.omoMli.cr1o0l9e:p3i2d9o-p3te3r6a.Venezolana. Bol. Entomol.
studyusuallybegins with a checklist or a catalogue of Ven. 10:(l-2), 1-336(1956),(3-4),pi. I-CX(1957).
described species as an inventory to document those Dyar, H. G. 1914. Reportonthe Lepidopteraofthe Smithsonian
that have alreadybeen described. A checklist mayre- bMiuosl.og4ic7a:l13s9u-r3v5e0y.ofthe Panama Canal Zone. Proc. U.S. Nat.
fine the placement oftaxa and can clearly marktaxon Hampson, G. 1896. On the classification ofthree subfamilies of
transfers, aswellasprovide otherinformation, such as moths ofthe family Pyralidae: the Epipaschiinae, Endotrichi-
misplaced taxa. By definition, a checklist or catalog Hasennafeusasn,dIP.yr1a9l6in0a.e.DTireanLsa.rvEanltsoymsotle.maStoick.dLeornZduonns.le4r:.4A5c1-a5d5e0m.ie
does not adequately state or discuss the taxonomic Verlag,Berlin. 263pp.
problems solvedorthose that remaintobe solved. Klots, A. B. 1956. Lepidoptera, pp. 97-110 In S. LTuxen (ed.),
We have described the morphological reasons for Taxonomist'sglossaryofgenitaliaininsects.EjnarMunksgaard,
Copenhagen.
theplacementoftaxainthePyraliniorEndotrichiniin Meyrick,E. 1890. OntheclassificationofthePyralidinaoftheEu-
two checklists (Shaffer et al. 1996, Shaffer & Solis ropeanfauna.Trans.Entomol.Soc.London.(PartIH):429-492.
1995). We have also summarized the taxonomic and Minet,J. 1982. Les Pyraloideaetleursprincipalesdivisionssyste-
matiques. Bull. Soc. Entomol. France86:262-280.
phylogenetic status ofthe two tribes included in the . 1985. Etude morphologique et phylogenetique des or-
Pyralinae. The genitaliaofthe Endotrichiniareclearly ganestympaniquedesPyraloidea.2,Pyralidae,Cambidae,pre-
miere partie (Lepidoptera Glossata). Ann. Soc. Entomol.
different from those ofthe Pyralini, but authors have
France(N.S.)21(l):69-86.
dealt only with a few genera in both taxa and, as we Munroe, E. G. 1983. Pyraloidea, pp. 67-85. In Ronald W.
have shown, have usedthe sameplesiomorphic orho- Hodges,etal., ChecklistoftheLepidopteraofAmericanorth
ofMexico.UniversityPress,Cambridge.
moplasious characters since Meyrick (1890) to define
higherleveltaxa.Weretainthetwotribesintheclassi- Neo.tr1o9p9i5c.alPyrLaeupsitdionpatee,rpap.ch5e3c-k7li9s.t:InPJa.rtB.2.HepAspsnoecri.atAitolnasfoorf
ficationsystemforthesakeofstabilityandretentionof NeotropicalLepidoptera,Gainesville,Florida.
Munroe, E. &M. Shaffer. 1980. ArevisionofVitessidia Roths-
character information, but acknowledge that the child & Jordan and Vitessa Moore (Lepidoptera: Pyralidae).
Pyraliniislikelyaparaphyleticgroup. Moreover, apre- Bull. Brit. Mus. (Nat. Hist.)Entomol.39(4):241-360.
liminarystudyofanindependentcharacterset,thelar- Ragonot, E. L. 1891. EssaisurlaclassificationdesPyralites.Ann.
val stage, provides no obvious synapomorphies forthe Soc. Entomol. France (1890):435-488 (January), 489-546
(May).
Pyralinae orthe Pyralini. Schaus, W. 1940. Insects of Porto Rico and the Virgin Islands,
Our observations ofthe genitalia and larvae ofthe mothsofthefamiliesGeometridaeandPyralididae. NewYork
Pyralini and Endotrichini are made with the expecta- Acad. Sci. 12:291-417.
Sein Jr., F. 1930. The SugarCane Root caterpillarandothernew
tion that they may prove useful in a future phyloge- rootpestsinPuertoRico. Dept.Agric.PortoRico.14:167-191.
netic studyofthe Pyralini generathatincludes an en- Shaffer,J. C. & M. A. Sous.J. 1994. PyralidaeofAldabraAtoll2.
tiresuiteofadultandimmaturecharacters. Anyfuture AElpidpaabsrcahiainndaea:noLtehpeirpafsrcohmiaBugernk.inna.,, Fwaistoh,aWensetwAsfpreicciae.sEfnrtoom-
study should also include pupal and perhaps behav- mol. Scand.25:311-320.
10 Journalofthe Lepidopterists' Society
Shaffer, M. & M. A. Sous. 1995. Pyralinae,pp. 80-81. InJ. B. Sous,M.A.&C. Mitter. 1992. Reviewandphylogeneticanalysis
Heppner(eel.),AtlasofNeotropicalLepidopterachecklist:Part ofthesubfamiliesofthe Pyralidae(.sen.sustricto)(Lepidoptera:
2. Association for Neotropical Lepidoptera, Gainesville, Pyraloidea). Syst. Entomol. 17:79-90.
Florida. Whalley,P.E.S. 1961. Achangeofstatusandaredefinitionofthe
Shaffer, M., E. S. Nielsen & M. Horak. 1996. Pyralidae, subfamilyEndotrichinae (Lep. Pyralidae)withthedescription
pp. 164-199. In E. S. Nielsen, E. D. Edwards & T. V. Ransi, ofanewgenus.Ann. Mag. Nat. Hist.3(13):733-736.
ChecklistoftheLepidopteraofAustralia. MonographsonAus- . 1963. A revision ofthe world species ofthe genus En-
tralian Lepidoptera, CSIRO Publications, East Melbourne. dotricha (Zeller) (Lepidoptera: Pyralidae). Bull. Brit. Mus.
529pp. (Nat. Hist.)Entomol. 13(2):395-454.
Sous, M. A. 1993. A phylogenetic analysis and reclassification of
thegeneraofthePococeracomplex. NewYorkEntomol.Soc. Receivedforpublication 15 February 1998; revisedandaccepted
J.
101(l):l-83. 29September1998.
MANUSCRIPT REVIEWERS, 1998
The merit ofa scientificjournal depends on the quality ofits reviewers as well as its authors, but the former are
usuallyunknown to the readers ofthe published articles. TheJournal relied on the expertise of60 reviewers last
yeartoprovide66evaluations ofmanuscripts. ItiswithmuchgratitudethattheJournalacknowledgesthe services
ofthe people listed below, from whom manuscript reviews were received in 1998. Those who reviewed two or
more manuscripts are denotedbyasterisks.
Adamski, David,Washington, DC Landry, Bernard, Aylmer, QB
Aiello, Annette, Balboa, Panama Landry,Jean Francois, Ottawa, ON
Arnold, Richard, Pleasant Hill, CA Lederhouse, Robert, Lockport, NY
Austin, George, LasVegas, NV MacNeill, Donald, San Francisco, CA
Ballmer, Greg, Riverside, CA McCabe, Timothy, Albany, NY
Boggs, Carol, Stanford, CA *Metzler, Eric, Columbus, OH
Borkin, Susan, Milwaukee, WI Miller, Lee, Sarasota, FL
Bowers, Deane, Boulder, CO Miller, William, SaintPaul, MN
Brower, Andrew, Corvallis, OR Peigler, Richard, San Antonio, TX
DC MD
Brown, John, Washington, Poole, Robert, Rockville,
Brown, Richard, Mississippi State, MI *Powell, Jerry, Berkeley CA
Burns, John,Washington, DC Prowell, Dorothy, Baton Rouge, LA
Calvert,William, Austin, TX Pyle, Robert, Grays River, WA
Carde, Ring, Riverside, CA Rindge, Frederick, NewYork, NY
Cho, Soowon, Berkeley, CA Robbins, Robert, Washington, DC
Collins, Michael, NevadaCity, CA Scholtens, Brian, Charleston, SC
Covell, Charles, Louisville, KY Scriber, Mark, East Lansing, MI
Drummond, Boyce, Florissant, CO *Shapiro, Arthur, Davis, CA
Dussourd, David, Conway AR Shaw, Scott, Laramie,WY
Eichlin, Tom, Sacramento, CA Shepard, Jon, Nelson, BC
Emmel, Tom, Gainesville, FL *Shuey, John, Indianapolis, IN
Epstein, Marc, Washington, DC Sims, Steve, Maryland Heights, MS
Ferguson, Douglas,Washington, DC Singer, Michael, Austin, TX
Fink, Linda, Sweet Briar,VA Strand, Michael, Madison, WI
Friedlander, Timothy, North Potomac, MD Swengel, Ann, Baraboo, WI
Gall, Lawrence, NewHaven, CT *Tuck, Kevin, London, England
Goldstein, Paul, NewYork, NY Webster, Reginald, Fredericton, NB
Hawkins, Bradford, Irvine, CA Weller, Susan, SaintPaul, MN
Jenkins, Dale, Sarasota, FL West, David, Blacksburg,VA
CO NY
Kelley, Scott, Boulder, Williams, Ernest, Clinton,