Defense mechanisms in Pyralidae and Choreutidae: Fecal stalactites and escape holes, with remarks about cocoons, camouflage and aposematism

8 Pages·2003·4.8 MB·English

by  Annette Aiello

.
JournaloftheLepidopterists'Society
57(3),2003,168-175
DEFENSE MECHANISMS IN PYRALIDAE AND CHOREUTIDAE: FECAL STALACTITES AND
ESCAPE HOLES, WITH REMARKS ABOUT COCOONS, CAMOUFLAGE AND APOSEMATISM
Annette Aiello
SmithsonianTropicalResearchInstitute, Box2072, Balboa,Ancon,RepublicofPanama,email:[e-mail]
AND
M. Alma Sons
SystematicEntomologyLaboratory, PSI,ARS, USDA, SmithsonianInstitution, P.O. Box37012, NationalMuseumNaturalHistory,
E-517, MRC 168,Washington, D.C.20013-7012,USA,email:[e-mail]
ABSTRACT. Anovelbehavior,linkingfecalpelletswithsilkontheundersideofleaves,toformwhatlooklikeslenderbrownstalactites,is
reportedintwospeciesofPyralidaeandtwospeciesofChoreutidaefromtheRepublicofPanama.Thesefecalstalactites,constructedinasso-
ciationwithescapeholescutinleaves,mayfunctionaslandmarkstolocatethoseholesand/orasdecoysorcamouflage.Wediscussfecalstalac-
tites,camouflageandaposematism,andcocoonsintheselarvaeaspartofamultipledefensesystem.Weprovidethefirstlarvaldescriptionfor
Monoloxisflavicinctalis
Additionalkeywords: Abaera, Brenthia,Monoloxis,Panama,hostplants.
Larvae ofsome lepidopterans livingwithin enclosed Materials and Methods
spaces, e.g., leaf-eating species that roll leaves or cut
and foldleafshelters, remove fecalpellets, attimes ex- The four species were collected as larvae (repre-
plosively, fromtheirhabitations (e.g., Friedlander 1987, sentingvarious stadia), or pupae, on the dates and at
Rawlins 1984, M. Weiss pers. com., AA pers. obs.). the localities (all in the Republic ofPanama) listed in
These behaviors mayhave evolved to avoid pathogens Table 1. Theywere reared in petri dishes or in small
(Rawlins 1984) ortoeliminateolfactorycuesthatmight cages fashioned from petri dishes andwindowscreen-
attract predators or parasitoids (Stamp & Wilkens ing, andplacedinZiploc®bagswith folded, moistened
1993). Contrarytothis, somelarvaeuseandlivewithfe- papertowelstripstoregulatehumidity. Theirbehavior
cal pellets in their habitations, e.g., manypyraloid lar- was observed and recorded daily (with few excep-
vae, includingstoredproductpests (MAS pers. obs.). tions), andshedheadcapsules andpupal exuviaewere
We documentanoveluse offecalpellets bythe lar- collected and mounted. Larvae were preserved by
vae offour moth species. We use "fecula" and "fecal bringing them to a boil in distilled water, then drop-
pellets" to refer to larval excrement, reserving "frass" pingthem into 80% ethanol.
for "The chips or particles cast aside bywood borers" The two Brenthia are not identified to species.
(Frost 1959). The larvae of Monoloxis flavicinctalis Brenthia species-levelidentifications are possible only
(Sepp, [1852]) (Pyralidae: Chrysauginae) (Figs. 2-4), with genitalic dissection ofmales and onlyifthe spec-
Abaera nactalis Walker, [1859] (Pyralidae: Chrysaugi- imenbelongstoaspecies describedbyMeyrickandil-
nae) (Figs. 6, 7), andtwospeciesofBrenthia Clemens, lustratedbyClarke (1969) (V. Beckerin litt).
1860 (Choreutidae: Brenthiinae) chew one or more Adult specimens and exuviae of Maracaijia chlo-
escapeholes neartheblade midvein, thenlinktheirfe- risalis Walker (Aiello lot 1978-45) and Monoloxis
cal pellets using silk to form what look like slender flavicinctalis (Aiello lot 1979-73) are deposited at the
brownstalactites suspendedfromthe undersideofthe National Museum of Natural History (NMNH),
leaf (Figs. 4, 7, 8). In addition, M. flavicinctalis and Smithsonian Institution, Washington, D.C, U.S.A. All
Brenthia sp. 1 are here reported to construct cocoons material relating to the remaining rearings, including
offecula. other specimens ofM.flavicinctalis, andplantvouch-
We discuss the function offecal stalactites and es- ers, areatthe SmithsonianTropical Research Institute
cape holes, briefly address the evolution offecal sta- (STRI), RepublicofPanama.
lactites inBrenthia species, incontrasttothose species In the accounts to follow, lot numbers are those of
that do not construct them, and review other struc- Aiello, andconsistoftheyearplusasequentialnumber.
tures that appearsimilarto fecal stalactites, with simi- Whenmorethanoneindividualisreared, anindividual
lar or dissimilar functions. We also address the host number (#) is appended. Thus "lot 1979-73 #2" refers
plants ofthese four moth species and possible apose- toindividual#2ofthe 73rdlotfortheyear1979. These
matism oftheirlarvae. numbers appear on the labels ofall reared specimens
Volume 57, Number 3 169
Table 1. Collectionanddevelopmentaldata(numberofdaysineachstadium)andoutcomes. Numbersincludedaysspentpreparingfor
moltingorpupation,i.e.,noteating. Finaldateisthedateofeclosion,death,orpreservation,andisnotincludedindurations. Ifcocooncon-
tentswerenotvisible, onlytheduration in thecocoonisgiven. Eclosion dates are forthe morningimmediatelyfollowingadultnocturnal
emergence. Minimumdurations (>)aregivenforthestagescollectedorforstagescutshortbypreservationornaturaldeath. A=indicates
amoltthatmayhaveoccurredonadaywhenobservationswerenotmade.Alowercase"p"nexttoanindividualindicatesitwasparasitized.
Instar Instar Instar Instar Instar Cocoon/ Final
Nameandcollectiondata Lot# Indiv# a b c d e pupa date Outcome
Pyralidae(Chrysauginae):
Monoloxisflavicinctalis,onLacistema
aggregatum
CanalArea,BarroColoradoIsland
Snyder-MolinoTrail-5.9
25May1979,A.Aiello
1979-73 >2 18 >12P 27 Jun Pupadied,discarded
1979-73 >1 12 15P 23 Jun Adult
CanalArea,BarroColoradoIsland
BrokawBidge(offBalboaTrail-10)
25August 1988,A.Aiello&E. Leigh
1988-19 >7 30C 1 Oct Adult
1988-19 >7 1 Sep Larvapreserved
PanamaProvince,Arraijan
LomadelBio
12December2001,A.Aiello
2001-44 >5 39C + 15P 9 Feb Pupadied,pointed
2001-44 >1 13 Dec Larvapreserved
Abaeranactalis,onCordiapanamensis
CanalArea,Summit,Old
GamboaBoad
27June1990,D.Windsor
1990-7 >2 =6 8 9 37C 3 Sep Adult
Choreutidae(Brenthiinae):
Brenthiasp. 1,onCojobarufescens
PanamaProvince,Arraijan
LomadelBio
12January1992,A.Aiello —
1992-5 1 12 Jan Emptycocoon
1992-5 p2 27 Jan Parasitized,wasp
pupadied
1992-5 3 >5 2C+ IIP 30 Jan Adult
1992-5 4 >15C 27 Jan Adult
Brenthiasp.2,onCalatheasp.
CanalArea,BarroColoradoIsland
26August 1993,D.Windsor
1993-70 1 >12C 7 Sep Adult
1993-70 2 >12C 7 Sep Adult
1993-70 3 >12C 7 Sep Adult
Brenthiasp.2?,onCalatheasp.
PanamaProvince,CerroJefe
ConservationTrail
10September1993,A.Aiello,D.Windsor&
Miller
J.
. 1993-73 pi >1 1C + 8P 20 Sep Parasitized,wasp
adult
1993-73 2 >1 11 Sep Larvapreserved
170 Journalofthe Lepidopterists' Society
Figs. 1-4. Monoloxisflavicinctalis (Sepp) (Pyralidae: Chrysauginae), reared on Lacistema aggregation (P. J. Bergius) Rusby (Flacour-
tiaceae),asAiellolot1979-73. 1,Pinnedadult.2,Maturelarva,eating.3,Larva(anteriorportion),movingtouppersurfaceofleaf,throughes-
capehole.4,Larva(posteriorportion),returningtolowersurfaceofleaf,bybackingthroughescapehole. PhotographsbyA.Aiello.
andtheir associatedparts, and correspondto numbers through the hole to the upper surface of the leaf,
on dailydataforms maintainedbyAiello at STRI. which was totally bare (Fig. 3). After a few seconds
without further disturbance, it backed down through
Results the hole into its silk tangle (Fig. 4). Except for these
escape maneuvers, larvae remained within their silk
Monoloxisflavicinctalis (Sepp, [1852]) tangles, extendingoutofthemonlyto eatsurrounding
(Pyralidae: Chrysauginae) leaftissue. In the only instance that new shelter con-
(Figs. 1-4, 9) structionwaswitnessed (lot2001-44#1),thelarvapro-
All sixMonoloxisflavicinctalis larvaewere foundon ducedthe escape hole before anythingelse.
mature leaves ofLacistema aggregation (R Rergius) Molting tookplace within the silk tangles. Cocoons
Rusby (Flacourtiaceae). Table 1 gives collJe.ction and were constructed offecula and silk on the leafor on
developmental data. Ofthese sixindividuals, twowere the container floor. The adults were brown, with or-
preserved as larvae, two died as pupae, and two were ange forewing apices (Fig. 1), and rested with the
reared to adults. In each larva the head was dark red forewings coveringthe hindwings, in abroad, flat, tri-
and the bodywas purplish black, with white markings angle. Plant vouchers for lot 2001-44 are Aiello 1582
(dorsalandlateralonTl andA7, andlateralonA2, A5, and 1635.
and A8-A10) (Fig. 2). The larvae lived on the under- Though the setal pattern ofM.flavicinctalis is typi-
surfaces of leaves, each within a wispy tangle of silk cal of the Chrysauginae, the larvae are distinctively
We
supportedby a series ofthree or four, flexible, brown patterned, an unusual trait within the Pyralidae.
stalactites, severalmm apartalongone sideofthemid- provide the first larval description for the genus, to-
vein, and constructed by linking as many as 50 fecal gether with an illustration (Fig. 9). A pupa was not
pellets and securingthem with silk. Each larva rested available fordescription.
with its headnearasmall, neatlyroundedholelocated Larva(Fig. 9): Length: 18mm (n = 1) (finalinstar). Headwith
reddishbrownplatelets;beigeorwhitebetweenplatelets. Epicra-
to one side ofthe midrib and near a fecal stalactite. nial suture present. White between LI and A3, medially across
When touched, the larva moved rapidly head first headandonfronsdorsaltoFl,oneithersideofepicranialsuture,
Volume 57, Number 3 171
Figs.5-8. AbaeranactalisWalker(Pyralidae:Chrysauginae),rearedonCordiapanamensisL.Riley(Boraginaceae),asAiellolot1990-7.5,
Pinnedadult. 6,Penultimatelarvawithinfeculaandsilktanglebeneathleaf;thelargestdorsal,yellowmarksareonAl. 7, Larvacamouflaged
withinfeculaandsilktangle;darkpatchesarefeedingareas. 8,Larvalshelterback-lighted,showingfecalstalactites,escapehole(arrow),and
scrapedfeedingareas. PhotographsbyCarlHansen.
and ventroposterior to stemmata. Frontoclypeus area ventral to Abaera nactalis Walker, [1859]
Fl,includingclypeusdarkbrown; anteclypeuswhite; labrumyel- (Pyralidae: Chrysauginae)
lowish brown. Adfrontal area light brown. Mandibles yellowish
brown with dark brown margins. Tl-3 and Al-10 integument (Figs. 5-8)
lSiDgh1tloynruAg8o.seP.roWtihtohrasccilcersohtiiezleddwrhiintges,adtatrhkebbraoswensbofetDw2eeonnDT23aanndd The single Abaera nactalis larva (lot 1990-7) was
SD2 alongposteriormargin and extendinglength ofmedial line. foundon a mature leafof Cordiapanamensis L. Riley
XD1,XD2,SD1withsmalldarkbrownpinacula.Tlwithlobesan- (Boraginaceae). Table 1 gives collection and develop-
teriortothoraciclegsandprothorax;2Lsetaebelowandanterior
to spiracle on brown pinaculum. Tl-3 legs with basal segments mental data. The head was patterned with white and
sclerotized dark brown; tarsus white. T2-3 with D1-D2, SD1- darkbrown, andthe bodywas checkered darkbrown,
SD2,andL1-L2onsamepinaculum;SV1andL3withonesetaon pale brown, and bright yellow (Fig. 6). Like M.flavi-
separate pinacula. Tl-3 and Al, light brown ventrally; A2-A10
whiteventrally. A2, A5, andA7withwhiteareas anteriortospira- cinctalis, it lived within a loose, silk tangle supported
cleand SD1 andextendingdorsalto SD1,butventral to D setae. by flexible fecal stalactites, and had an escape hole
wAbhe7ittawele;seopnwiinStaDhc1ualwauhnmidtdeDalarrkeaabnrdjooibwnnei.tnwgAeD1en02boporntimbhaordDii2lyssiedwethsai.et.AeA8w9iwtiphtrhibmwrahroiiwtlney f(Faicges.of6d-i8e)lteharfowuhghenwhwiechdiitstsucroboetdedit.toUntlhiekuepMp.efrlasvuir--
mottlingbetweenSD2, D2,andDl.Al-8withLIandL2present cinctalis, this larva decorated the silk tangle with nu-
on separate pinacula ventral to spiracle; SD1 on a large, brown merous individual fecal pellets, which, in conjunction
pinaculum dorsal to spiracle, except A2, A5, and A7 where the
pinaculum is small, dark brown, triangular; Dl and D2 setae on with the complex markings of the larva, provided
separate small, round, darkbrown pinacula. Al-6 with three SV messy but highly effective camouflage (Fig. 7). Early
setae,A7-8withtwo SVsetae. Al-8withone L3seta. SD2ofAl instars ateonlythetissueoftheleafundersurface,pro-
anteriorto SD1, SD2 ofA2, A3, A4, A6, andA8 anterodorsal to
spiracle; SD2 notpresent on A2, A5, andA7. SD1 pinaculum on ducing extensive, brown scraped patches bounded by
A8 protruding, seta at least 20 times the length ofother setae. the secondaryveins (Fig. 7). The final instar ate areas
oStpihrearcalebdoonmiAn8alatslpeiarsatcltews.icAe9aswiltahrgtehraenedLslsiegthatleyomnosraemdeorpsianlactuhlaan; ofleaf, veins and all.
Dl, D2, and SD1 on separate pinaculum. Prolegs with crochets Portions ofthree shelter-building efforts were ob-
biordinalinacircle. served. The first ofthese new shelter building events
172 Journalofthe Lepidopterists' Society
abdominal abdominal
prothorax mesothorax metathorax segment6 segments8-10
Fig.9. Monoloxisflavicinctalis(Sepp)(Pyralidae:Chrysauginae),setalmapsoflarvabody(above)andhead(below).IllustrationsbyA.Solis.
tookplace 5 July on the original leaf, and consisted of night of2-3 September. It held the wings in a slightly
an escape hole near the midrib, one fecal stalactite, a tentedtriangle.
silktangle sprinkledwithfecula, andtwoscrapedfeed-
ingareas. When theleafbegan to turnblack, the larva Brenthia Clemens, 1860, sp. 1
was prodded onto a fresh leaf, thus precipitating the (Choreutidae: Brenthiinae)
second building episode. Within 30 minutes the larva One larva and three fecula and silk cocoons (lot
had movedtothe underside ofthe leafandchewedan 1992-5) of a small species of Brenthia (6 mm wing
escape hole next to the midrib. During the next 45 span) were found on the undersurface of mature
mm
minutes, it constructed a fecal stalactite (4 long) leaflets ofCojoba (-Pithecellobium) rufescens (Benth.)
nearthe hole. The next morningasecond fecal stalac- Britton & Rose (Fabaceae: Mimosoideae), in sec-
mm
tite (10 long), a silk tangle, and a small feeding ondary growth. Table 1 gives collection and develop-
area had been added. And the day after that a third mental data. The three cocoons were suspendedhori-
stalactite (17 mm long) appeared. The thirdnewshel- zontally among stalactites, within fecula-sprinkled silk
tereffort tookplace 20July, afterthe newlymolted fi- webbing. Cocoon #1 contained onlya cast larval skin.
nal instarwas transferredto a fresh leaf. The larvafirst Cocoon #2 held a pupa, which protruded from its
chewedanewescapehole, thenconstructedseveralfe- housing andwithin which awasp pupa could be seen
cal stalactites and a silk tangle. The following day clearly; the wasp pupa failed to develop, then molded
(21July),itfashionedacocoon-likeshelteroffeculaand andwas discarded.
silkinsteadofmore stalactites, andbegan eatingwhole The only larva (#3) was among fecula-sprinkled
leafratherthan simplyscrapingtheblade surface. webbingandhad madeseveralneatlyroundedholesin
The maturelarvacurledintoan nexttothe midrib the leaflet, permitting rapid passage to the uppersur-
at aleafbase andconstructedacocoon ofsilkandleaf face. Each hole hadaflexible fecal stalactite nexttoit.
material. The finished cocoonwas cream color on the The larva ate the undersurface leaf tissue only pro-
inside, dark brown on the outside, and was covered ducing small scraped patches. Cocoon #4 contained a
with tufts of leaf trichomes. The adult male, brown, healthy pupa. Individuals #3 and #4 yielded adults.
withpowder-blueforewingapices (Fig. 5), eclosedthe Priortoeclosion, pupaeprotrudedfromtheircocoons.
Volume 57, Number 3 173
Adults displayed in their petri dishes. This and the tite were the first items to appear in the shelter con-
next species are among the microlepidopteran "pea- struction sequences, underscoringtheirimportanceto
cock moths" that are seen frequentlyin the Canalwa- larval survival. Unlike the larval behavior of the two
tershed area, performing a raisedwing display (Aiello Brenthia species describedhere, severalotherspecies
& Beckerin prep.) on foliage, sometimes several indi- of Brenthia fashion escape holes, but not stalactites.
vidualsperleaf. Specimen information and previous descriptions of
Brenthia biologyintwodifferentpartsoftheWorlddo
Brenthia Clemens, 1860, sp. 2 not report the building of fecal stalactites. Brenthia
(Choreutidae: Brenthiinae)
pavonacella Clemens, reared by Busck in the United
Table 1 gives collection and developmental data. States (NMNH: specimens and leaf remains), lived
Three cocoons (lot 1993-70) of a larger Brenthia among fecula-dotted silk tangles beneath leaves. In
species (1 cmwingspan,versus 0.7cm)werefoundon Japan, B.japonica Issiki (Arita 1971, Issiki et al. 1975)
the undersurface of Calathea sp. (Marantaceae) and B. pileae Arita (Arita 1971) have been reportedto
leaves. Escape holes, short fecal stalactites, silk tan- fashion escape holes and fecula-laden silk tangles;
gles, and scraped feeding areas on the leafundersur- fig. 230in Issikietal. (1975) shows alarvathathas "es-
face indicated a larval life style similar to that ofthe caped" through its hole. Similar behavior is found in
precedingBrenthia species. However, this specieswas an unrelated, more primitive, microlepidopteran
found on a monocotyledonous plant, and instead of species, Compsistis Meyrick (Elachistidae: Depres-
constructing a cocoon of fecula and silk, it spun a sariinae), whose larvae cut escape holes but do not
white silkcocoon. The cocoonwas composed ofthree construct fecal stalactites, beneath mature leaflets of
parts. The main part, an elongate spindle-shaped Pseudobombyx septenatum (Jacq.) Dugand (Bomba-
structure that housed the pupa, rested suspended caceae) (AA pers. obs. lot 1993-94). Because escape
within a cloud ofwispy silk between two silk sheets; holes appeared before fecal stalactites in shelter con-
the top sheet was flat and had numerous holes, espe- struction andbecausesome species ofBrenthia create
ciallytowardsits margins, andwasanchoredallaround escape holes without fecal stalactites implies that es-
totheleaf; thebottom sheetwascreasedlengthwiseto cape holes mayhave come firstin the evolutionaryse-
form a V-shaped trough, shorter than the top sheet, quence ofthis behavioral pattern. This idea could be
and anchored to it alongits sides. The final larval skin tested by conducting aworldwide phylogenetic study
had been pushed out of one end of the spindle and ofBrenthia.
into the silk cloud. Prior to eclosion the pupae pro- There are several structures in arthropods, includ-
jected from their cocoons. All three cocoons yielded ingin other Lepidoptera, that are reminiscent offecal
adultfemales. And all three moths dashed about their stalactites. The sand pillars constructed by some fid-
petri dishes displaying in the same manner as the dler crabs help them locate theirburrows rapidlyand
smallerspecies. thus avoid predation (Christy 1991, 1995). In Mono-
One oftwo larvae found on CerroJefe (Table 1, lot loxis and Brenthia, fecal stalactites may function sec-
1993-73),alsoontheleavesofCalatheasp.,spunawhite, ondarilyas decoylarvae, andtheyrecallthe "fake"lar-
three-layered cocoon like those oflot 1993-70, and al- vae fabricated by early instar Adelpha hasiloides
mostsurelywasBrenthia,verylikelyBrentliia sp. 2. Fol- (Bates) (Nymphalidae) (Aiello 1984). Fecal stalactites
lowing pupation, it pushed its final larval skin out of remind us ofthe horizontal and more rigid fecal rods
one end of the spindle-shaped cocoon. On the mis- produced by early instars of many Nymphalidae, ei-
taken conviction thatan adultwouldbe obtainedfrom theras continuations ofleafveins oras formations an-
it, theotherlarvawaspreserved. Alas, asmallbraconid choredto the leafmargin. The techniques involvedin
waspemergedfromthecocoon andnoadultswereob- construction offecal rods and stalactites maybe simi-
tainedfrom this rearing. lar, but their functions are quite different. Fecal rods
are used as restingand moltingperches byearliest in-
Discussion
starnymphalids and are thoughttoprovide both cam-
Thelarvalconstructions,i.e., fecalstalactites andes- ouflage and safe haven from ants and otherpredators
cape holes, and associated behavior described above (Machado& Freitas2001). Manynymphalidsenhance
are part of a complex defense system that includes those protections bybarricadingthe base ofthe struc-
camouflage and possibly aposematism. Fecal stalac- ture with loosely attached leaf bits (Muyshondt &
tites are always nearan escape hole, and appearto act Muyshondt 1979) or, inAdelpha spp., clusters offecal
aslandmarkstohelplarvaelocatetheholes andescape pellets (Aiello 1984).
quickly. The escape hole and its accompanying stalac- Though fecula cocoons are not common among
174 Journalofthe Lepidopterists' Society
Lepidoptera, examples are found in several families a shrub or small tree that also is host to an as yet
besides those reported here for Pyralidae and Chore- unidentifiedsexuallydimorphic,wasp-mimickingdiur-
utidae, e.g., Synanthedon spp. (Sesiidae) (Barrett nal moth (Arctiidae: Ctenuchinae) (AA pers. obs. lot
1997), Mimallo amilia (Cramer) (Mimallonidae) (AA 1999-8). The Flacourtiaceae belong to the Violales, a
pers. obs. lots 1985-131, 1987-45, 1990-54, 1997-33, cluster offamilies notable for supporting an array of
2002-27). Feculacocoons mighthelpprotecttiieiroc- aposematic lepidopterans, e.g., Heliconius spp.
cupants from parasitoids and predators, but their ef- (Nymphalidae) on members of the Passifloraceae
fectiveness has not been tested. Among known Bren- (Benson et al. 1976, Trigo2000),Josia draconis Druce
thia, Brentliia sp. 1 is unique, so far, in using fecal (Notodontidae: Dioptinae) on Turnera panarnensis
pellets to construct its cocoon. All other Brenthia Urb. (Tumeraceae) (AApers. obs. lots 1994-37, 1994-
species forwhich we have information spin white silk 39, Miller 1996); Siderone marthesia (Cramer)
cocoons: B. coronigera Meyrick in India (Fletcher (Nymphalidae) on Casearia guianensis (Aubl.) Urb.
1920, NMNH: Rangi specimen), B. japonica (Arita (Flacourtiaceae) (AApers. obs. lots 1990-25, 1996-28,
1971, Issiki et al. 1975), B. pavonacella (NMNH: 2000-37); Zunacetha annulata Guerin (Notodontidae:
Busckspecimens), B. pileae (Arita 1971), andBrenthia Dioptinae) on Hybanthus prnnifolhis (Humb. &
sp. 2 in Panama (this paper). As well, white silk co- Bonpl.) Schulze-Menz (Violaceae) (Wolda & Foster
coons among webbing are found in another member 1978,AApers. obs. lots 1977-25, 1979-26, 1997-11), to
of the Choreutidae, Hemerophila albertiana (Stoll) mention afew.
(AApers. obs. lot2001-39). Itwouldbeachallenge for AposematismtoprotectcamouflagedPyraloidealar-
apredatoryorparasitoidwasp to breach one ofthese vae against chance exposure maybe a more common
multi-layeredsilkcocoons. defense mechanism than has been reported in the lit-
Additionally, our observations indicate that though erature. In additiontothetwochrysauginespeciesdis-
camouflaged,thelarvaeof'M.flavicinctalis andA. nac- cussed above, the first author has reared the larva of
talis may also be aposematic, exhibiting both warning Maracayia chlorisalis Walker (Crambidae: Spilomeli-
coloration and unpalatability (Bowers 1993). It is nae), whose clear cuticle and large, black pinacula
known that the degree of pigmentation in lepi- camouflage it beneath a silk and fecula tangle on the
dopteran larvae tends to correlate positivelywith de- broad, succulent leaves ofits foodplant, an epiphytic
gree ofexposure tovisuallyhuntingpredators (Stamp cactus,Epiphijllwnphi/llanthus (L.) Haw. (Cactaceae)
& Wilkens 1993). Exposed feeders include mimetic, (lot 1978-45); and the aposematic (white, ornamented
camouflaged, orcrypticallypatternedspecies aswellas with black pinacula and yellow supraspiracular
colorful aposematic ones (Bowers 1993, Stamp & blotches) larvaofPalpitaflegia (Cramer) (Crambidae:
Wilkens 1993). The larvae ofhidden feeders tend to Spilomelinae) that eats the leaves of a toxic plant,
be colorless, or they may appear green or brown due Thevetia ahouai (L.) A. DC. (Apocynaceae) (lot 1984-
to their gut contents, orwhite due to fat body, and in 60). The adults ofthe latter two are white, the color
species that extend from their shelters to feed, orthat most conspicuous and therefore most aposematic at
reside in moveable cases, the head and prothorax are night. The evolution ofthe abilityto sequester defen-
pigmented and the rest ofthe body is not (AA pers. sive compounds as larvae and retain them into the
obs.). In contrast to conventional notions that most adult stage has notbeenwell studied (Bowers 1993).
camouflaged pyraloid larvae are watery-looking cater- It is doubtful that Brenthia larvae, being small and
pillars with pale or clear cuticles, M.flavicinctalis and inconspicuous, relyon chemical protection from their
A. nactalis arewell-pigmented. host plants. If any do so, the most promising host
The larva ofA. nactalis is strikingly colored yellow plants, as far as plant secondary compounds are con-
and brown, and well camouflaged within its fecula- cerned, wouldbe the Sapindaceae, which is hostto B.
sprinkledwebbing. As well, it maybe protected from elongata HeppnerintheWestIndies andB. sapindella
chance exposure to predators by chemicals obtained BusckinCuba(Heppner 1985). Anothergroupofsec-
from its food plant, Cordia panarnensis, a short-lived, ondary compound candidates among known Brenthia
secondgrowthtree thatalsohosts sixspeciesofmetal- host plants would be the Fabaceae, which are known
lically colored tortoise beetles: two species of Omo- tosupportanumberofLepidopteraaposematicas lar-
cerus Chevrolat and four of Discomorpha Chevrolat vae and/or adults, i.e., Ormetica sicilia Druce (Arcti-
(Chrysomelidae: Cassidinae) (Windsor et al. 1992). idae) onInga sp. (AApers. obs. lot 1980-44), Utetheisa
The larva of M. flavicinctalis, less well camouflaged ornatrix L. (Arctiidae) on Crotalariacajanifolia Kunth
than that ofA. nactalis, quite likely derives chemical (Fabaceae: Papilionoideae) (Trigo 2000), Melanispixe
protection fromits foodplant, Lacistema aggregation, (Boisduval) (Riodinidae) on Albizia adinocephala
Volume 57, Number 3
(Donn. Sm.) Britton & Rose (AA pers. obs. lots 1988- Benson,W.W„K.S.Brown,Jr.&L.E.Gilbert. 1976. devolu-
30, 1991-5). In addition to Brenthia sp. 1 in Panama, tionofplantsandherbivores:passionflowerbutterflies. Evolu-
tion29:659-680.
the Fabaceae are hosts to at least two other Brenthia Bowers, M. D. 1993. Aposematic caterpillars: life-styles of the
species: B. albipunctata Aritaon Spatholobus compar waminglycoloredandunpalatable, pp. 331-371. In Stamp. N.
Craib in Thailand (Arita 1987) and B. pavonacella on Ear.y&coCnastsreayi,ntTsoMn.f(oerdasg.i),ngC.atCehrpaipllmaarn: e&coHlaolgli,caNleawndYoervko.lution-
Desmodium sp. in the U.S.A. (NMNH: Busck speci- Christy, H. 1991. Comparativestudiesofreproductivebehavior
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mens from Great Falls, Virginia, 17 and 18 July 1913; inmantisshrimpsandfiddlercrabs.Am.Zool.31:329-337.
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Inga vera Willd. at Lares, Puerto Rico, where ". . . in Clarke,J.F.G. 1969. CatalogofthetypespecimensofMicrolepi-
November 1931, Mr. Francisco Sein found them dopterainthe British Museum (Natural History) describedby
abundant, feedingon the underside ofthe leaves. ." EdwardMeyrick.Vol.VI,pp.24—39. London: British Museum
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two mechanical constructions that mayenhance larval Friedlander,T. 1987(88). Taxonomy,phylogeny, andbiogeogra-
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Acknowledgments MachCao.d,oL,tdG..,O&saAk.a.L.23F7repipt.as. 2001. Larval defence against ant
OurthanksgotoJonLewis(SystematicEntomologyLaboratoiy, predation in the butterfly Smyrna blomfddia. Ecol. Entomol.
USDA) forhis generous helpwith the literatureandwith locating 26:436^39.
specimens;toVitorO.Becker(ResearchAssociate,Universidadede Miller,J. S. 1996. Phylogenyofthe NeotropicalmothtribeJosi-
Brasilia, Brasil) for identifying the moths; to Donald Windsor ini(Notodontidae: Dioptinae):ahiddencaseofMullerianmim-
C(aSrTlRIH)anfsorencolfolrecpthiongtotghreaAp.hinnagcttahleisfoarnmderC;omtposWiaslttiesrlAairvealel,oa(nDdukteo Muysihcroy.ndZoto,l.JrJ..,LAi.nn&.AS.ocM.u1y1s8h:1o-n4d5t.. 1979. Notesonthelifecy-
University Medical Center), Naoki Takebayashi (Duke University cle andnatural historyofbutterflies ofEl Salvador. IIIc. His-
DepartmentofBiologicalSciences),andK.T. Park(KangweonNa- torisodiusandCoeaacheronta (Nymphalidae:Coloburinae).J.
tionalUniversity,Chuncheon,Korea)forkindlytranslatingtheJapa- Lepid. Soc.33:112-123.
nNeMsNeHt)ex,t;DtaoviMdarR.cSEmpisttheiannd(DSteepvaertLminegnatfeolfterSy(sStyesmtaetmiactiBcioElnotgoy-, RawlWihnese,lJe.r,E.Qu1e9n8t4i.nM&ycBolapchkaweglyl,inMLeerpeiddiotphte(redas,.)p,pF.u3ng8u2s^-2i3ns.ecInt
mology Laboratoiy, USDA), Bernard Landry (Museum d'histoire relationships.ColombiaUniversityPress, NewYork.
naturelle, Geneva, Switzerland), Priya Davidar (Pondicherry Uni- Stamp,N.E.&R.T.Wilkens. 1993. Onthecrypticsideoflife:be-
versity, India), Jean-Philippe Puyravaud, and an anonymous re- ingunapparenttoenemiesandtheconsequences forforaging
viewerfortheirhelpful reviews ofthe manuscript; andto Martha and growth ofcaterpillars, pp. 283-330. In Stamp, N. E. &
Weiss, Georgetown University, forinsights intothe opposite larval Casey,T.M.(eds.),Caterpillar:ecologicalandevolutionarycon-
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