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Journal of The Lepidopterists' Society

Volume 50                                     1996                                      Number 4

Journal of the Lepidopterists' Society 50(4), 1996, 273-289

POPULATION BIOLOGY OF HETEROSAIS EDESSA

(NYMPHALIDAE) AND ITS ASSOCIATED ATLANTIC FOREST

ITHOMIINAE COMMUNITY

Andre Victor Lucci Freitas

Curso de Pos Graduagao em Ecologia, Instituto de Biologia, Universidade Estadual de Campinas, C.R 6109, 13083-970, Sao Paulo, Brasil

ABSTRACT. The early stages of Heterosais edessa, an advanced species of Ithomiinae (Nymphalidae) are described. Adult population size, sex ratio, longevity, food resources and movement are given for this species and compared with those of 17 other members of the local Ithomiinae community.

Additional key words: mark-recapture, Godyridini, Cestrum, Solanaceae, K-strategist.

Long term population studies may reveal important features of ecology and adaptation in tropical butterflies (Ehrlich 1984). DeVries (1994) emphasized the importance of such studies as a basis for understanding Neotropical butterfly communities, encouraging studies of life histories. The Ithomiinae, along with the Heliconiini, attracted the attention of early naturalists as central models of mimicry rings in the Neotropics (Bates 1862, Muller 1879), but there are few works concerning population ecology of ithomiine butterflies (Brown & Benson 1974, Drum-mond 1976, Haber 1978, Vasconcellos-Neto 1980, 1986, 1991, Young & Moffett 1979, Freitas 1993).

Heterosais edessa is a common ithomiine species found in humid primary and secondary forests, mostly in coastal southeastern Brazil, being rare on mountain slopes and nearly absent on the central plateau. Other closely related species or subspecies extend throughout the Amazon and as far northwest as Costa Rica. Heterosais is considered one of the most advanced genera within the Ithomiinae (Brown & Freitas 1994), and the study of ecological parameters in this genus may help in the understanding of the evolution of this subfamily.

This paper describes the juvenile biology of H. edessa and its adult population ecology in relation to other species of the associated ithomiine community.

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Study Site And Methods

Observations were made from July 1988 to July 1992, in the locality of "Morro do Japui" (46°24'W 23°59*S), Sao Vicente, Sao Paulo, Brazil The study area is covered by submontane rain forest (Ururahy et al 1987), with an annual rainfall near 2500 mm and an average annual temperature of 21°C (Setzer 1949, Prodesan 1969, Nimer 1972). A large part of the area is secondary forest, with a predominance of forest edge plants. A detailed map of the locality is given in Freitas (1993).

In order to study the life cycle of H. edessa, eggs and larvae were collected on individuals of the host plant species, Cestrum laevigatum Schldl. (Solanaceae), in the forest. Larvae were reared on leaves of this host in plastic boxes, cleaned daily. Egg size is presented as length and diameter, and the head capsule size of larvae as the distance between the most external ocelli (as in Freitas 1991, 1993, and Freitas & Oliveira 1992). All measurements were made using a microscope fitted with a calibrated micrometric ocular.

A mark-recapture census for Heterosais edessa and 18 other Ithomi-inae was conducted from May 1991 to May 1992. Visits were made from 1 to 5 times per week, except from December 1991 to April 1992 (less than once a week). Data on some species were also gathered from 14 January to 11 March 1989. Butterflies were captured with an insect net, individually numbered on the underside of the forewings with a felt-tipped pen, and released at the site of capture. The characteristics of each individual (sex, age, site of capture, source of nectar, and other activities) were recorded for later analysis. The age of individual butterflies was estimated in six categories based on wing wear (Ehrlich & Davidson 1960, Brussard & Ehrlich 1970, Ehrlich & Gilbert 1973, Brown et al. 1981, Freitas 1993). Males and females were analyzed separately. Time of residence (permanence) in the population was estimated following Brussard et al. (1974), as days elapsed between marking and last recapture. This parameter is related to the survivorship of the adults.

Rates of flower visitation were recorded during the 12 months of population studies of Heterosais edessa. A plant species was classified as highly visited if 50 or more individuals of H. edessa were observed feeding on its flowers, intermediate if 10 to 49 individuals were observed, and low if fewer than 10 individuals were observed on it during the entire study (as in Freitas 1993).

Results Juvenile Biology of H. edessa Drummond and Brown (1987) recorded Cestrum amictum Schldl. (Solanaceae) as the larval host of//, edessa in coastal Sao Paulo. In the

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FlG. 1. Distribution of body setae on the first instar larva of Heterosais edessa.

Morro do Japui study site, females followed oviposited only on Cestrum laevigatum Schldl. Eggs were laid singly on the underside of old leaves. In general, only one or two eggs were present on each leaf, but as many as six eggs were observed on the same leaf on plants protected from direct sunlight. Although this was not quantified, eggs and larvae were more commonly found on the host plants from April to August.

After hatching, caterpillars ate the entire egg shell. The caterpillars were very sensitive to low humidity and died rapidly in drier environments. Although solitary, larvae were not cannibalistic; several larvae of different instars could be reared together without losses. The caterpillars rested in a J-shaped position on the underside of the leaves. The caterpillars ate all the leaf tissue, even in the first instar. When disturbed, caterpillars could suspend themselves from silk threads, but this behavior was not frequently observed. Pupation usually occurred off the host plant, on neighboring plants but some pupae were found on leaves and stems of C. laevigatum.

Egg. White, ovoid (truncated at bottom) with 23 longitudinal ridges and 11 transverse ridges; diameter 0.7 mm, height 0.6 mm. A line drawing of the egg was published by Brown and Freitas (1994).

First instar. Translucent white with transparent head capsule, becoming pale green after feeding; average width of head capsule 0.39 mm (sd=0.02, n=4); distribution of setae as in Fig. 1. Duration 3—4 days.

Second instar. Pale green with a transparent head capsule, dark intestinal contents evident; average width of head capsule 0.58 mm (sd=0.04, n=ll). Duration 3-4 days.

Third and fourth instars. Similar to previous instar, with progressively darker green pigmentation; head capsule entirely transparent. Fourth instar larvae with narrow lateral yellow stripe (see also Brown & Freitas 1994). Average width of third instar head capsule 0.9 mm (sd=0.04, n = 13); duration 3-4 days. Average width of fourth instar head capsule 1.35 mm (sd=0.05, n = 8); duration 3—4 days.

Fifth instar. Green, with narrow lateral yellow stripe (see Brown & Freitas 1994). Head capsule entirely transparent with dark area in ocellar region. Dorsal vessel clearly

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TABLE 1. Mark-recapture study of Heterosais edessa in Monro do Japui, Sao Paulo, Brazil. L = maximum longevity, M = maximum movement, m = males, f = females, cap = total, recap = total recaptured.

Multiple Recaptures

CAP                     RECAP                                                                                              male;

Month                       ni              f             in             f            Pm            Pf            Mm              Mf 12 3 12

May 91            18        2203—        50        — 2 — — — —

Jun91             11        0        4        0        2      —      100        — 4 — — — —

Jul 91              84      54      17      11      65      21      300      100 14 2 1 11 —

Aug 91            49      33      12        9      23       13      280      100 10 — 2 7 3

Sep 91              7      11        3        6      10      39      100        50 3 — 3 3

Oct 91              2        1        0        0      —      —        —        ______

Nov 91              6        1        0        0      —      —        —        ______

Dec 91              1        0        0        0      —      —        —        ______

Jan 92               3        3        00      —      _______ _

Feb 92              0        0        0        0      —      —        —        ______

Mar 92             0        0        0        0      —      —        —        ______

Apr 92              0        0        0        0      —      —        —        ______

May 92           10        2        1        0      17      —      100        — 1 — — — —

Totals            192    107      37      26      65      39      300      100 32 2 3 21 6

visible. Maximum length 26 mm. Average width of head capsule 2.69 mm (sd=0.21, n=9). Duration 4-6 days.

Pupa. Green, strongly humped (bowed), 0.9 x 1.2 mm; light green first day, acquiring gold stripes after one or two days. A few scattered black dots on wing cases and in ocular area (see Brown & Freitas, 1994). Duration 9 to 11 days.

Adult Ecology of H. edessa

In total, 299 adult H. edessa were captured during 12 months of study, with 63 later recaptured. Fig. 2 shows individuals present per day; when an individual was recaptured it was considered present in the population on all previous days since the day of first capture. Adults were most abundant from May to September, with abundance decreasing after October and maintaining low population levels during the following months (Table 1). During a short capture period, from January to March 1989, 23 individuals of H. edessa were captured on flowers of Adenos-temraa viscosum Forst. (Asteraceae) inside the forest. The population appeared to be breeding continuously, with some peaks (Fig. 2).

The sex ratio of H. edessa in the field deviated from 1:1 (197 males and 107 females marked; X2=24.2, df=l, p<0.001). Males were the dominant sex throughout the study, except in September 1991 and January 1992 (Fig. 3). The proportions of recapture of males (19.3%) and females (24.3%) can be considered equal (;^2=0.71, df=l, p>0.20).

For the analysis of age structure, the six initial age categories were grouped into three: new, intermediate, and old (as in Freitas 1993).

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100%

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; percent of males in

Data for males and females were grouped to increase the total data and enhance perception of age patterns along time. The age structure of H. edessa was dominated by new and intermediate individuals during most of the study. Old individuals appeared mainly after July, with little increase in number during the following months (Fig. 4). No emergence peaks of new individuals were observed throughout the study, and new individuals reached 100% only on days with only one or two individuals captured.

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Fig. 4. Age structure of H. edessa (males and females grouped) in Morro do Japui, May to September 1991: black=fresh (new) individuals, hatched = intermediate, white=old, as percent of total for each day.

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Table 2. Permanance of marked Heterosais edessa: days elapsed between marking and last recapture represent the minimum permanance (MP) for each individual.

1-5 6-10 11-15 16-20 21-25 26-30 31-35 36-40 > 40

16

7 4

3

2

1 1

3

25

12

6

6

7 2 1

2 3

The movement of adults was limited, and the distance between release site and recapture rarely exceeded 100 m. Of the 63 recaptures, 28 males (75.7%) and 23 females (88.5%) were less than 100 m from the site of first capture, and the other 9 males and 3 females were 100-300 m away. Males had a residence time (12.9 days, sd=15.1, n=37) practically equal to females (13.5 days, sd=10.3, n=26) (Table 2).

Nectar sources utilized by H. edessa are available during the entire year, as shown in Table 3. Some individuals were also observed on bird droppings and broken stems of several Eupatorium species. Adults were often encountered near a nectar source, but sometimes they were observed flying low inside the forest or near streams and valleys many times away from any nectar source. Adults were frequently found on flowers of Asteraceae, flying erratically near the ground (rarely over 1 m), like other transparent species of Ithomiinae found in the study area (such as Pseudoscada ernica (Hew.), Oleria aquata (Weymer), Pterony-mia carlia Schaus, Hypoleria adasa (Hew.), Ithomia drymo Hiibner, /. agnosia zikani D'Almeida and /. lichyi D'Almeida).

The Ithomiinae Community in Morro do Japui

Besides H. edessa and the seven species mentioned above, another 12 ithomiine species were present in the study area, including: Melinaea lu-dovica paraiya Reakirt, M. ethra (Godart), Placidula euryanassa (Feld. & Feld.), Methona themisto (Hiibner), Mechanitis I. lysimnia (Fabri-cius), M. polymnia casabranca Haensch, Epityches eupompe (Geyer), Hypothyris ninonia daeta (Boisduval), Dircenna dero celtina (Burmeis-ter), Prittwitzia hymenaea (Prittwitz) (one individual in six years), and Episcada clausina striposis Haensch. All 12 of these species were also marked and recaptured at the same time as H. edessa, but some species were not abundant enough to permit continuous graphs of population dynamics (Fig. 5). Other Ithomiinae found in hills up to 5 km around

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Journal of the Lepidopterists' Society

Table 3. Nectar sources visited by adult Heterosais edessa during 1989-1992 in Morro do Japui, S. Vicente, Brazil. Continuous lines are periods of intense flowering, broken lines indicate few flowers available (1—5 individuals with flowers). Level of visitation of nectar sources: *** = high, ** = intermediate, * = low. Vernonia condensata and Persea americana are introduced plants in the study area.

Nectar source                                                      JFMAM          J          JASOND

ASTERACEAE

Eupatorium vitalhae DC. *                                                        .._____________..

E. laevigatum Lam. ***                                                   ..__________...

E. punctulatum D.C. *                                                                                            .._______..

E. inulaefolium H.B.K. *                                   .._________...

Mikania hindiana D.C. *                                   .._____...

M. rnicrantha H.B.K. *                                           ..____...............

M. cordifolia (L.F.) Willd *                                     .._______......

Trixis antimenorrhoea Mart, ex Baker **                            ..._____...

Vernonia condensata Baker ***                                             .._______..................

V beyrichii (Less.) *                                                 .._______...

V. scorpioides (Lam.) *                                                       ...______________________...

Adenostemma viscosum Forst. ***             _______...............                                ..____

Ageratum conyzoides Linn. *____________________________________________

Bidens pilosa Linn. *____________________________________________

Emilia sonchifolia D.C. *____________________________________________

ROSACEAE

Rubus rosaefolius Smith. ***                                           ..._____________________.........

LAURACEAE

Persea americana Mill **                                                                         .____....

BORAGINACEAE

Cordia verhenacea D.C. *                        ....................____..................................................

the study area were Aeria olena Weymer, Callithomia lenea (Cramer), Thyridia psidii cetoides (Rosenberg & Talbot), Pteronymia euritea (Cramer) and an as yet undescribed subspecies of Pseudoscada quadri-fasciata Talbot. These five species are probably present erratically in the study site, perhaps even as established colonies, since the foodplants are available and the characteristics of the vegetation are the same as in the adjacent forests. As the Sao Paulo coast receives rainfall all year round, no Ithomiinae "pockets" were observed during the six years of constant visits to the study area, but some small aggregations were found near streams on hot summer days.

Sex ratios. The sex ratios of all species in the study site were male-biased, except in those species with very few individuals marked (Table 4). The sex ratios showed few changes from month to month, and only I. drymo showed a tendency for increase in the number of females during the study (Table 5). The two species oiMelinaea (studied in 1989), showed an extreme case of male-biased sex ratio (Table 6). During the

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study no females of Melinaea ethra and only two of M. ludovica were recorded, both flying inside the forest.

Longevity. Permanence in the population (an indirect predictor of life span) was measured only for a few species, due to the low rates of recapture for most. In three species studied in 1991-1992 (M. lysimnia, D. dero and H. edessa) and in the two Melinaea studied in 1989, average permanence was typically more than 10 days (though near this in females of D. dero), except in P. euryanassa, (average less than 10 days; Freitas 1993). In all species studied, maximum permanence was over one month, exceeding two months in some species (Table 7).

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Table 4. Sex ratios of 18 Ithomiinae species captured in Morro do Japui, S. Vicente, Brazil, from 26 May 1991 to 26 May 1992. Asterisks indicate sex ratios significantly different (p<0.05) from 1:1 using chi-square. Percent total = percentage of all captures represented by that species. Days present = number of days a given species was present during the entire study.

Table 5. Sex ratios for four Ithomiinae species marked in Morro do Japui, given as percent (boldface) of males in the total (T) for each month.

Table 6. Populational parameters for Melinaea ludovica and M. ethra, 14 January 1989 to 11 March 1989. TM = total marked, TC = total captured, MOV = maximum movement (meters), PER = maximum permanance (days), x = average permanance (with standard deviation), m = males, f = females. All males collected were feeding on Adenos-temrria viscosum.

TM Multiple recaptures __________TR           MOV         PER__________________________________

f               in               m               in                        x                               12               3             4

M. ludovica 113 2 32 300 55 14.1 ± 14.1 25 5 2 0 M. ethra 115 0 33 300 47 13.7 ± 12.9 25 4 2 2

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TABLE 7. Permanence in the population for four species of Ithomiinae studied in Morro do Japui, 1991-1992. x = average permanance (days, with standard deviation).

Species/sex                                                        x                                      range                                        SD                                      N

P. euryanassa males P. euryanassa females M. lysimnia males M. lysimnia female D. dero males D. dero females H. edessa males H. edessa females

nales

All species studied showed a peak of abundance between July and August (Figs. 2, 5; see also Freitas 1993 for abundance graphs of P. euryanassa), except the two Melinaea species, which were most abundant from November to March (summer months); in 1991—1992 they were not as abundant as in 1989 and 1990.

Food sources. Adult Ithomiinae were usually observed on flowers, especially of Asteraceae (Table 8). Nectar sources in the study site were available all year round (Table 3). All species showed a preference for flowers in the Asteraceae, except the transparent blue Ithomiinae species (H. edessa, I. dry mo, I. lichyi, I. agnosia, P. erruca, P. carlia and O. aquata), that prefered flowers of Rubus rosaefolius (Rosaceae), all year (Table 8; other transparent blue species not present in the Table were also captured mostly on these flowers: H. adasa, 17 of 22 feeding records; P. carlia, 12 of 23; O. aquata, 6 of 7). The apparent strong pref-

TABLE 8. Frequency of flower visitation for 8 species of Ithomiinae studied in Morro do Japui, 26 May 1991 to 16 May 1992. Plants with an asterisk are known PA sources. Bold numbers indicate that visitation is male biased. Results presented as males/females. See Table 3 and text for full names of plants and butterflies {A.cura. = Asclepias curassavica).

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Journal of the Lefidopterists' Society

erence of P. euryanassa for flowers of Persea (Table 3) may have been due to chance, since this seemed to be the most abundant flower during the population peak of this species (Freitas, 1993). In general, the records of males and females were near 1:1 on all nectar sources. On E. laevigatum males were more abundant for most of the species. In 1989, Adenostemma viscosum also attracted mainly males of Melinaea (Table 6), as well as H. edessa (25 males and no females) and O. aquata (21 males and 1 female). Adult Ithomiinae individuals of both sexes were also observed puddling on mud, and feeding on bird droppings, rotting fruits, and broken stems of several Eupatorium species.

Species composition. Data on relative species abundance were analyzed only for the period from 26 May to 23 November 1991, when individuals were most abundant and the censuses were more frequent. Throughout this period, four species (P. euryanassa, H. edessa, M. ly-simnia and D. dero) accounted for at least 70% of the captured Ithomiinae on any day (see also Table 4 and Fig. 6A). The number of species captured per day varied from 2 to 13 (mean 6.96, sd=2.6, n=54), with at least 6 species present on most days (Fig. 6B). The most abundant species were also the ones that were seen on most days (Table 4), showing that they were well distributed throughout the year (see also Fig. 5, and Freitas 1993). The days with more than six species captured were mostly in July and August, also the period when the largest number of individuals were seen per day (Fig. 6B). The abundance of some species varied from year to year: Ithomia lichyi was very abundant in 1989 and 1990, but /. agnosia and E. clausina were always sparse from 1988 to 1993. Some abundant species may be rare in other years: Pseudoscada erruca, a moderately common species from 1988 to 1993, was practically absent in the first part of 1994 in Morro do Japui, although it continued to be common in other forests near the study area.

Discussion

Biology of H. edessa

Heterosais edessa has immatures typical for its tribe (Godyridini), with solitary larvae, little pigment in the larval cuticle including cephalic capsule, and a bent, green pupae; all these are considered derived traits in the Ithomiinae (Brown & Freitas 1994). Although larvae are solitary, they do not exhibit cannibalistic habits, common in solitary larvae of Heliconiini (Brown 1981 and pers. obs.). Several other Ithomiinae with solitary larvae were also not aggressive towards other smaller larvae (pers. obs.); this trait may have allowed the development of gregarious behaviour in Ithomiinae larvae. In fact, gregarious larvae are considered to be a polyphyletic trait in Ithomiinae, appearing erratically in many Ithomiinae radiations (see Brown & Freitas 1994).

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Fig. 6. Upper: species ratio of Ithomiinae captured in Morro do Japui from May to November 1991. Bottom: total of individuals of all Ithomiinae captured in Morro do Japui from May to November 1991 (filled circles) and number of Ithomiinae species captured per day in the same period (open circles).

The general behavior of H. edessa larvae, like the J-shaped resting position and the suspension by silken threads when disturbed, is very similar to that of other known Ithomiinae (Brown & D'Almeida 1970, Young 1972, 1974a, 1974b, 1974c, 1978a, 1978b, Ackery 1987). Larvae of H. edessa are basically cryptic in color pattern and behavior.

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The Ithomiinae Community of Morro do japui

Although more abundant in the winter months, H. edessa is found in Japui all year round; the low numbers in some months may be due mostly to the low number of plants in bloom. This pattern of fluctuation seems to be present in varying degrees in most Ithomiinae (Vasconcellos-Neto 1980, Freitas 1993). The vagility of//, edessa is low, with individuals usually tracking their nectar sources. Because the larval food plants are present throughout the area, females do not need to search widely to find them, so both male and female movement is more related to nectar sources.

Sex ratio was male biased in H. edessa and seven other ithomiine species. Male biased sex ratios in the field have been reported for many species of butterflies, even when sex ratio in the laboratory was 1:1 (Brussard & Ehrlich 1970, Brussard et al. 1974, Brown & Ehrlich 1980, Matsumoto 1984, 1985, Freitas 1993). In the Ithomiinae, the predominance of males in field collections has been attributed to their appearance at sources of pyrrolizidine alkaloids (PAs) (Brown 1985, Trigo 1988, Freitas 1993), as seen with E. laevigatum in this study. The fact that male biased sex ratios were also detected on other plants (e.g., R. rosae-folius) may be explained by the existence of behavioral differences between males and females (Ehrlich 1984), resulting in males generally flying in the same places as the collectors, with females more dispersed throughout the habitat.

The residence times of H. edessa and the other Ithomiinae in the study area were relatively high compared to species in other butterfly families, but were lower than those for species of Heliconius in the same area (Rogner & Freitas, unpubl. data). Some individuals may survive for more than two months, and two females older than 30 days were observed ovipositing and with apparently distended abdomens, indicating continual egg production as in Heliconius species (Dunlap-Pianka et al. 1977). The relatively short lifespan of Ithomiinae in this study site in relation to that in other regions (Vasconcellos-Neto 1980, Brown 1985) may be due to the continuously humid climate, and concomitant lack of reproductive diapause in this region. The age structure of adult H. edessa shows that the population has continuous recruitment of new individuals throughout the year, though probably diminished from July to December. These data suggest that H. edessa, like most other Ithomiinae in the study site, is more K-selected (Pianka 1970) than Placidula euryanassa (Freitas 1993).

The Ithomiinae community in Morro do Japui is typical for the Sao Paulo coastal region. The absence of montane species results in a low richness in relation to the interior (28 species in the "Serra do Japi," in-

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terior of Sao Paulo; Brown 1992), although some species like H. edessa are absent in the interior, or occur only as migrants from the coast (the two Melinaea and O. aquata). Also, several species in the coastal Atlantic transparent yellow complex (Brown & Benson 1974) are absent in the region (like Scada karschina), appearing only 100 km east of Sao Vicente in Sao Paulo state (Sao Sebastiao municipality).

The number of species captured per day throughout the year was almost always more than six, reaching 13 on some days, with high numbers of individuals, reflecting a relatively high diversity. Eleven species are common enough to be captured on almost any day in the study site. A relatively complete list of Ithomiinae of a given coastal Sao Paulo site may be obtained in four to five days of intense work from June to September, especially with the use of Heliotropium indicum baits to attract males of rarer species, or by searching for the right flowers (habitat "pockets" that concentrate adult Ithomiinae in winter are virtually absent in most of the Sao Paulo coastal region; see also Brown 1972). A survey of the 26 Ithomiinae species of Sao Sebastiao was made in only seven collecting trips from May to September 1992.

The data presented here provide valuable information for two kinds of community studies. Firstly, the study of plant-based food webs is of great importance in the understanding of tropical ecosystems. For example, the data obtained for a given taxonomic group of herbivorous insects can be useful in studies of diversity and health of a given plant assemblage. This is especially true for groups like Ithomiinae which are rapidly sampled and depend on a diverse set of plants for larval and adult hosts (Gilbert 1980). Ithomiinae have been suggested as especially good indicator organisms for this kind of study (Brown 1991, Beccaloni & Gaston 1995). Secondly, an understanding of the seasonal patterns of each species, as well as their behavior, habits and preferences can lead to greater efficiency in collecting data in a given area, especially if many areas must be surveyed in little time. Some sites are difficult to reach, so that the knowledge of which months are better to census butterflies is valuable to those who intend to carry out inventory and monitoring.

The present paper attempts to define some of these patterns for the Sao Paulo coastal forests, to help future work in this region. Similar work is needed in other areas to reveal the different patterns, especially in places with marked seasonality or with different climatic regimes.

Acknowledgments

This research was supported by the Fundacao de Amparo a Pesquisa do Estado de Sao Paulo (FAPESP) (Fellowship 90-0082-3). I thank K. S. Brown Jr., L. E. Gilbert, P. S. Oliveira, R. B. Francini, J. R. Trigo, A. Gomes-Filho, J. R. Lima and two anonymous reviewers for comments on the manuscript. R. B. Francini and C. M. T. Apostolo helped during field and laboratory work.

288

Journal of the Lepidopterists' Society

Literature Cited

ACKERY, P. R. 1987. The danaid genus Tellervo (Lepidoptera: Nyinphalidae): a cladistic

approach. Zool. J. Linn. Soc. 89:293-274. Bates, H. W. 1862. Contributions to an insect fauna of the Amazon Valley, Lepidoptera:

Heliconidae. Trans. Linn. Soc. Lond. 23:495-566. Beccaloni, G. W. & K. J. Gaston. 1995. Predicting the species richness of neotropical

forest butterflies: Ithomiinae (Lepidoptera: Nymphalidae) as indicators. Biol.

Cons.71:77-86. Brown, I. L. & P. R. Ehhlich. 1980. Population biology of the checkerspot butterfly, Eu-

phydryas chalcedona. Structure of the Jasper Ridge colony. Oecologia 47:239—251. Brown, K. S. Jr. 1972. Maximizing daily butterfly counts. J. Lepid. Soc. 26:183-196. ----------. 1981. The biology of Heliconius and related genera. Ann. Rev. Entomol.

26:427-456. ----------. 1985. Chemical ecology of dehydropyrrolizidine alkaloids in adult Ithomiinae

(Lepidoptera: Nymphalidae). Rev. Bras. Bio. 44:435-460. ----------. 1991. Conservation of neotropical environments: insects as indicators, pp.

499—504. In Collins, N. M. & J. A. Thomas (eds.), Conservation of insects and their

habitats. Academic Press, London. ----------. 1992. Borboletas da Serra do Japi: diversidade, habitats, recursos alimentares e

variacao temporal, pp. 142-186. In Morellato, L. P. C. (Org.), Historia Natural da

Serra do Japf. Ecologia e preservacao de uma area florestal no Sudeste do Brasil, Edi-

tora da UNICAMP, Campinas, SP. Brown, K. S. Jr. & R. F. D'almeida. 1970. The Ithomiinae of Brazil (Lepidoptera:

Nymphalidae). II. A new genus and species of Ithomiinae with comments on the tribe

Dircennini D'Almeida. Trans. Am. Entomol. Soc. 96:1—18. Brown, K. S. Jr. & W. W. BENSON. 1974. Adaptive polymorphism associated with multiple mullerian mimicry in Heliconius numata (Lepid. Nymph.). Biotropica 6:205—228. Brown, K. S. Jr. & A. V. L. FREITAS. 1994. Juvenile stages of Ithomiinae: overview and

systematics (Lepidoptera: Nymphalidae). Trop. Lepid. 5:9—20. Brown, K. S. Jr., A. J. Damman & P. Feeny. 1981. Troidine swallowtails (Lepidoptera:

Papilionidae) in southeastern Brazil: natural history and foodplant relationships. J.

Res. Lepid. 19:199-226. Brussard, P. F. & P. R. Ehrlich. 1970. The population structure of Erebia epipsodea

(Lepidoptera: Satyrinae). Ecology 51:119—129. Brussard, P. F, P. R. Ehrlich & M. C. Sincer. 1974. Adult movements and population

structure in Euphydryas editha. Evolution 28:408-415. Devries, P. J. 1994. Patterns of butterfly diversity and promising topics in natural history

and ecology, pp. 187-194. In McDade, L. A., K. S. Bawa, H. A. Hespenheide & G. S.

Hartshorn (eds.), La Selva. Ecology and natural history of a neotropical rain forest.

Univ. Chicago Press, Chicago. DRUMMOND, B. A. III. 1976. Comparative ecology and mimetic relationships of ithomiine

butterflies in eastern Ecuador. Unpubl. Ph. D. Thesis, Univ. Florida, xvi + 361 pp. Drummond, B. A. Ill & K. S. Brown Jr. 1987. Ithomiinae (Lepid.: Nymphalidae): summary of known larval foodplants. Ann. Missouri Bot. Garden 74:341—358. Dunlap-Pianka, H., C. L. Boggs & L. E. GILBERT. 1977. Ovarian dynamics in heliconi-

ine butterflies: programmed senescence versus eternal youth. Science 197:487—490. Ehrlich, P. R. 1984. The structure and dynamics of butterfly populations, pp. 25—40. In

Vane-Wright, R. I. & P. R. Ackery (eds.), The biology of butterflies. Academic Press,

London. EHRLICH, P. R. & S. E. Davidson. 1960. Techniques for capture-recapture studies of

Lepidoptera populations. J. Lepid. Soc. 14:227—229. Ehrlich, P. R. & L. E. Gilbert 1973. Population structure and dynamics of the tropical

butterfly Heliconius ethilla. Biotropica 5:69—82. FREITAS, A. V. L. 1991. Variacao morfologica, ciclo de vida e sistematica de Tegosa claud-

ina (Eschscholtz) (Lepidoptera, Nymphalidae, Melitaeinae) no Estado de Sao Paulo.

Rev. Bras. Entomol. 35:301-306.

Volume 50, Number 4

289

----------. 1993. Biology and population dynamics of Placidula euryanassa, a relict ithomiine

butterfly (Nymphalidae: Ithomiinae). J. Lepid. Soc. 47:87-105.

FREITAS, A. V. L. & P. S. Oliveira. 1992. Biology and behavior of the neotropical butterfly Eunica bechina (Nymphalidae) with special reference to larval defence against ant predation. J. Res. Lepid. 31:1—11.

GILBERT, L. E. 1980. Food web organization and the conservation of neotropical diversity, pp. 11-33. In Soule, M. E. & B. A. Wilcox (eds.), Conservation biology: an evolutionary-ecological perspective. Sinauer, Sunderland, Massachusetts.

HABER, W. 1978. Evolutionary ecology of tropical mimetic butterflies (Lepidoptera: Ithomiinae). Unpubl. Ph. D. Dissertation, Univ. Minnesota.

MATSUMOTO, K. 1984. Population dynamics of Luehdorfia japonica Leech (Lepidoptera: Papilionidae). I. A preliminary study on the adult population. Res. Popul. Ecol. 26:1-12.

----------. 1985. Population dynamics of the Japanese clouded apollo Parnassius glacialis

Butler (Lepidoptera: Papilionidae). I. Changes in population size and related population parameters for three successive generations. Res. Popul. Ecol. 27:301—312.

MOLLER, F. 1879. Ituna and Thyridia; a remarkable case of mimicry in butterflies (translated by R. Meldola). Proc. Entomol. Soc. London: 20-29.

NiMER, E. 1972. Climatologia da Regiao sudeste do Brasil. Introducao a climatologia dinamica. Rev. Bras. Geogr. 34:3—48.

Pianka, E. R. 1970. On r- and K-selection. Am. Nat. 104:592-597.

PRODESAN, A. 1969. Estudo da viabilidade do distrito industrial de Santos. 91 pp.

Setzer, J. 1949. Contribuicao para o estudo do clima do estado de Sao Paulo. Edit. Esco-las Profissionais Salesianas, Sao Paulo. 239 pp.

TRIGO, J. R. 1988. Ecologia quimica na interacao Ithomiinae (Lepidoptera: Nymphalidae )/Echitoideae (Angiospermae: Apocynaceae). Unpubl. M. S. Thesis, Univ. Estad. de Campinas, Campinas, SP. xiv + 199 pp.

Ururahy, J. C, J. E. R. Collares, M. M. Santos, & R. A. A. Barreto. 1987. 4. Vege-tacao. In Projeto RADAMBRASIL. Vol. 32. fls. sf 23-24 (Rio de Janeiro e Vitoria). Edit Ministerio das Minas e Energia, Brasilia, DF.

Vasconcellos-Neto, J. 1980. Dinamica de Populacoes de [thomiinae (Lepidoptera: Nymphalidae) em Sumare-SP. Unpubl. M.S. Thesis, Univ. Estad. de Campinas, Campinas, SP. vi + 206 pp.

----------. 1986. Interactions between Ithomiinae (Lepidoptera: Nymphalidae) and

Solanaceae, pp. 366-377. In D'Arcy, W. G. (ed.), Solanaceae, biology and systematics. Columbia Univ. Press, New York.

----------. 1991. Interactions between ithomiine butterflies and Solanaceae: feeding and reproductive strategies, pp. 291—313. In Price, P. W., T M. Lewinsohn, G. W. Fernan-des & W. W. Benson (eds.), Plant-animal interactions. Evolutionary ecology in tropical and temperate regions. John Wiley & Sons, Inc., New York.

YOUNG. A. M. 1972. On the life cycle and natural history of Hymenitis nero (Lepidoptera: Ithomiinae) in Costa Rica. Psyche 79:284-294.

----------. 1974a. On the biology of Godyris zavaleta caesiopicta (Lepidoptera: Nymphalidae: Ithomiinae). Entomol. News 85:227-238.

----------. 1974b. A natural historical account of Oleria zelica pagasa (Lepidoptera:

Nymphalidae: Ithomiinae) in a Costa Rican mountain rain forest. Stud. Neotrop. Fauna 9:123-140.

----------. 1974c. Notes on the biology of Pteronymia notilla (Ithomiidae) in a Costa Rican

mountain forest. J. Lepid. Soc. 28:257-268.

----------. 1978a. The biology of the butterfly Aeria eurimedia agna (Nymphalidae: Ithomiinae: Oleriini) in Costa Rica. J. Kansas Entomol. Soc. 51:1—10.

----------. 1978b. Notes on the biology of the butterfly Hypoleria cassotis (Bates) (Nymphalidae: Ithomiinae) in northeastern Costa Rica. Brenesia 14-15:97-108.

Young. A. M. & M. W Moffett. 1979. Studies on the population biology of the tropical butterfly Mechanitis isthmia in Costa Rica. Am. Midi. Nat. 101:309-319.

Received for publication 14 May 1995; revised and accepted 12 November 1995.