Adelpha erotia erotia form "Lerna" (Nymphalidae): Exploring a corner of the puzzle

2006·3.4 MB·

Volume 60, Number 4 181
JournaloftheLepidopterists'Society
60(4),2006, 181-188
ADELPHA EROTIA EROTIA FORM "LERNA" (NYMPHALIDAE):
EXPLORINGA CORNER OFTHE PUZZLE
AnnetteAiello
SmithsonianTropicalResearchInstitute, Box0843-03092Balboa,Ancon,RepublicofPanama email:[email protected]
ABSTRACT.ThelarvaeandpupaofAdelphaerotiaerotia"lema"aredescribedandfigured,anditisconcludedthatthetaxonbelongstothe
A.mesentinaspecies-group. Cecropialongipesisreportedasthelarvalfoodplant. RelationshipsamongthesixA.mesentinagroupspeciesare
explored.
Additionalkeywords: lifehistory,Panama,Bombacaceae,Cecropiaceae,Malvaceae, Urticaceae
Introduction other limenitidine genera, Adelpha is extremely
The immature stages of any group of Lepidoptera, homogeneous morphologically, both in wing pattern
and the affiliations among their larval food plants, can and genitalia, thus providing few reliable characters for
provide valuable insights into species identities and phylogenetic analysis. Character systems other than
alliances, and they can prove especially valuable in adult morphology thus are essential for achieving a
better understanding ofAdelpha evolution and extant
helping untangle a perplexing group, such as the
speciose Neotropical genusAdelpha Hiibner. species relationships. The time is ripe for a molecular
Adelpha dorsalwingpatterns are confusing, and that study of Adelpha. As well, the immatures and food
has led to numerous misidentifications in widely plants offer a wealth of taxonomic information, but
consulted works, and hence to increasing confusion. detailed and reliable knowledge ofthem is available for
Thus, formore than acentury,Adelpha defiedattempts only about 25% of known Adelpha species (Miiller
to sortoutits more than 350 publishedtaxa, thoughthe 1886, Moss 1933, Aiello 1984, 1991, Otero & Aiello
best didtry (Godman & Salvia 1879-1901, Fruhstorfer 1996, Freitas et al. 2001, Willmott 2003b, & references
1907, Forbes unpublished manuscript). Butdifficulties therein). For that reason, it is important for all who
ofidentificationarenowlargelybehindus, thankstothe posses newinformation on Adelpha biologv, to publish
recent revision ofAdelpha by Willmott (2003b), who it. Even a small bit ofnew information can provide a
has taken us a giant leap forward in our understanding valuable clue to species relationships, or offer support
ofthis bewilderinggroup. After evaluatingall available fororagainstsuspectedones, andthus alsoshedlighton
type material and all published names, Willmott the reliability of other characters. Here, I present
concluded that Adelpha comprises 85 species and 209 informationaboutthelarvae, pupa, andlarvalfoodplant
taxa. His detailed descriptions and color photographs ofAdelpha e. erotia form "lerna" (Hewitson, 1847), in
makeitpossible atlasttoidentifyvirtuallyanyspecimen the Republic of Panama, and point out possible
with nearcertainty. alliances within the A. mesentina group, to which it
The task remains, to devise a satisfying classification belongs.
forthe group, one that reflects natural relationships. A Materials and Methods
major obstacle to understanding relationships within
Adelpha is that the dorsal wing patterns are deceptive; Thirteen individuals were collected, 11 as larvae
species ofsimilarappearanceare notnecessarilyclosely (representing various stadia) and two as pupae, on die
related, and closely related species do not necessarily dates, at the localities (all in Panama), under the lot
resemble each other. numbers, and bv the collectors listed in the appendix.
There is substantial evidence that dorsal wing Ofthese 13 individuals, one was preserved as a larva,
patterns are mimetic, perhaps through Miillerian five died as larvae and were preserved, one was
mimicrybasedondifficultyofcapture (Mallet & Singer preserved as a pupa, and six were reared to adults.
1987, Srygley 1994) orthrough Batesian mimicry(Aiello Preservations were made bybringingthe larva orpupa
1984, Prudicetal. 2002). Thoughventralwingpatterns to a boil in distilled water, then dropping it into 80%
do provide some reliable characters useful in species ethanol. Adultswere frozen, then pinned andspread.
identification, they also are subject to some variation Reared individuals were housed in petri dishes until
directly resulting from selection on the dorsal wing fourth or early fifth stadium, then transferred to small
pattern (Willmottpersonalcommunication). Like many cages fashioned from petri dishes and window
182 Journalofthe Lepjdopterists' Society
screening, and placed in Ziploc® bags with folded, the Urticaceae.
moistened paper towel strips to regulate humidity.
Results
Pupae were suspended in the same type of cages, by
taping their support silk to the covers. All reared All 11 larvaewerefoundon Cecropia longipes Pittier,
individuals were maintained in an air-conditioned lab. 1917 (Urticaceae) (plantvouchersAiello 1588, 1612), in
Behaviors were observed and recorded daily (with few Parque Natural Metropolitano (PNM), on the Pacific
exceptions), and shed head capsules and pupal exuviae side ofthe isthmus. Though C.peltata L., 1759, is more
were collectedand mounted. commonin PNM,noevidenceofAdelpha wasfoundon
Lotnumbers forrearedindividualsconsistoftheyear it, and one larva (lot 2000-11) to which C. peltata was
plus a sequential number. When more than one offered, ate some, thenproduced liquid feculaand died
individual was reared, an individual number (#) is the next day. On the Atlantic side of the isthmus, C.
appended. Thus "lot 2000-20#2" referstoindividual#2 insignis Liebm., 1851, the only Cecropia accessible
ofdie 20thlotfortheyear2000. These numbers appear from the Fort Sherman canopy crane, yielded no
on the labels of all reared specimens and their Adelpha. ThoughC.obtusifolia BertoL, 1840, ispresent
associated parts, and correspond to numbers on daily in PNM, and, C. longipes, C. obtusifolia, and C. peltata
data forms maintained by Aiello at the Smithsonian are foundat Fort Sherman, dreywere outside the areas
Tropical Research Institute (STRI), Republic of accessible from the canopy cranes, and thus could not
Panama. All material relating to these rearings, be searched.
includingplantvouchers, is at STRI. Egg. Notseen.
The first individual reared (lot 1996-24) was inpFoisristtioInnsstMaDr.1,HPeIa,dPb2r,oLwIn,,AwFi1t,hsAeFt2a,esFelt,iCnIb,eiAgle,oSrI,grSa2v,pSi3n,aScSuIl,a
identified by Keith Willmott, from a photograph andSS2,andwith3setaeoneachhalfoflabrum. Bodypalegreenish
showing dorsal and ventral views ofthe adult (Fig. 1), brown, with beige pinacula and bands of tiny beige spots diat
resemblepinaculabutlacksetae-
and is alluded to on page 174 ofhis Adelpha revision Second Insta—r. Headbrown,denselyda—ppledwithpale,broadly
(Willmott2003b). conicalchalazae eachwithat—erminal seta represen—tingthreesize
First instar setal terminology follows Stehr (1987). groups. The—principalchalazae largestandplumpest arearranged
infourseries 7posterior,4medial,2anterior, 1parafrontal. Among
these, ml and al are the most prominent. The spaces among the
principalsare filledbyafewmedium-sizedandmanysmallchalazae.
Thelongestsetaearefoundon thesmalle—stchalazae, locatedo—nthe
lowerportionofthehead. Allplain setae i.e.,setinpinacula are
confinedtothe mouthparts. Bodyyellowocherabove,brownonthe
sides,withshort,yellowocherscoliandgraychalazaeandpinacula.
Third Instar. Head upper fourth, dark brown, with ocher
chalazae;lowertiiree-fourthsgray,withsmallsalmon-coloredchalazae
andstillsmallergrayones;principalchalazaearrangedinfourseries—
7posterior,4medial,3anterior, 1parafrontal(minuteorabsent);the
basesofthelargestlumpyduetominutechalazae. Bodypaleocherto
salmon-brown dorsally, and darkgrayto darkbrown laterally, those
ground colors mostly obscured by densely set, gray pinacula and
chalazae;scolistubbv,ochertosalmon-brown,arrangedassub-dorsal
andsupra-spiracularonT2,T3,andA2-A7,andasonlysub-dorsalon
ASandA10;spiraclesdarkbrown,almostblack.
Fourth Instar. Head nowpale, with brow—npits and numerous
tinychalazaesurroundingtheprincipalchalazae otherwisesimilarin
form andarrangementtothethirdinstar. Bodylikethirdinstar,but
Fig. 1,Adelphaerotiaerotia "lema" adult, dorsal (left),ven- widi a more pronounced 2-toned appearance, i.e., the sides ofthe
tral(right),lot 1996-24. thoraxmuchdarkerdiandierestofthebody.
— — Fifth Instar (Figs 2 top left, top middle, and bottom; and 4
Head chalazae positions second instar on do not bottomleft). Headcream-colortopalebeige,withflat-bottomedpits
ofthe same color, numerous setae set in minute chalazae, brown-
correspond to first instar setal positions, thus, chalaza marked frons, and dark brown mandibles; chalazae white, widi
and scolus terminology (Fig. 2 top) was devised and conspicuous black-tips, and arranged in four series: 7 posterior, 6
developed by Aiello and Willmott over the course of medial.4anterior(alanda3largerthana2anda4),and1parafrontal.
Body widi thoraxyellowocher, mottled with red dorsally, chestnut
previouspublications (Aiello 1984,Willmott2003b) and brown laterally; abdomen yellow ocher with red mottling dorsally,
incorrespondencebetweenusduringthewritingofthis cream-colorlaterally;simple,widelyspacedsetaearrangedasseveral
paper. rings per body segment; chalazae white, ranging from tinyto long;
subdorsalscoli(dorsalofStehr1987)(onT2-AS)large(excepttinyon
Plant classification follows Stevens (2001 onwards).
Al),white,with severaltiersofblack-tipped, black-basedbranches;
Note that Bombacaceae, Malvaceae, Sterculiaceae, supraspiracularscoli(onT2—A7)similartosubdorsals,buttinyand2
Tiliaceae, and several odier families are now combined or3branched;subspiracularscolipale,tiny;gradingfromsessileand
forked on anterior segments, to subsessile and 3—4 branched on
as die Malvaceae, and that Cecropiaceae is included in
Volume 60, Number4 183
Fig. 2 Top, fifth instarheadcapsules:Adelphaerotiaerotia "lerna" (lot2000-29#1) frontview (left) andlateralview(middled
Adelphaserpacelerio (lot 1984-10#2), lateralview (right); bottom, fifth instarhabitus:Adelpha e. erotia "lema," (lot2000-29#31.
Chalazae: a= anterior, m = medial,p =posterior,pf=parafrontal.
posteriorsegments;caudalscolussimple,anteriorlycurved;spiracles shimmering gold reflection on the dorsum ofthe thorax, from die
pinkishbeigewithdarkbrownborders. innercurveofaT2middorsalprojectionthroughtheinnercurveofa
Abeautifullarva,whosewhitescoliwithblacktipsandjoints, and A2middorsalhook. TheA2middorsalhookhuge,fairlystraight,\ridi
numerous white chalazae viewed against the yellow body give it a awider,slightlycurvedapex,farsurpassingdiebluntlvroundedtipof
frostedappearance(Fig.4bottomleft). theT2middorsalprojection,andreachingalmosttoheadlevel. Head
In mature larvae, abdominal segment 2 has a large, rounded, "horns"small, rounded,verysimilarto,butnarrowerdiandioseofA.
middorsal hump that corresponds to the huge A2 hook of the lycoriasmelanthe(Bates, 1864). Pupationsilkchestnutbrown. Bodi
impendingpupa. field-collectedpupae,werefoundsuspendedbeneathleaves.
Larval behavior. First instars constructed fecula and silk rods Developmentandeclosiontimes. Seeappendix.
anchored to the leaf lobe margin, i.e., they were not leaf vein
extensions,andrestedheadoutwardonthem betweenfeedingbouts POSITION WITHINADELPHA
andduringmolting. Theserodswerelengthenedgradually,untilthey
reachedseveraltimesthebodylength ofthelarva. Secondthrough The 2-toned, uniform-scoli larvaanddie "hugehook"
fourthinstarsaddedtinyleafbitstothebaseoftheirfecalrods, and pupa ofA. erotia erotia "lema" indicate diat tliis taxon
continuedtorestonthemasbefore. Firstthroughthirdinstarfeeding
damagewascharacterizedbytinyholes alloverthe leaf, and fourth belongs to dieA. mesentina group (Aiello 19S41,which,
and fifth instars ate whole leaf. Following the molt to the final so far comprises six species as outlined bv Willmott
stadium, larvae abandoned their fecal rods entirely, and when (2003b, as phylaca group) [=group II ofAiello (19S4.
disturbedassumedaheaddown.T1-A2arched,A7-A10upposition.
Pupa (Fig. 3). Entirelydarkbronzybrown exceptforan areaof 1991)], and includesA. mesentina (Cramer, 1777), the
184 Journalofthe Lepidopterists' Society
type species ofthe genus. The final instars ofthe four
A. mesentina group taxa reared in Panama (Fig. 4) also
share similar head morphology; they have 7 posterior
and 4 anterior chalazae, whereas mostAdelpha have 6
and 2 respectively (see Key). Neitherpreserved larvae
nor head capsules of the two Brazilian species were
available forexamination.
Willmott (2003a, 2003b) tentatively includes A.
pollina Fruhstorfer, 1915 in the mesentina group,
though he suspects that it is misplaced there. In his
revision (Willmott, 2003b) he places Adelpha
hesterbergi Willmott & Hall, 1999 in the A. capucinus
group, whereas in his cladograms (Willmott 2003a) it
appears loosely allied with the mesentina group (as
"phijlaca group"), but is not commented on in the text.
The immature stages and larval food plant(s) are
unknown for either taxon, and their logical affiliations
remain to be discovered.
Final stadium larvae of A. e. erotia "lerna" and A.
lycorias melanthe share a number ofcharacteristics not
reported forotherAdelpha species; both have afrosted
appearance due to the contrastingdark tips and branch
bases of their pale scoli and to the numerous tiny
chalazae that further ornament the bodv; and in both,
the A2 subdorsal scolus terminates in a whorl of Fig. 3,Adelphaerotia erotia "lerna" pupa, lateralhabitus and
subequalbranches. dorsalheadandthorax(lot 1996-24).
By contrast, final instar A. phylaca pseudaethalia
Hall, 1938 and A. messana messana (C. & B. Felder, Those unique features aside, thepupae ofA. e. erotia
"lerna" andA. lycorias melanthe resemble one another,
1867) have uniformly brownish scoli and chalazae, set
and differ from A. phylaca pseudaethalia and A. m.
amongplain setae; and one ofthe terminal branches of
messana in havingthe A2 "hook" apex slightlywidened
the A2 subdorsal scolus is elongate and upwardly and truncate, instead oftapering to a rounded tip; and
inclined. Theyalsoshare adark, elliptical, lateral patch
the "head horns" small and rounded, instead of
across the junction of abdominal segments 4 and 5, a
outwardly curved points, or, as in A. mesentina andA.
feature not present in either A. e. erotia "lerna" or A. thesprotia, outwardlycurved andleaf-like (Moss 1933).
lycorias melanthe.
Though the illustrations in Moss (1933) are sufficiently
Moss (1933) does not comment on larval scolus form
& detailed to comment on some characteristics of the
or color pattern inA. mesentina orA. thesprotia (C.
latter two taxa, examination of preserved pupae or
B. Felder, 1867), but does note diat the twowere "...so exuviaewouldbe necessarytoverifythe form ofdieA2
similar...in larva and pupa that no special observations
"hook."
were made." However, his illustrations are sufficiently
Combining published information on the group
clearto showthat the larvae ofthose two species share
(Miiller 1886, Moss 1933, Aiello 1984, 1991) and the
die relativelyunomamented body and dark contrasting present report, we now have descriptions ofthe larvae
elliptical spot with A. phijlaca pseudaethalia andA. m.
andpupae ofat least one subspecies foreach ofthe six,
messana.
well-accepted members of the A. mesentina group.
theThAe2p"uhpoaoko"fAn.eea.rleryotsitara"ilgehrtn,a"anisddissutripnactsisvienginthhaeviTn2g Based on those descriptions, A. e. erotia "lerna" andA.
lycorias melanthe appear to form a subgroup, and the
projection, whereas in each of the other five A.
other four taxa another. Within the second subgroup,
mesentina group species (Fig. 4 shows the four
and based upon the form ofthe pupal "head horns,"A.
Panamanian species) it is curved and directed towards phylaca pseudaethalia appears to pair with A. m.
dietipoftheT2projection,withwhichitforms anearly messana, andA. mesentina withA. thesprotia.
closedcircle. InA. e. erotia "lerna," theT2projectionis The intra-group relationships suggested by the
short and broadly rounded, whereas in die other five
immature stages of the mesentina group are rather
taxait is drawn outto aroundedpoint.
Volume 60, Number4 185
Fig.4,Toprow:Adelpha messana messana final instar, adult,pupa]exuviae (lateral,ventral) (alllot 19S9-22#2): Secondrow:A.
phylacapseudaethalia finalinstar, adult, pupa(lateral,ventral) (alllot 1983-78);Third row:A. hjcorias melanthefinalinstar, adult,
pupa(lateral,ventral) (all 1983-8#3); Bottom row:A. e. erotia "lerna" finalinstarandadult (lot2000-20#5),pupalexuviae (.lateral,
ventral) (lot1996-24).
186 Journalofthe Lepidopterists' Sixiiety
Table 1. Larva] foodplant genera and families reported forAdelpha phylaca Group taxa. forspecimens with identifications
verified. Plantclassification follows Stevens (2001 onwards). Divisionsreflectpossiblespeciesaffinities.
AdelphaTaxon Order Rosales Ordeb Malvales Reference
Ulmaceae Urticaceae Malvacea
A. lycoriasmelanthe Trema Aiello 19S4(lot83-S),asmelanthe;Aiellolot03-14
A. lycoriaslycorias Cecropia Miiller18S6,asisis
A. lycoriaslycorias Coussapoa Miiller1S86,asisis
A. lycoriaslycorias Pourouma Miiller1886,asisis
A.erotiaerotia "lema" Cecropia Aiellolots98-45,00-20,02-29
A. mesentina Pourouma Moss 1933
A. thesprotia Pourouma jmbax Moss 1933,asdelphicola
A. phylacapseudaethalia Cecropia Aiello 1984(lots81-70,82-39,82-40,83-78),asphylaca
aethalia;Aiellolot00-27
A. messana messana Luehea Aiello 1990(lot89-22),asixialeucas
different from thoseobtainedbythe cladisticanalysisof onward), Urticaceae (including Cecropia) and
Willmott (2003a), which relies strongly on adult Ulmaceae, together with Cannabaceae and Moraceae,
characters. It is tobe hopedthat molecularstudies will form awell-supported clade. Furthermore, within the
provide both a clearer concept of intra-group Malvaceae, which comprise 9 subfamilies, the two that
relationships and an assessment of the relative concern us here, Malvoideae and Bombacoideae, form
importance ofimmature andadult charactersets. theirownwell-supportedclade. Thus, theA. mesentina
group taxaare restrictedtotwo small plantclades.
LARVAL FOOD PLANTS OF THEADELPHA
Two of the three plant families just discussed, have
MESENTINA GROUP
been reported as larvalhosts in anotherAdelpha group,
Though three plant families have been reported for theA. sei-pa group: A. celerio on Ochroma (Malvaceae)
members of the A. mesentina group (Table 1), the (lot 1982-41)\mdCecropia (Coleylot 15inAiello 1984);
Urticaceae clearly dominate these records, and at least and an unidentified Adelpha taxon on Heliocarpus
one member ofeach of the three A. mesentina group (Malvaceae) (lots 1982-75, 1983-68, 1985-122).
species-pairs has been reported on that plant family. Whetherlarval hostplant switches and expansions have
The dominance ofUrticaceae may prove greater once played an important role in Adelplia speciation cannot
additional records (DeVries 1986, 1987; Janzen & be determined until we have clearer knowledge of
Hallwachs 2004) of A. lycorias melanthe and A. m. species relationships within Adelpha, and more
messana on Cecropia are verified by Keith Willmott. complete information on host plant associations
Within the Rosales, as outlined by Stevens (2001 throughoutthe ranges ofAdelpha taxa.
Keyto PanamanianAdelpha Mesentina-GROUP SPECIES BASED ON LARVAE AND PUPAE
p = posterior. medial, a = anterior
la. Larvaheadwith6posteriorchalazae,4medialchalazae (in.serpn-Groupml islongandslender,andm2-m4arelow,roundedbumps),
and 2 anteriorchalazae (see fig. 2); frons bordered by a stripe ofsmooth relativelypit-less cuticle; body usuallywith A2 subdorsal scolus
noticeably different in form and/or size from rest; tiiorax not significantly different in color or pattern from rest ofbody; pupa A2 dorsal
projectionnotmassiveandnotformingacirclewithorsuqiassingtheT2dorsalprojection non-mesentina groups
lb. Larvaheadwith7posteriorchalazae,4medialchalazaeplus2or3tinyintercalatedextras(sometimesnotwellaligned),and4anterior
chalazae;pit-lessstripeabsentorpoorlydefined;mainbodyscolifairlyuniforminshapeandsize;sidesofthoraxconspicuouslydarkerthanrest
ofbody; pupawith ahugeA2 dorsal hookthatcurves inward, nearlvtouchingand formingacirclewiththeT2 dorsalprojection, ornearly
straightandsurpassingdieT2projection 2
2a.Larvaheadchalazaetippedwidiblackordarkbrown;bodymainscolipale,withblackorbrownbranchtipsandbases;bodyalsoclothed
withdenselyset,short,palechalazae;alldieabovecombinetogivethelarvaafrostedappearance;headextramedialchalazaeconspicuousand
usuallywell aligned with principals, forming fairly uniform line; terminal branches ofbody A2 scolus similar to each other in length and
inclination;abdomenwithoutdarklateralpatches;pupaheadhornssmallandrounded. 3
Volume 60, Number4 1ST
2b. Larvahead and bodvscoli not dark-tipped, though largerbranchjunctions ofmain bodyscoli maybe darkish: bodyalsowith short
chalazae, but these not densely set, except perhaps laterally; head extra medial chalazae minute or small and often not well-alignedwith
principals;oneterminalbranchofbodyA2subdorsalscolus noticeablylongerthanrest,andtiltedupward; abdominalsegment5withwell-
defined,darkbrownlateralpatch;pupaheadhornsoutwardlycurved,small,roundedoracutepoints,oroutwardlycursedandleaf-like. 4
3a. Larvaheadchalazaeslenderconical,withsomewhatrounded, conspicuouslyblacktips;pitsindistinct, samepalecoloras surrounding
cuticle;pupaA2dorsalhooknearlystraight,andsurpassingtheT2dorsalprojection;T2dorsalprojectionshortandbroadlyrounded
A. erotiaerotia "lerna"
3b.Larvaheadchalazaeslendertapered,thelargeroneswithacute,darktips;pitswell-defined,afewdarkerthansurroundingcuticle;pupa
A2dorsalhookcurvinginwardtoformnearlyclosedcirclewithT2dorsalprojection;T2dorsalprojectiondrawnouttoaroundedpoint
A.lycoriasmelanthe
4a.Pupaheadhornsoutwardlycurved,small,roundedoracutepoints 5
4b.ThelarvaeofA. mesentinaandA. thesprotiahavenotbeenexamined,butiftheykeytothiscouplet,thenpupaheadhornsoutwardly
curvedandleaf-likeseparatesthemfromthetwospecieskevedincouplet5 A. mesentina andA. thesprotia
5a. Larvaheadwith 3 extramedial chalazae nearly same size as principals, one each between ml/m2, m2/m3, and m3/m4, butnotwell
alignedwithdiem,resultinginaraggedline;pupaheadbonisverysmall,outwardlycurved,roundedpoints A.messana messana
5b.Larvaheadvvitfi2tiny,extramedialchalazae,oneeachbetweenni2/m3andni3/m4,usuallywellalignedwithprincipals;pupaheadhorns
outwardlycurved,pointedandleaf-like A.phylacapseudaethalia
Acknowledgements inventoryofthe macrocaterpillarfauna, andits foodplants and
parasitoids, of the Area de Conservacion Guanacaste (ACG),
Many thanks to the following colleagues at the Smithsonian northwesternCostaRica, http://janzen.sas.upenn.edu.
TropicalResearchInstitution(STRI): SunshineVan Baelforhergifts Mallet, &M.C.Singer. 1987. Individualselection,kinselection,
oflarvae,JoeWrightandMimaSamaniegoforapupa,RicardoCortez andtJ.heshiftingbalanceintheevolutionofwarningcolours: the
LfoirnalogGiostnizcaalleszup(pSorTtR.IDDoingintaalCIomnalgoinnfgorLahbe)rmfeotripcruelpoaursinilglutshteratcioolnos,r Mosse,viAd.eMn.ce1f93r3o.mbSutotmerefligeesn.erBailoilz.atJ.ioLnisnno.nSAocd.el3p2h:a3,37a-3n5e0o.tropical
figures, and Angel Aguirre (STRI Library) for obtaining copies of genus ofnvmphalid butterflies ofthe group Limenitidi. Nov.
important publications. Thanks also to Keidi Willmott (Natural Zool.39:12-20+pis. 1&2.
History Museum, London) for butterfly identifications, valuable Muller,W. 1886. Sudamerikanische Nvmphalidenraupen,versuch
commentsonthemanuscript,andforhisenormouscontributionsto einesnaturlichen SystemsderNymphaliden. Zool.Jahrb.,Zeit.
ourunderstandingofthegenusAdelpha. Thethoughtfulcomments Syst,Georg. Biol.Thiere 1:417-678+pi. 12-15.
of an anonymous reviewer are gratefully acknowledged. Special Otero, L. D. & A. Aiello. 1996. Descriptions ofdie immature
diankstotheSTRIadministrationfortheirsupportovertheyears. stages of Adelpha alala (Nymphalidae). Lepid. Soc.
J.
50(4):329-336.
LITERATURE ClTED Prudic, K. L.,A. M. Shapiro,&N. S.Clayton. 2002. Evaluatinga
AlELLO, A. 19S4. Adelpha (Nymphalidae): Deception on the wing. putative mimetic relationship between two butterflies,Adelpha
Psyche91:1-45. bredowiiandLimenitislorquini. Ecolog. Entom.27:68-75.
AlELLO,A. 1991. Adelpha ixialeucas: Immaturestagesandposition Srygley,R. B. 1994. Locomotormimicryinbutterflies? Theassocia-
withinAdelpha(Nymphalidae). Lepid. Soc.45:181-187. tions ofpositions ofcenters ofmass amonggroups ofmimetic,
DeYrCioesst,aP.RiJ.ca1n98b6u.tteHrofslitepsla(nPtaprielcioornJ.didsaae,ndPineartiudraael,haisntdorNyynmoptheaslio-n 3un4p3r:o1f4i5t-a1b5l5e. preyW. Philosophic. Trans. R. Soc. Lond. B.
dae). Res.Lepid.24(4):290-333. Stehr, Frederick 1987. Immature Insects. Volume 1.
DeVrHiiesst,orJPy...JP.ap1i9l8i7o.niTdahee,BPuitetreirdfalei,esNyomfpChoasltiadaRei.caParnidnctehteoinrUNnaitvuerra-l SteveKnesn,daPl.l/F.Hu(n2t0,01Duobnwuaqrudes,).IAownag,ioxsipve+rm75P4hpvpl.ogenvWebsite.Ver-
sityPress, xxii+327pp. sion 7, May2006. http://wvvw.niobot.org/MOBOT/research/AP-
Forbes, W. T M. Unpublished, incomplete manuscript in the web/
archives ofthe Museum ofComparative Zoology Library, Har- Willmott,K. R.2003a.CladisticanalysisofdieNeotropicalbutterfly
vardUniversity. genusAdelpha (Lepidoptera; Nymphalidae),withcommentson
Freitas, A. V. L., K. S. Brown, Jr., A. Aiello. 2001. Biologyof thesubtribalclassificationofLimenitidini. SvstematicEntomol-
Adelpha mijthra feedingonAsteraceae, anovelplant familyfor ogy28:279-322.
theneotropical Limenitidinae (Nymphalidae), andnewdataon Willmott,K.R.2003b.TheGenusAdelpha:ItsSvstematics,Biology,
Adelpha "species-GroupVII." Lepid. Soc.54(3):97-100. and Biogeographv (Lepidoptera: Nymphalidae: Limenitidini).
Fruhstorfer,H. 1907. Adelpha.InJ.:Seitz,A.TheMacrolepidoptera ScientificPublishers, Gainesville,Florida,viii+322pp.
oftheWorld. Div.II,Vol.5,pp.510-533+pis. 106-110a. Receivedfor publication 2 March 2005; revised and accepted 11
Godman, F. D., & O. Salvin. 1879-1901. Adelpha. In Biologia September2006
Centrali-Americana or Contributions to the Knowledge ofthe
Fauna and Flora of Mexico and Central America. Insecta.
Lepidoptera-Rhopalocera. Vol. I,pp.294-311;Vol. II,pp. 691,
692;Vol.Ill,pi.28-30& 109.
Janzen, D. H. &W. Hallwachs. 2004. Philosophy, navigationand
use ofa dynamic database ("ACG Caterpillars SRNP") for an
188 Journalofthe Lepidopterists" Society
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