Table Of Content41: 76-83,2002 (2009) 77
an important phenomenon in butterflies (Willmott & DeVries, P. j. 1987. The butterflies of Gosta Rica atid their Nattiral
Freitas, 20()(5;Janz et ai, 2006). In Nymphalinae, recent History. Princeton University Press, New Jersey.
H.wwARt), K. J. 1969. Datos para el esiudio de la ontogetiia de
studies have shown that the plasticity in host plants
lepidopteros argentinos. Miscelauea. Instituto Miguel Lillo.
ranges may be related to diversification processes Universidad Nacional de Tucittnan 31:1-142.
(Weingartner et al, 2006; Nylin & Wahlberg, 2008). In Janz, N., S. Nvt.iN & N. WAiii.itERC. 2006. Diversity begets diversity:
host expansions and the diversification of jtlant-feeding insects.
this sense, H. bella may be a model for understanding
BMG Evolutionaiy Biology 6: 4.
the importance of local adaptation in the evolution
Lorenzi, H. 2000. Plantas daninhas do Brasil: terrestre, aciuaticas,
of host plant use in Neotropical butterflies. parasitas, toxicas e medicinais. 3rd ed. Editora Plantarum,
Nova Odessa.
Nvi.tN, S. & N. W.Mtt.iiERG. 2008. Does plasticity drive s]teciatioti?
Acknowledgements
Host plant shifts and diversificatioti in nymphaline butterflies
(Lepidoplera: Nymphalidae) during the tertiary. Biological
We thank Adriano Cavalleri and Crisliano A. Lserhard for
Journal of the l.innean Society 94: 11,5-130.
their lielp in the field, and Niklas Wahlberg and Keith S. Brown
Scon , J. A. 1986. fhe Butterflies of North America: a natural
If. for reading the last version of the nianuscri|tt. To CNP<] for
histoiy and field guide. Stanford University Press, Stanford.
the doctoral fellow'shijr granted to lAK (Proc. 140183/2006-0).
Toi.Ktx), Z. D. A. 1973. Eauna del noroeste argctitino. (iontribucion
AVLF thatiks FAPESP (grants 00/01484-1 and 04/05269-9) atid
al conocimietito de los lepidopteros argentinos III. ilypanarlia
the BIOTA-FAPESP program (98/05101-8), the Brazilian CNPq
hella (E.) (Rhopalocera-Nymphalidae). Acta Zoologica Lilloatia
(fellovvshi|) 300315/200.5-8) and the National Science Foundation
30: 23-35.
grant DEB-0316505. This work was partially supported by the CNPq
Tot.M.w, T, & R. Lewincton. 1997. Butterflies of Britain atid Fairope.
(Proc. 478787/2001-4).
Harper Gollins, London.
WEtNC.ARTER, E., N. WAtit.BERC & S. Nvi.in. 2006. Dytitituics of
Literature cited host plant use and species diversity in Polygoiiia butterflies
(Nymphalidae). Journal of Evolutiotiary Biologv' 19: 483-491.
At.wo,P. & L. l lERNANttE/.. 1987. Atlas de las inariposas diurnas Wit.t.Mon, K. R., J. P. W. H.\tt & G. LA\t.\s. 2001. .Systematics of
de Cuba (Lepidopteni: Rhopalocera). Editorial Cientifico- f/ypanarlia (Lepidoptera: Nymphalidae: Nymphalinae), with
Tecnica, La Habatia. a test for geographical .s|)eciatiou mechanisms in the Andes.
BEC.ou.oNt, G. W., S. K. HAi.t., A. 1,. Vit.oRt.x & G. S. RoitixsoN. 2008. .Systetnatic Entotnolog\' 26: 369-.399.
Catalogite of the ho.st|)lants of the Neotroirical butterflies / Wit.t.MOTt, K. R. & A. V. L. FREit.vs. 2006. Higher-level phylogeny
Catalogo de las plantas huesped de las inan|)o.sas Neotro|)icale.s. of the Ithomiinae (Lepidojitera: Nymphalidae): classification,
in: ni3ni - Monografias Tercet Milenio, Vol. 8. S.F..A.. RIBES- liatterns of larval hostplant colonization and diversification.
CYTEl), The Natural Mistoiy Museum, Instituto Venezolano Gladistics 22: 297-,368.
de Investigaciones Cientt'ficas, Zaragoza.
Biiczanko, C. M. 1949. Acraeidae, fleliconiidae et Nym|thalidae Lucas A. Kaminski, PPG-Ecologia, Departamento de Zoologia,
de Pelotas e sens arredores (CotUribuicao ao conhecimento Universidade Estadual de Gampinas, GP 1)109, 13083970, Gampinas,
da fisiografia do Rio Grande do .Sul). Livraria do Cilobo, Sao Paulo, Brazil.
Pelotas. lucaskatninski @yah oo. com. hr
Brow'n, K. S., Jr. 1992. Borboletas da Serra do Japi: Diversidade,
habitats, recimsos tilimentares e variagao tem|toral, p. 142-187. Andre V. L. Freitas*, Departamento de Zoologia, Instituto de
In\ L. P. G. MORELLATO (ed.). Historia Natural da Serra do Biologia, Universidade Estadual de Gam|)iuas, GP 6109, 13083970,
Japi: Ecologia e preservayao de nma area florestal no sudeste Gatnpinas, Siio Paulo, Brazil.
do Brasil. UNIGAMP/FAPESP, f iampinas. baku@u n ka mp. hr
t)'AtiRER.\ B. L. 1987. Butterflies of the Neotro|5ical Region. Part IV.
Nymphtilidae (partim). Black Rock Hill House, Victoria. *(Mrn'spo>i(ting o ill hoc
Duration of molt in a Neotropical butterfly (Lepidoptera: Nymphalidae)
In thi.s note I present the results of an observational medium-sized (forewing length 29-32 mm), relatively
study on the length of molting in caterpillars of common butterfly inhabiting the Reserva Biologica do
Hypothyris ninonia daela (Boisduval, 1836), an ithomiine Po^o D’Auta (RPA), a 277 ha forest fragment in Juiz de
(Lepidoptera: Nymphalidae: Ithomiinae) butterfly of Fora, MG, Brazil (21°4,5’S, 43°20’W) (Costa, 2002).
southeastern Brazil. To determine the proportion of Hypothyris has five larval instars that feed
time spent molting I meastired the time caterpillars exclusively on Solanum (Solanaceae) species (Costa,
spend in molting relative to total development time. 1999). Ovipositing females lay their eggs singly on
I was able to do this using a specific external marker the underside of hostplant leaves. Neonates are
(see below) that enabled me to recognize the start approximately 1.5-2 mm in length and are translucent
and the end of each molt period. white. Their color turns from white to greenish
The species, hereafter referred to as Hypothyris, is a after feeding. However, since food consumption
78 /. Res.Lepid.
stops during tlie molt process, and the larva partially etc. For computation, I assumed that all changes
evacuated its gttt in preparation for molt, the distal end between growth and molting behavior occurred at
of the body changes coloration from a characteristic the midpoint of the time interval between any two
gray-green to translucent white. This color change of consecutive surveys. Larval size (body length to the
the last larval body segments thus provides a simple nearest 0.5 mm) was measured at least once during
and reliable external marker that indicates the start each instar.
of each larval molt, thereby facilitating the precise The climate at Juiz de Lora is characterized by
determination of the dttration of each molt. a warm rainy sttmnier and a cool diy winter. The
The color change mediated by gut contents does average monthly temperature is 18.7°C, varying from
not occtir in caterpillar species that have a pigmented 21.6°C (Lebruary) to 16.1"C (July). The diurnal range
or otherwise ojjaqite body wall. For stich cases, it is in air temperature greatly exceeded this average
difhctth or impo.ssible to determine the start of molt monthly change of only 5.5°C. On the other side, the
process based tipon visual inspection. Consequently, monthly distribution of precipitation is very unequal
Hypothyris larvae provide an ideal subject for the study throughout the year, varying from 272 mm (January)
of molting behavior. For the ptipal molt, color changes to 19 mm (July). Mean annual precipitation is-1,500
from gray-green to brownish and then to whitish. In mm (Costa, 1991).
all cases, shedding of the old head capsule marks the To test whether the duration of the molt period
end of molt. is a function of the growth period of the same instar,
I collected Hylmlhyris 'xmnvAixiYes from a previoitsly I tised linear regression analysis (least square). As a
used study site at RPA (Costa, 1996) with all eggs and complement, I also used regression analysis to test
larvae between 21.IV (n = 14) and TV. 1995 (n - 2). if the time spent in each stage of development is a
The 16 specimens were reared under semi-natttral constant proportion of the total developmental time.
conditions, indoors, snbjected to natural photoperiod Statistics (tests, .symbols, and terminology) followed
and air temperatitre fluctuations, between 22.IV and Sokal and Rohlf (1981)
23.VI.1995 through adult emergence (8 $$, 5 SS)- Eleven of thirteen reared larvae had the usual five
Each larva was reared individually in a glass jar with larval stages, but two caterpillars molted to an extra
moist hlter paper. Larvae were fed Solanuvi cernuum sixth instar before pupation. The latter two larvae
Veil., an abttndant hostplant (Costa, 1999). Frass were much smaller in body length (ca. 15 mm) when
was removed with leaves and moisttire were stipplied they underwent the extra instar. These exceptions are
as necessary. concordant with the hypothesis that there is a critical
The data for the study were from: (1) five minimal length for a caterpillar prior to pupation
immatures individuals (4 $$: # B7, B8, BIO, B12; 1 S'- (Nijhout, 1975; see also Esperk et al, 2007).
B5) reared from egg to adulthood (used to compute Table 1 gives the average time in the growth stage
species-specific parameters); (2) four immattires of Hypothyris caterpillars increasing with sticcessive
individtials (Bl, B2, B3, B4) that were collected as
larva (three L3, one L5) and reared to adulthood; (3)
two immattires individuals (B9, Bl 1) reared from egg
Table 1. The time duration of Hypothyris caterpillars during
to adulthood but that molted to an extra sixth instar
different instars (LI -L5) and corresponding molts. Values
before pupation; and (4) two immatures individtials are means ± SD from five larvae.
(PI, P2) collected as eggs on 1 .V and that were reared
to adtihhood in different (plastic) containers. The
other three immattires (B6, B13, B14) collected as
Phase (length, mm) Duration ± SD (hours)
eggs on 21.IV died during their second instar.
All caterpillars were examined by means of a LI (1..5-3.5) 73.31 ± 6.85
magnifying lens 3-6 times daily for approximately 1- L1-L2 molt 14.45 ±5.21
3 minutes each to determine the time spent during L2 (3.5-5.5) 118.75 ± 54.69
each larval instar dttring both growth (feeding) and L2-L3 molt 21.14 ±2.55
molting (non-feeding) periods. This resulted in 139
L3 {6.()-8.()) 119.1 1 ±.50.72
surveys between 22.IV and 29.V. 1995, when the last
L.3-L4 molt 20.65 ± 3.26
larva pupated. For each determination, I categorized
L4 (8.5-13.0) 116.51 ±2.3.07
caterpillar behavior as either “growth” (feeding,
resting, moving around) or molting. I also noted L4-L5 molt 28.03 ± 3.25
activities stich as mode and place of feeding, mode 1,5 (13..5-27.0) 169.84 ±23.33
and jilace of resting, body aspect dttring molting. L5-piipa molt 36.93 ±6.41
41: 76-83,2002 (2009) 79
larval instars, changing from 73.31 hours ± 6.85 SD in a fixed proportion of instar length.
LI to 169.84 hours ± 23.33 SD in L5. In parallel, the Like other ectotherms (e.g., Casey, 1993), the
average time spent in the molt stage also increased performance of//y/ioZ/iyrh caterpillars is temperature-
with larval instar, changing from 14.45 hours ± 5.21 SD sensitive and varies in different months of the year.
(L1-L2 molt) to 36.93 hours ± 6.41 SD (L5-pupa molt, For instance, studies on the performance of these
including prepupa). However, considering larvae B9 caterpillars in the held have shown that the growth
and B11, these values are higher: 212.53 hours ± 43.85 rate increases linearly with temperattire (unpublished
SD (L6) and 41.4 hours ± 11.58 SD (L6-pupal molt). data). Thus, the results of this study, which were
The duration of growth and molting periods of obtained dtiring autumn when the number of
the same instar are related; longer feeding periods immatures in the held is decreasing, may overestimate
correspond to longer molting periods (linear the values dtiring warm seasons, spring and especially
regression analysis for data of Table 1: R2 = 0.8544; t summer, when immature density is higher (Costa,
= 4.19; p < 0.05). This relationship remains significant 1991) and development is probably faster. I believe
even including in the regression analysis data from all the restilts of this study show that molting and growing
13 caterpillars (Fig. 1). periods of//y/io/Ziyr/.s caterpillars are correlated, so that
The results of the regre.ssion analysis with data larval growth rate can be reduced directly (i.e., low
of Table 1 indicate that approximately 85% of the temperattires prolong molt period) or indirectly (i.e.,
variation in the length of molting was explained by low temperatures prolong growth period that implies
variation by dtiration of growth. The restilt appears iti a molt period correspondently longer).
relevant and more so considering that the sample
included caterpillars from different instars growing Acknowledgements
simultaneously under a naturally fluctuating daily
I would like to acknowledge two anonvtuous reviewers, they
temperattire regime.
made comments that encouraged me to revise and improve the
In relative terms, the dtiration of molt periods
manuscript. 1 am also gratefnl to Jennifer R. A. Taylor and Jorge
varied from 14.7% (L3-L4 molt) to 19.4% (L4-L5 R. Litna, for helji with the Etiglish text on earlv version of the
molt) of the total time that larva spent in each instar mannscri|tt; atid Rndi Mattoni, for his generous help with the lititil
version of this [tiiper.
period (Table 1). However, the percentage of time
spent by caterpillars during molting was not a ftmction
Literature cited
of the total dtiration (growth phis molting) of each
instar (linear regression analysis: R2 = 0.098; t = 0.57;
C.\si£v,T. M. 1993. Effects of tetuperatttre on foraging of caterpillars,
p > 0.9), as would be expected if molt duration were pp. .6-28. In: N. E. Statnj) & T . M. Casey (eds.). Cater]jillars:
ecological and evolntiomiry cotistraints on foraging. Chajitnati,
New York.
Co.sT.v, E A. P. L. 1991. Sohre a tttilizayao de Solanum rmuiiiw Veil.
(Solanaceae) como planta hospedeirti por Hypolliyri.i ninonia
Figure 1. Relationship between growth and molting
daeta (Bdv., 1836) (Lepidojjtera: Nytnphalidae: Ithomiinae).
periods of the same instar (from LI to L5 plus two extra
MS thesis, Universidade Estadiittl de Catnpinas, Cainjiitias.
L6) for 13 Hypothyris caterpillars. The linear regression
Co.STA. E. A. P. I.. 1996. nemografia foliar: contando folhas no
analysis for all these data also produced significant results
hrayo-de-pregnica. Ciencialloje 123:22-24.
(R2 = 0.2708; t = 4.52; p « 0.001). In this figure, seven of
Co.ST.v, E. .V. P. L. 1999. New records of larval hostjtlants for
the original 57 points are coincident: two full triangles (LI),
Ithomiitiae butterflies (Nytnphalidae). Revista Brasileira de
three empty circles (L3), and two empty quadrats (L4).
Biologia .69: 46.6-469.
Costa, E. A. P. L.. 2002. Bttrboletas Ithomiinae (Lejiidoptera:
Nytnphalidae) da Reserva Bioldgica do Poyo D'Anta (|ni/, de
60 Eora, MG). Revista Brasileira de Zoocieticias 4: 14.3-149.
Esi’KRK, T, T.VMNtARU, T. & S. Nu.t.x. 2007. Intraespecific variability
JO in tnimber of larval instars in insects. Journal of Economic
Entomology 100: 627-646.
Ni|tiocT, H. E. 1975. A threshold size for metamorphosis in the
iLl
tobacco hornworm, Mandura sexta (L,.). Biological Bulletin
aL2
149:214-22.6.
oL3
•a SoK.\t, R. R. & E. J. Roiii.f. 1981. Biotnetry: the pi incijiles and
S 20 dL4
practice of statistics in biological research, 2 ed. Ereeman,
«LJ
San Erancisco.
10 • L6
0
Feupe a. P. L. Costa, Caixa Postal 201, Viyosa, MG, Brazil 36.670-000.
250
meiterer@h(}t mail, com
Gro^vth tune (hours)
