.• .. ' ,.' ... . J.· �.;i.. . ) . . Catej¡'fJPlar$�ªs.9naIityin a central Br�li� .cerrado �"�::t7rT-��;,'tP;·'",; · �----�r-�-:·-;':---·�'""F::" ���-�- - ._--���-�-�.. �' .:,.._--- ._.. i_��• ..• ,�=,_ ....

H�lena c .. .MoJ!ai�l,IvóneR.·Diniz2andDelanoM. S. Silval

, Dep�e��Q deEcologi� Univ�rsicl�e de �r�Qia, 70910-900Bras ilia, DP, B,rasil. E-lIllIil;morais@�nb.br

2 bepaÍt�efitb" qéZoologia, UiMeJ'Sidade., deÍlrás �. I!.;709JO-9OÓ&í;¡Isíliá,))F, Brasil �l" E-mái}:; [email protected] , ,.�¡. , "�:�J '1-' ,. -' , t '¡'J�¡ . ,,'1 ' ;. �._ ; ,c: , , ""i:tl" 1'1< )� �:.i ,.Rt;�b�"e(Í2�J¡.n�199� .. Coí'Í'ected 2"VII�19�9. .;pcep�ed· 8.�VU"l 999.

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fb'\i�fltsn.�WJllj.:wmc(\flli'ín!t�,t1Kp· i�fl.tlence.\;¡.·· i:ltJ'UIl'dailOO an4ufe'} ;.' .tr(Jlp.c,aI.·..:l·I iS6l�ts . in ..'s�aIlpn�y ;diy'! .;ttopic�(; . (seiitDeAlingeF0�t�g6;·).W,oi{;t¡l::¡¡; . (lQS$;,j�h.il993 fo�;tef�rences).Maq:yf¡ff>P� .. 1 ", • .JI. 4�Cal�inseé'�'po�al '" .. · �i��,Seasómtt1)N\íántI Yátiationh�;. O�fttt��p;f¿Piréi�t���tbhfai��· ·"'.'· ".. �."Hik, fruhf�ehf�Re$� abun,��:ci¿��ri� 'ifItheW�!,/ seás�n�:·g.t�zem�973; Wi>lda,198��¡1trse� ' ."'. son�:{órest;a.�nim�';;Qrlepi�0Pter¡pl 'speci,�s may diapa�iúl$ . . ' 'tarlarV'aeinsidepupªl ;,::cocoo�S��' .. :.���t;th�·f�lo:Wjtig�et " se�oJl¡;('Aiil10,ti99�. iOtliéi�, tt',:'ay.pass�tbe:dry:ij �'tY';"::,;." :'�,}' " f';;��;� 'f:'>;:· . . ,,',',: /·;1?> ,�\ . 1026 REVISTA DE BIOLOGÍA TROPICAL studies regarding seasonal actlvltIes of the mation on the vegetation of these sites can be immature stadia of . Caterpillar season­ found in Ratter (1980), Felfiliand Silva (1993) ality is particularly evident in the tropical dry and Felfili et al. (1993). forest of Costa Rica, where caterpillar fauna Brasília is at an altitude of approximately fluctuates enorroously in biomass and propor­ 1,000 m and rainfall is concentrated in the six tional species composition within a year months between October and March, with an (Janzen 1993). average monthly precipitation of 207 mm. The In contrast to the various studies for sea­ dry months, from May through September, sonally dry forest of the neotropics, there are have a mean monthly precipitation of 24 mm. no studies of caterpillar seasonality in the The precípitation varíes among years (Fig. la). neotropical savanna. The central Brazilian cer­ Although the majority of the woody cerra­ rado (edaphíc woodland savanna) is highly do plant species are deciduous, flushing their seasonal, with marked wet and dry seasons, new ¡eaves between September and October at and most plant species are only briefly decid­ the beginning of the rainy season (Mantovani uous, despite the accentuated seasonality in and Martins 1988; Morais et al. 1995), this rainfall. The cerrado region exhíbits a very occurs in a gradual asynchronous process rich lepidopteran fauna, with around 1,000 among individuals of the same species and butterfly species (Brown and Mielke 1967). It between species, and many species retain their has been estímated that the fauna ranges leaves throughout the dry season (evergreen from 5,000 to 8,000 species (V. O. Becker and semideciduous). Sorne species lose old pers. com.). Plant-herbivore interactions may leaves and flush new ones synchronously, be qualitatively different from those of tropi­ remaining leafless for about one month during cal deciduous forests because leaves remaín the dry season. This is the case for three of the on plants throughout much of the dry season. host plant species examined in the present The present work describes the seasonal study (Erythroxylum deciduum, E. tortuosum abundance of caterpillars, the temporal occur­ and Qualea grandiflora). rence of caterpíllarspecies, and the duration of From April 1991 to April 1993, all cater­ the pupal stage for species collected in the pillars encountered on any dicotyledonous fieId at different times of the year. Thisstudy host plants in 1 ha block at FAL were collect­ represents a first description of the seasonality ed. Starting in May 1993 to May 1995 the of caterpillar abundance for the Brazilian cer­ number of exarnined plant species was rado, and ineludes sorne of the firstdata avail­ reduced to nine. Fifteen individuals of each of able on seasonal variation in species composi­ the nine host plant species in three genera were tion for any group in a cerrado area. examined for caterpillars once a week in FAL, JBB and PNB: Byrsonima coccolobifolia (Spr.) Kunth, B. crassa Nied., B. verbascifolia MATERIALS AND METHODS L. Rich. (Malpighiaceae), Erythroxylum deciduum St. Hill, E. suberosum St. Hin, E. The fieId work was conducted in four tonuosum Mart. (Erythroxylaceae), Qua/ea one-ha blocks of cerrado sensu stricto grandifloraMart., Q. multifloraMart. , and Q. (Goodland 1971) withín threepreserved areas parviflora Mart. (Vochysiaceae). For all near Brasília, DF, (15° 45' S, 47° 50' species, individual plants examined varied W): University of Brasília Experimental Farro from 0.50 to 2.50 m in height. The plants were (Fazenda Água Limpa - FAL), Botanic Garden censused once a weú during morning hours of Brasília (Jardim Botanico de Brasília - from 8:00 to 12:00 amo JBB), and National Park of Brasília (Parque Externally-feeding caterpillars were col­ Nacional de Brasília - PNB). Detailed infor- lected and indivídually reared in plastic jars on MORAIS et aL: Caterpillar seasonality in a central Brazilian cerrado 1027

¡eaves of the plant on which they were found. and then declined sharply until the lowest abun­ The duration of the pupal stage, and parasitoid dance in early-wet season (October) (Fig. 1b). emergence were accompanied under uncon­ The decrease in caterpillar abundance coincided trolled conditions (temperature, humidity, with thesecond half of the dry season in the cer­ light) in the laboratory. The durations of the rado regíon (Fig. la), during which the host pupal period were pooled by month for each plants are leafless or have senescent leaves species. The monthly means averaged over all (August-S eptember). species were used. This procedure reduces the The most frequent species tended to occur influence of those species that may occur throughout the year (see Table 1 for examples). abundantly during specific months. These multivoltine lepidopteran species belong Caterpillars were classified as morphos­ to various taxonomic groups and may be pecies, but because the identification of mOf­ polyphagous or restricted to one plant genus in phospecies among collectors was not always the study areas (Diniz and Morais 1995, 1997). consistent, for the analyses of temporal species 400 o a x o + Q composition we considered only the species 350 b E a 300 + • -5 + �Mea that emerged as adults. This results in an 250 .� 200 • 5 � 1990 underestimate of species' occurrence. The g 8 ""- ISO • 1991 O " adults were identified by Vitor O. Becker 'H lOO x 1992 o: O (EMBRAPA - Brasília) and Keith S. Brown JI. 50 e 1993 O o 1994 J F M A M J J A S O N D (UNICAMP). Voucher specimens were A 1995 deposited in the Departamento de Zoologia Month Collection at the Universidade de Brasília. b

RESULTS

Of the approximately 4,000 caterpillars collected in 80 species, 61 genera and 41 farn­ ilies of plants, we successfully reared about JFMAMJ JASOND Month 1,500 lepidopteran adults representing 247 species from 32 families. Lists of species of caterpillars and their host plants are presented c by _ Diniz and Morais (1995). One-hundred­ fifty-one (61 %) of the 247 lepidopteran species occurred at low density (Iess than four times in four years). Between May 1993 and 1995, we con­ ducted censuses of nine host plants JFMAMJ JASOND 10,800 Month species, 15% of which had externalIy-feeding Fíg. 1. A) Mean monthly precípitatíon for Brasília, DF, caterpillars. The 137 lepidopteran species (24 Brazil (1980-1995). Data from the IBGE meteorological farnilies) reared from the three genera of cer­ station. Mean annual precipítation:l,469mm; Mean annu­ rado host plants during the quantitative survey al temperatura=2 ! o C. B) Percentage of plants wíth cater­ are listed in Diniz and Morais (1997). pillars. C) Precentage of caterpillar (unit used=plant with Caterpillar abundance (expressed as the caterpillars) from which parasitoids emerged. Data for B and e were collected between May 1993 and May 1995 in propmtion of plants with caterpillars) increased cerrado areas near Brasília. nine host plant species were sharply in mid-April and remained high until examined: Byrsonima spp. (3), Erythroxylumspp. (3), and mid-July, during the first half of the dry season, Qua/ea spp. (3). Theshaded area represents the dryseason. 1028 REVISTA DE BIOLOGÍA TROPICAL

TABLE 1

Lepidopteran species and the occurrence of their caterpillars during the year in the cerrado sensu stricto of the Federal District, Brazil. The period between May to September corresponds to the dry season.

M o n t h

Family Species J F MA OND

Oecophoridae Cerconota achatina (Zeller, 1855) x x x x x x x Hesperiidae Chiomara punctum (Mabille, 1878) x x x x x x Geometridae Cye/omia mopsaria (Guenée, 1857) x x x x x x Dalceridae Dalcerina tijucana (Schaus, 1892) x x x x Lyrnantriidáe Eloria subapicalis (Walker, 1855) x x x x x x x Arctiidae Fregela semi/una (Walker, 1854) x x x x x x x Oecophoridae Gonioterma indecora (Zeller, 1854) x x x Oecophoridae lnga haemataula (Meyrick, 1912) x x x Sphingidae lsognathus caricae (Linnaeus, 1764) x x x x x x x Megalopygidae Megalopyge albicollis (Walker, 1855) x x x hipparchia (Cramer, 1777) x x x Oecophoridae Stenoma sp. x x x x Oecophoridae Stenoma staudingerana (Maassen, 1890) x x x x x

Nymphalidae Siderone marthesia nemesis (Illiger, [1801]) x x x x x x Urodidae Urodus sp. x x

Oecophoridae Eonichla sp. x Megalopygidae Trosia dimas (Cramer, 1775) x

Mimallonidae Mimallonidae sp. x

The caterpillar species collected in the the wet season (Fig. 2) due, in part, to the pres­ field were in various stadia, so complete life ence of species with pupal diapause, such as cycles are not known. This study was restricted Trosia aimasand Eonichla sp. (Table 2). to the presence of caterpillars in the field and pupal duration under laboratory condiuons, and no infonnation is available for eggs and adults. 45 The pupal period was variable among individ­ {4� r.r uals and within species (Table 2), and varied ; �) � f according to the time of the year (Fig. 2). � t � i At the beginning of the dry season (Fig. (191 I I ¡ la) the insects tended to have shorter pupal . 11,2) � t periods (Fig. 2), resulting in more generations �) � t which coincided with the seasonal peak of caterpillar abundance in the field during this period (Fig. lb). The duration of the pupal JFMAMJJASOND period increased as the dry season progres sed Month

and decreased sharply at the transition between Fig. 2. Average pupal period (± se) for 247 lepidopteran the dry and wet seasons. Mean pupal duration, species which pupated in different months, [n]. as well as its variance, increased at the end of The horizontal bar correponds to the dry season. MORAIS et al.: Caterpillarseasonality in a central Brazilian cerrado 1029

TABLE 2

Lepidopteran species and their average pupal period in days, under laboratory conditions.

Pupal period (days)

Species n mean sd min max

Cerconota achatina (Zeller, 1855) 131 14,0 6,23 5 31 Chiomara punctum (Mabílle, 1878) 13 17,8 20,63 5 72 Cyclomia mopsaria Guénée, 1857 14 11,1 5,43 4 22 Dalcerina tijucana (Schaus, 1892) 8 20,4 2,20 18 24 Eloria subapicalis (Walker, 1855) 10 15,4 9,87 9 41 Fregela semi/una (Walker, 1854) 43 22,0 3,95 13 31 Gonioterma indecora (Zeller, 1854) 20 16,8 19,64 7 96 Inga haemataula (Meyrick, 1912) 13 19,8 8,92 6 34 Isognathus caricae (Línnaeus, 1764) 8 24,0 9,87 8 32 Megalopyge albicollis (Walker, 1855) 28 66,8 31,15 28 144 Phobetron hipparchia (Cramer, 1777) 9 71,2 52,07 20 148 Stenoma sp. 55 98,6 54,31 7 183 Stenoma staudingerana (Maassen, 1890) 16 20,2 20,10 5 81

Siderone marthesia nemesis (Illiger, [1801]) 5 14,6 1,95 12 17 Urodussp. 4 9,5 2,08 7 12

Eonichla sp. 235 Trosia dimas (Cramer, 1775) 266 Mimallonidae sp. 351

Sornespecies have short pupalperiods and period of 14 days (Table 2). However, the can occur throughout the year or be restricted caterpillarscollected in the field during the dry to one particular season. For example Siderone season remained as larvae twice as long in the marthesia nemesis was found during February laboratory (54.56 ± 27.2 days, n = 82) as those and March in four consecutive years (second collected during the wet season (27.5 ± 19.0 half of the wet season), feeding on mature days, n = 31; t = 5.08 P < 0.001). These cater­ leaves of Casearia sylvestris (Flacourtiaceae) pillars did not show any tendency for dor­ (Morais et al. 1996). Its pupal stage lasted 15 mancy and normally continued to feed. days. The adults also occur in the gallery forest Larvae reared on new leaves of B. coccolobi­ (Pinheiro and Ortiz1992) where they may have folia produced heavier pupae and bigger more generations. Other species, such as adults (1. Andrade pers. com.). Urodus sp., was found only in November in Other species had very long pupal periods four consecutive years and always on the same during the dry season (Table 2, Fig. 2). In host plant species (Erytroxylum deciduum). Megalopyge albicollis the duration of the Others, for instance Fregela semi/una, pupal period decreased between early-Julyand Cyclomia mopsaria and Cerconota achatina, late-August, and the adults emerged synchro­ have short pupal periods independent of the nously during the wet season in a 20-day span time of the year Oess than 31 days). Cerconota between the end of October and the beginning achatina for example, fed on the three of November. In Stenoma sp. the long pupal Byrsonima species (Andrade et al. 1995) period is variable among individuals and adult throughout the year and had an average pupal emergence is asynchronous, but 65% of the 1030 REVISTA DE BIOLOGíA TROPICAL

June to August initiated pupae emerged as reasons to explain low caterpillar density at the adults in November. beginning of the rainy season in the cerrado: Relative abundance of parasitoid on (5) the accumulated water stress in the late-dry caterpillars (unit used was plant with caterpil­ season; and (6) the delay of adult emergence. lars) remains low until the beginning of the In the late dryseason there is low precip­ wet season, in October, when it increases itation (Fig. la), the relative humidity of air is rapidly (Fig. lc). low (often below 20% during warmer hours in September), and the majority of leaves are senescent (poor quality of available food) DISCUSSION which, together, probably constrain larval per­ formance and/or increases caterpillar mortality Relative abundance of caterpillars varied at this time. Even a number of lepidopteran monthly in the cerrado. The highest caterpillar species that normaHy occurs throughout the abundance occurred during May and July, and year was often absent the field in the months then declined and remained low until mid­ between August and September (Table 1). A April (Fig. 1b). The caterpillar abundance similar pattem, with leaf quality reduction and variation in the field was. not related to leaf decreased reproductive activity as the dry sea­ phenology. The higher caterpillar abundance son progresses was shown for sorne butterfly occurred when leaf production was very low species in tropical savannas of Australia (Morais et al. 1995, Price et al. 1995), and (Braby 1995) and in tropical dry forests of most of the existing leaves were old. Soon Costa Rica (Odendaal 1990). after (September-October), the majority of cer­ Sornespecies had very long pupal periods rado plant species flushed their leaves, fol­ (between 8 and 12 months), suggesting that lowed by the peak of newly expanding leaves they may be univoltine and that the pupal stage in November-December. In spite of aH these may function as a resting stage (Table 2). In changes of leaf phenology, caterpillar abun­ Costa Rican tropical dry forest a large propor­ dance remained low fromAugust to April. tion of moth species spend all or part of the dry One partial explanation for the low cater­ season as dormant pupae, and many of them pillar abundance found during the early rainy are, indeed, univoltine (Janzen 1987). In the season could be leaf pubescence. In fact, leaf cerrado areas the proportion of univoltine phenology could be a major factor limiting species appears to be low. damage on leaves in this cerrado area The delay in adult emergence at the begin­ in the early rainy season (unpublished data), ning of the wet season may be related to the as many cerrado plant species are pubescent unpredictable precipitation in October. Similar with young leaves more hairy than mature pattems were also found for moth species at ones. Thus, caterpillars that consume pubes­ Santa Rosa National Park in Costa Rica cent host plants may be rare during (Janzen 1993). In fact, the onset of rains is September and October when these plants are variable among years (Fig. la), and the species flushing newleaves. that have long pupal periods during the dry Janzen (1993) suggests four possible season tend to emerge in November. The explanations for the low caterpillar abundance longer development period in the dry season registered in the period from four to six weeks may be the result of reduced food quality, after the first rains at Santa Rosa National water stress, and lower temperature. During Park: (1) the delay or spread of oviposition; (2) October and November the number of cater­ the time required for eggs to hatch; (3) small pillars coHected was low and constitutes the species at low density (inconspicuous); and (4) fust generation after the dry season. high caterpillar mortality due to predation. In deciduous dry forest in Costa Rica, Here we suggest elements regarding two other Janzen (1987, 1988, 1993) found a peak of MORAIS et al.: Caterpillar seasonality in a central Brazilian cerrado 1031 caterpillar abundance in the second half of the Although there are no hard data demons­ rainy season. Our results contrast with what trating the theory of enemy-free space in the was found for Costa Rica. One of the explana­ cerrado area, the proportion of caterpillars at­ tions given for tbis phenomenon, could be the tacked by parasitoids was lower during drysea­ availability of host plant leaves throughout the son than in the early wet season (Fig; 1c). Pa­ year in the cerrado. In fact, in the cerrado, rasitoid density increases in the early wet sea­ many species retain their leaves through the son (October), coinciding with the time of 10- dry season, perrnitting a bro:;tder range of life west caterpillar abundance (Figs.lb, c). Appa­ histories than in the Costa Rican dry foresto rently, the first half of the dry season constitu­ However, the higher caterpillar abundance tes the temporary parasitoid enemy-free space. during the first three months of the dry season It is tempting to suggest, as did Janzen (Fig. 1b) is puzzling. Ecological opportunities (1987, 1988) for Costa Rica, that both para­ for phytophagous insects are broad, encom­ sitoid and predator pressures may regulate the passing such concepts as host plant availabili­ seasonal fluctuations in caterpillar abundance ty, bioclimatic conditions and enemy-free in cerrado area. space. As discussed before, the bigher leaf pro­ duction and most rains occur at a time when caterpillar densities are quite low. Aiello ACKNOWLEDGMENTS (1992) suggested that the decrease of para­ sitoids in Panama is a result of the decline in We thank V. O. Becker and K. S. Brown butterflynumbers during the severe dryseason for the identification of lepidopteran species; and, when humid conditions return, the tem­ R. Marquis (University of Mis�ouri-St. Louis), porarily enemy-free space (parasitoid) allows P. Price (Arizona State University), W. Benson the butterfly populations to increase rapidly. (University of Campinas), R. Macedo, J. Hay Consequently, when parasitoids find them­ and J. Marinho-Filho, (University of Brasília) selves in the center of abundance, they also for their helpful comments and criticisms on increase and quick1y become abundant. The the manuscript; J air Maia for bis help on sorne lag of time between the recoveries of hosts and figures, R. Henriques (University of Brasília) parasitoids make the temporary increase of for allowing us to consult bis organized cli­ butterfly populations possible. Similarly, matic data; and Aurora Bendicho (Camagüey, predator and parasitoid activities can play an Cuba) for the spanish abstracto The following important role in influencing the numbers of students provided field and 1aboratory assis­ caterpillars encountered in the field and may tance in different years: 1. Andrade, L. help keep caterpillar densities down during the Baumgarten, A. Botelho, D. Coelho, R. wet season in Brazilian cerrado. Cordeiro, H. Cunha, L. DelClaro, C. Días, E. It is well known that predators and para­ Emery, P. Estevanato, M. Lasneaux, R. sitoids can exert high pressures on caterpillar Malcher, N. Menezes, M. Millhomen, F. Pinto abundance (Stamp and Casey 1993, Marquis and H. Rocha. The research was supported by and Whelan 1994). The literature known for Brazilian Research Council (CNPq) grant cerrado shows that predator abundance is, 520255/95-0. D. Silva, was supported by grant indeed, bighest during the wet season. In 800083/86-5. Brasília, Polistes nests increase during the rainy season and colonies initiated in the dry season produced fewer pupae than those initi­ RESUMEN ated in the wet season (Ramos and Diniz Este trabajo se realizó desde 1991 a 1995 en cuatro 1993). Also, bird breeding activity is greatest áreas de cerrado sensu smcto en Brasilia, DF, Brasil. El between September and December (Femandez clima dela región es estacional, con las estaciones seca y llu­ 1978, Cavalcanti 1990, Macedo 1992). viosa bien marcadas.Nueve especies de plantas hospederas 1032 REVISTA DE BIOLOGÍA TROPICAL

(tres géneros y tres familias) fueron examinadas en 10 800 Diniz, J. R. & H. C. Morais. 1997. Lepidopteran caterpi­ censos desde 1993 a 1995 y en 15% de ellas se encontraron llar fauna of cerrado host plants. Biodiver. Conservo larvas. Se obtuvieron los adultos de 247 especies de lep­ 6: 817-836. idópteros (32 familias). La proporción de plantas con larvas aumentó rapidamente a partir del 15 de abril, permanecien­ Felfili, J. M. & M. C. Silva. 1993. A comparative study of do alta hasta julio, estación de seca y fue menor al comenzar cerrado (sensu stricto) vegetation in central Brazil. J. la estación lluviosa. La duración media del período de pupa Trop. Ecol. 9: 277-289. varia durante el año (multivoltinas). La baja densidad de lar­ vas al inicio de las lluvias podría deberse al estrés hídrico Felfili1, 1. M., M. C. Silva, A. V. Rezende, J. W. B. Macha­ acumulado durante la estación seca; la defensa química y do, B. M. T. WaIter, P. E. N. Silva & 1. D. Hay. 1993. fisica de las hojas nuevas; la demora en la emergencia de los Anal ise comparativa da florística e fitossociologia da adultos y la actividad de depredadores y parasitoides. vegetayilo arbórea do cerrado sensu stricto naChapa­ Probablemente el espacio libre de enemigos durante los tres da Pratinha, DF, Brasil. Act. Bot. Bras. 6: 27-46. primeros meses de la época seca favorece la abundancia de larvas en el campo. Femandez, R. A. N. 1978. O comportamento alimentar como fator de isolamento ecológico em oito espé­ cies de Tyrannidae (Aves) do PlanaltoCentral. Mas­ lers Thesis, University of Brasília, Brasília, DF, REFERENCES Brazil.

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