The arms race between heliconiine butterflies and Passiflora plants – new insights on an ancient subject

Heliconiines are called passion vine butterflies because they feed exclusively on Passiflora plants during the larval stage. Many features of Passiflora and heliconiines indicate that they have radiated and speciated in association with each other, and therefore this model system was one of the first examples used to exemplify coevolution theory. Three major adaptations of Passiflora plants supported arguments in favour of their coevolution with heliconiines: unusual variation of leaf shape within the genus; the occurrence of yellow structures mimicking heliconiine eggs; and their extensive diversity of defence compounds called cyanogenic glucosides. However, the protection systems of Passiflora plants go beyond these three features. Trichomes, mimicry of pathogen infection through variegation, and production of extrafloral nectar to attract ants and other predators of their herbivores, are morphological defences reported in this plant genus. Moreover, Passiflora plants are well protected chemically, not only by cyanogenic glucosides, but also by other compounds such as alkaloids, flavonoids, saponins, tannins and phenolics. Heliconiines can synthesize cyanogenic glucosides themselves, and their ability to handle these compounds was probably one of the most crucial adaptations that allowed the ancestor of these butterflies to feed on Passiflora plants. Indeed, it has been shown that Heliconius larvae can sequester cyanogenic glucosides and alkaloids from their host plants and utilize them for their own benefit. Recently, it was discovered that Heliconius adults have highly accurate visual and chemosensory systems, and the expansion of brain structures that can process such information allows them to memorize shapes and display elaborate pre‐oviposition behaviour in order to defeat visual barriers evolved by Passiflora species. Even though the heliconiine–Passiflora model system has been intensively studied, the forces driving host‐plant preference in these butterflies remain unclear. New studies have shown that host‐plant preference seems to be genetically controlled, but in many species there is some plasticity in this choice and preferences can even be induced. Although much knowledge regarding the coevolution of Passiflora plants and heliconiine butterflies has accumulated in recent decades, there remain many exciting unanswered questions concerning this model system.     

Sequestration and biosynthesis of cyanogenic glucosides in passion vine butterflies and consequences for the diversification of their host plants

The colorful heliconiine butterflies are distasteful to predators due to their content of defense compounds called cyanogenic glucosides (CNglcs), which they biosynthesize from aliphatic amino acids. Heliconiine larvae feed exclusively on Passiflora plants where ~30 kinds of CNglcs have been reported. Among them, some CNglcs derived from cyclopentenyl glycine can be sequestered by some Heliconius species. In order to understand the evolution of biosynthesis and sequestration of CNglcs in these butterflies and its consequences for their arms race with Passiflora plants, we analyzed the CNglc distribution in selected heliconiine and Passiflora species. Sequestration of cyclopentenyl CNglcs is not an exclusive trait of Heliconius, since these compounds were present in other heliconiines such as Philaethria, Dryas and Agraulis, and in more distantly related genera Cethosia and Euptoieta. Thus, it is likely that the ability to sequester cyclopentenyl CNglcs arose in an ancestor of the Heliconiinae subfamily. Biosynthesis of aliphatic CNglcs is widespread in these butterflies, although some species from the sara‐sapho group seem to have lost this ability. The CNglc distribution within Passiflora suggests that they might have diversified their cyanogenic profile to escape heliconiine herbivory. This systematic analysis improves our understanding on the evolution of cyanogenesis in the heliconiine–Passiflora system.     

Phenotypic plasticity in chemical defence of butterflies allows usage of diverse host plants

Host plant specialization is a major force driving ecological niche partitioning and diversification in insect herbivores. The cyanogenic defences of Passiflora plants keep most herbivores at bay, but not the larvae of Heliconius butterflies, which can both sequester and biosynthesize cyanogenic compounds. Here, we demonstrate that both Heliconius cydno chioneus and H. melpomene rosina have remarkable plasticity in their chemical defences. When feeding on Passiflora species with cyanogenic compounds that they can readily sequester, both species downregulate the biosynthesis of these compounds. By contrast, when fed on Passiflora plants that do not contain cyanogenic glucosides that can be sequestered, both species increase biosynthesis. This biochemical plasticity comes at a fitness cost for the more specialist H. m. rosina, as adult size and weight for this species negatively correlate with biosynthesis levels, but not for the more generalist H. c. chioneus. By contrast, H. m rosina has increased performance when sequestration is possible on its specialized host plant. In summary, phenotypic plasticity in biochemical responses to different host plants offers these butterflies the ability to widen their range of potential hosts within the Passiflora genus, while maintaining their chemical defences.     

Origin of the cultivated passion fruit Passiflora edulis f. flavicarpa and genomic relationships among species of the subgenera Decaloba and Passiflora

• Passiflora edulis f. flavicarpa is the most economically important species in the genus Passiflora. However, the origin of this yellow form of passion fruit remains unclear, being suggested as a hybrid (P. edulis f. edulis × P. ligularis) or wild mutant.
• Here, the origin and genomic relationships of P. edulis f. flavicarpa with some related species in the genus Passiflora (subgenera Decaloba and Passiflora) were investigated using genomic in situ hybridization (GISH). Genomic DNA of 18 species was used as probe, which was hybridized onto chromosomes of P. edulis f. flavicarpa.
• Of all genomic DNA probes tested, none allowed us to identify a specific chromosome set in P. edulis f. flavicarpa. Conversely, probes from the subgenus Passiflora, P. edulis f. edulis, P. alata, P. cincinnata, P. coccinea, P. nitida and P. vitifolia, produced intense and uniform hybridizations on all chromosomes of P. edulis f. flavicarpa. Moreover, probes from P. ligularis, P. foetida and P. sublanceolata produced more intense hybridizations in the terminal region of four chromosomes, corresponding to the DNAr 45S locus, and also dispersed, less intense, hybridization across all chromosomes. Probes from the subgenus Decaloba, P. biflora, P. capsularis, P. cervii, P. coriacea, P. micropetala, P. morifolia, P. rubra and P. suberosa, produced hybridizations restricted to the DNAr 45S sites.
• The hybrid origin of P. edulis f. flavicarpa could not be supported based on the GISH results, and it is suggested that this species is conspecific with P. edulis f. edulis, because the probe with DNA of this form hybridized strongly throughout the target genome. The other putative parent species, P. ligularis, showed only a distant relationship with the target genome. The results also suggest that species of the subgenus Passiflora share many repetitive sequences and that the relationship between subgenera Decaloba and Passiflora is very distant.

     

Male pheromone composition depends on larval but not adult diet in Heliconius melpomene

1. Condition‐dependent traits can act as honest signals of mate quality, with fitter individuals being able to display preferred phenotypes. Nutrition is known to be an important determinant of individual condition, with diet known to affect many secondary sexual traits. 2. In Heliconius butterflies, male chemical signalling plays an important role in female mate choice. Potential male sex pheromone components have been identified previously, although it is unclear what information they convey to the female. 3. In the present study, the effect of diet on androconial and genital compound production is tested in male Heliconius melpomene rosina. To manipulate larval diet, larvae are reared on three different Passiflora host plants: Passiflora menispermifolia, the preferred host plant, Passiflora vitifolia and Passiflora platyloba. To manipulate adult diet, adult butterflies are reared with and without access to pollen, a key component of their diet. 4. No evidence is found to suggest that adult pollen consumption affects compound production in the first 10 days after eclosion. There is also a strong overlap in the chemical profiles of individuals reared on different larval host plants. The most abundant compounds produced by the butterflies do not differ between host plant groups. However, some compounds found in small amounts differ both qualitatively and quantitatively. Some of these compounds are predicted to be of plant origin and the others synthesised by the butterfly. Further electrophysiological and behavioural experiments will be needed to determine the biological significance of these differences.     

Feeding behavior of Heliconius erato phyllis (Fabricius) (Lepidoptera: Nymphalidae) larvae on passion vines

Of the 15 species of passion vines recorded for the Rio Grande do Sul State, Brazil, nine are used by Heliconius erato phyllis (Fabricius, 1775). The larvae of this species feed preferentially on Passiflora misera H.B.K., which confers greater performance despite presenting smaller amounts of nutrients than other host plants. Thus, the performance associated with the consumption of a passion vine is possibly related not only to its nutritional content but also to the morphological and behavioral mechanisms involved in the feeding. In this study, the difficulties in accessing food imposed by the hosts (P. misera, Passiflora suberosa L., Passiflora caerulea L., Passiflora edulis Sims, and Passiflora alata Dryand.) were evaluated. Focal observations were performed every 2 min during 6-h sessions (sequential sampling), and the relative time spent for different behaviors (resting, feeding, walking, tasting, and vein cutting) was quantified on both young and old leaves. Larvae devoted more time feeding on P. misera in most cases. Larvae observed on P. alata devoted more to time resting and less time feeding, performing one or two meals of small duration, every 6 h. First instar caterpillars on old leaves of P. suberosa and P. caerulea spent more time walking, searching for a favorable feeding site. The hardness of leaves may be a limiting factor for the initial instars in this heliconian.     

Determination of antioxidant activity and phenolic content of extracts from in vivo plants and in vitro materials of Passiflora alata Curtis

Passiflora alata Curtis, commonly known as sweet passion fruit, is one of the commercially cultivated species of the genus Passiflora, whose fruits can be consumed in natura due to their sweet taste. It is also used worldwide as an ornamental and in folk medicine. The goal of this work was the evaluation of the antioxidant potential of extracts from in vivo plants, and in vitro-derived materials of P. alata. Leaves from in vivo plants were used for the optimization of parameters that affect the efficiency of extraction of antioxidant compounds (proportions of ethanol:water, maceration period, solvent:plant tissue ratio, and number of extraction stages), by employing the 2,2-diphenyl-1-picrylhydrazyl (DPPH) assay. The antioxidant activity and the extract yields were significantly influenced by the proportion of ethanol:water and maceration period. The optimized protocol was applied to obtain the extracts of in vitro-derived materials. Total phenolic content was determined using the Folin–Ciocalteu method. Higher antioxidant activities and phenolic contents were observed in extracts from leaves of in vivo-seed derived and from acclimatized plants when compared to in vitro plants, calluses and suspension cultures. Differences in the reaction kinetics of DPPH scavenging activity were also observed.     

Influence of method of presentation on results of plant-host preference tests with two species of grasshopper

Leaf material of two Piper spp. (Piperaceae) was presented to two different generalist herbivores (Orthoptera: Acrididae) to determine whether method of leaf presentation in preference trials affects choice. For the grasshopper Leptomerinthoprora brevipennis, the method of presentation had no effect on preference ranking: more of Piper sancti-felicis was eaten than of P. arieianum, regardless of whether leaf material was presented as leaf disks, whole leaves supplemented with water, or whole leaves without water. In contrast, preference of the grasshopper Abracis flavolineata depended on presentation method. In that case, whole leaves of P. scancti-felicis supplemented with water were preferred over the same of P. arieianum, while dry weight consumption of leaf disks of P. arieianum was greater than that for disks of P. sancti-felicis. When presented with leaves not supplemented with water, A. flavolineata showed no significant preference for one plant species over another.

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Résumé Des feuilles provenant de deux espèces de Piper (Piperacae) ont été proposées à deux Orthoptères, Acrididae, herbivores généralistes, pour déterminer si la méthode de présentation des feuilles modifiait le choix. Pour Leptomerinthoprora brevipennis, la méthode de présentation est sans influence sur l'ordre de préférence; la consommation de Piper sancti-felicis est toujours supérieure à celle de P. arieianum, quelque soit le mode de présentation: disques de feuilles, feuilles entières humidifiées, ou feuilles entières sans addition d'eau. A l'opposé, l'ordre de préférence de Abracis flavolineata dépend du type de présentation; les feuilles entières humidifiées de P. sancti-felicis sont préférées à celles de P. arieianum proposées de la même façon, tandis que la consommation de disques secs de P. arieianum est plus importante que celle des disques secs de P. sancti-felicis. En présence de feuilles entières humidifiées, A. flavolineata ne manifeste aucune préférence.

     

Ecological and genetic factors influencing the transition between host‐use strategies in sympatric Heliconius butterflies

Shifts in host‐plant use by phytophagous insects have played a central role in their diversification. Evolving host‐use strategies will reflect a trade‐off between selection pressures. The ecological niche of herbivorous insects is partitioned along several dimensions, and if populations remain in contact, recombination will break down associations between relevant loci. As such, genetic architecture can profoundly affect the coordinated divergence of traits and subsequently the ability to exploit novel habitats. The closely related species Heliconius cydno and H. melpomene differ in mimetic colour pattern, habitat and host‐plant use. We investigate the selection pressures and genetic basis underlying host‐use differences in these two species. Host‐plant surveys reveal that H. melpomene specializes on a single species of Passiflora. This is also true for the majority of other Heliconius species in secondary growth forest at our study site, as expected under a model of interspecific competition. In contrast, H. cydno, which uses closed‐forest habitats where both Heliconius and Passiflora are less common, appears not to be restricted by competition and uses a broad selection of the available Passiflora. However, other selection pressures are likely involved, and field experiments reveal that early larval survival of both butterfly species is highest on Passiflora menispermifolia, but most markedly so for H. melpomene, the specialist on that host. Finally, we demonstrate an association between host‐plant acceptance and colour pattern amongst interspecific hybrids, suggesting that major loci underlying these important ecological traits are physically linked in the genome. Together, our results reveal ecological and genetic associations between shifts in habitat, host use and mimetic colour pattern that have likely facilitated both speciation and coexistence.     

A REVISION OF PASSIFLORA SECTION DYSOSMIA

Revision of Passiflora subgenus Passiflora section Dysosmia DC. (Passifloraceae). The twenty one species and 10 varieties (31 taxa) comprising subgenus Passiflora section Dysosmia DC are described and some are illustrated with photographs and line drawings. Two species, Passiflora vesicaria L. and Passiflora ciliata (Dryand) Mast., are removed from synonymy and reinstated to species rank. Passiflora baraquiniana Lemaire is removed from synonymy and given varietal status. The species Passiflora santiagana (Killip) Borhidi is reduced in rank to varietal status. Of the 38 varieties of Passiflora foetida L. described by E. P. Killip in his 1938 monograph, 28 are reduced to synonymy, four are reassigned as varieties of other species and four remain as varieties of P. foetida. A new variety, Passiflora foetida var. ellisonii Vanderplank, is described with photographs and line drawings.     

Freshwater crabs (Decapoda: Pseudothelphusidae) increase rates of leaf breakdown in a neotropical headwater stream

1. Freshwater crabs are the largest macroconsumers in many neotropical headwater streams, but few studies have examined their roles in ecosystem processes such as leaf litter breakdown. As omnivorous macroconsumers, freshwater crabs affect multiple trophic levels. They may directly increase leaf breakdown through fragmentation and consumption or indirectly decrease breakdown by consuming other macroinvertebrates, including shredders and detritivores.
2. In a headwater stream in Monteverde, Costa Rica, we conducted an in-stream experiment with 40 enclosures to quantify the effects of pseudothelphusid crabs on both leaf breakdown and macroinvertebrate colonisation of leaves. Half of the enclosures were randomly selected to contain two crabs (mean carapace width = 30 mm) and half were controls without crabs. We sampled mixed leaf packs from the enclosures on days 11, 19, 28, 34, and 42. We found the leaves of one species (Koanophyllon pittieri) almost completely decomposed by day 28 in both treatments (crab versus no crab). The other two leaf species (Meliosma idiopoda, Quercus brenesii) composed the remaining leaf mass at the end of the experiment.
3. At 42 days, enclosures with crabs had faster rates of leaf breakdown than those without crabs (with crabs: k = −0.020; without crabs: k = −0.016; p = 0.034). This suggests that the magnitude of direct leaf breakdown by crabs, due to fragmentation, consumption, or manipulation of leaves, was greater than any indirect effects on leaf breakdown via crab consumption of other leaf-consuming species.
4. Macroinvertebrate composition based on taxa abundances or biomasses did not significantly differ between treatments (ANOSIM; p = 0.73 and p = 0.65, respectively). Shredder and detritivore abundances and biomasses increased significantly through time (ANOVA; p ≤ 0.001), but there was no evidence of an effect of crab presence (p > 0.2), nor were there significant interactions between crab presence and time (p > 0.3).
5. This is one of the first studies to quantify the effects of pseudothelphusid freshwater crabs on leaf breakdown rates. Our results suggest that these crabs can play a significant role in detrital processing in neotropical headwater streams. This study has also demonstrated that short-term enclosure experiments are useful in measuring in-stream effects of crab activity on leaf breakdown.

     

Invasive plant population and herbivore identity affect latex induction

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Comparison between two methods to measure herbivory. Is herbivory in the neotropics more severe than we thought?

The quantity of plant tissue consumed by herbivores can be recorded 1) by measuring herbivory in previously marked leaves or 2) by performing punctual measures. i.e. selecting leaves at random to measure the tissue absent in each leaf Punctual measurements are frequently used because they are a faster method to estimate herbivory. However, punctual measures do not include totally consumed leaves. therefore they underestimate the actual herbivory rates. In three species of understory shrubs (Palicourea sp. P. angustifolia and P. oralis: Rubiaccae) herbivory was measured using punctual measures and marking young leaves in order to determine the degree of herbivory underestimation by punctual measurements. Punctual measurements underestimated herbivory up to three times in the species with a high number of totally consumed leaves. In the species with a lower number of totally consumed leaves, herbivory rates recorded using both methods were similar. These results suggest that herbivory in neotropical forests could be more severe than what is currently suggested.     

Passiflora tacanensis, a new species of Passiflora subgenus Decaloba supersection Cieca from Mexico

Passiflora tacanensis, a new species from southern Mexico, is described and illustrated. It was discovered during the course of a taxonomic revision ofPassiflora subgenusDecaloba supersectionCieca and is assigned to that group of apetalous passion flowers.Passiflora tacanensis is known only from one fruiting herbarium specimen that was collected in southern Mexico on Volcán Tacaná in a high mesophytic forest. It is distinguished from its closest relatives,P. coriacea, P. sexocellata, P. mcvaughiana, andP. xiikzodz, by its foliose stipules, leaves that lack laminar nectaries but possess petiolar nectaries, and wide seeds with a chalazal beak that is not well developed.     

Predation and avoidance of tough leaves by aquatic larvae of the moth Parapoynx rugosalis (Lepidoptera: Pyralidae)

Abstract.

• 1 Aquatic larvae of the pyralid moth Parapoynx rugosalis Möschler repeatedly construct protective cases by cutting portions (discs) from leaves of the waterlily Nymphaea ampla and sandwich themselves between the disc and the underside of the host leaf. Construction of a new case requires leaving the old case, thus increasing exposure to predators and parasites.
• 2 In an experiment, larvae with protective cases experienced no mortality due to predation by fish, whereas larvae without cases experienced substantial predation.
• 3 In a series of choice tests, larvae preferentially selected young, tender leaves over old, tough leaves for construction of cases, and larvae spent significantly less time completing their shelters when cutting discs from young, tender leaves.
• 4 A partial explanation of why larvae select young, tender leaves for construction of their protective shelters may be that exposure time to predators during construction is minimized.
• 5 The same mechanism linking preferences for tender leaves to reduced exposure to predators during construction may also apply to other insect herbivores exhibiting leaf-rolling or case-building behaviour.

     

ThePassiflora problem in Hawaii; prospects and problems of controlling the forest weedP. Mollissima [Passifloraceae] with heliconiine butterflies

The introduced vine,Passiflora mollissima (Bailey), poses a threat to unique natural forest areas in Hawaii. While biological control is desirable, the proximity of the crop plant,P. edulis f.flavicarpa Degener requires control agents to be highly specific.Passiflora-specialist heliconiine butterflies may exhibit this necessary specificity. We examine the potential for heliconiines as pests and control agents ofPassiflora and report preliminary laboratory and field observations on host and habitat specificity.     

Leaf age and larval performance of the leaf beetle Paropsis atomaria

ABSTRACT.

• 1 Larval performance of the leaf beetle Paropsis atomaria Oliver was determined for larvae raised on both new and mature leaves of Eucalyptus blakelyi Maiden. Larvae were transferred to mature leaves at different ages; control larvae stayed on new leaves through all instars.
• 2 Only larvae reared on new leaves through the third instar survived to pupate on mature leaves; developmental time was prolonged by 20% and pupal weight was reduced by 50% in these larvae compared with larvae reared entirely on new leaves. Almost all larvae died when transferred to mature leaves as first, second or third instars.
• 3 Low survival and slow development on mature leaves was mainly due to failure by larvae to feed. Larvae palpated leaves and could discriminate among leaf ages immediately, without biting into the leaf tissue.
• 4 New leaves had higher concentrations of oil and tannins than old leaves, while there were no significant differences in nitrogen concentrations in the two types of leaves. Mature leaves were more than 3 times tougher than new leaves, in terms of g mm^?2 of penetrometer force.
• 5 In drought years E. blakelyi may not produce sufficient new leaves to supply specialist herbivores with their preferred food resource. We infer that drought years reduce P. atomaria larval performance significantly, and influence the population dynamics of the insect.

     

Leaf Colour as a Signal of Chemical Defence to Insect Herbivores in Wild Cabbage (Brassica oleracea)

Leaf colour has been proposed to signal levels of host defence to insect herbivores, but we lack data on herbivory, leaf colour and levels of defence for wild host populations necessary to test this hypothesis. Such a test requires measurements of leaf spectra as they would be sensed by herbivore visual systems, as well as simultaneous measurements of chemical defences and herbivore responses to leaf colour in natural host-herbivore populations. In a large-scale field survey of wild cabbage (Brassica oleracea) populations, we show that variation in leaf colour and brightness, measured according to herbivore spectral sensitivities, predicts both levels of chemical defences (glucosinolates) and abundance of specialist lepidopteran (Pieris rapae) and hemipteran (Brevicoryne brassicae) herbivores. In subsequent experiments, P. rapae larvae achieved faster growth and greater pupal mass when feeding on plants with bluer leaves, which contained lower levels of aliphatic glucosinolates. Glucosinolate-mediated effects on larval performance may thus contribute to the association between P. rapae herbivory and leaf colour observed in the field. However, preference tests found no evidence that adult butterflies selected host plants based on leaf coloration. In the field, B. brassicae abundance varied with leaf brightness but greenhouse experiments were unable to identify any effects of brightness on aphid preference or performance. Our findings suggest that although leaf colour reflects both levels of host defences and herbivore abundance in the field, the ability of herbivores to respond to colour signals may be limited, even in species where performance is correlated with leaf colour.     

Butterfly visual characteristics and ontogeny of responses to butterflies by a specialized tropical bird

The responses of two adult and three hand-reared, naive young rufous-tailed jacamars (Galbula ruficauda) to local butterflies were studied in feeding experiments. Four behavioural characteristics distinguish jacamars from other less specialized avian predators: (1) Exposed to butterflies for the first time, naive young jacamars would attack butterflies without showing signs of inhibition. Unacceptable butterflies, once captured, were taste-rejected quickly, and most survived the sampling. The few presumably unacceptable butterflies consumed by the birds were not observed to cause vomiting. (2) After gaining some familiarity with butterflies, young birds, like the adults, developed a reluctance to attack. They visually rejected certain classes of butterflies, often failing to attack them during an entire four-hour feeding trial. However, occasional attacks were made on butterflies in these ‘rejected’ classes. When this did occur, the insects proved to be actually easier to catch than those that were more often attacked. Once captured, however, the majority of these butterflies were taste-rejected. (3) For a given butterfly species, most individuals were either consumed or rejected. Thus, each species could be clearly classified as either acceptable or unacceptable to the jacamars. This consistency in jacamar responses resulted in a bimodal acceptability distribution of sympatric butterflies. (4) Young jacamars were capable of rapid associative learning and their responses were closely associated with butterfly visual characteristics in which colour pattern, flight behaviour, and morphology were also closely correlated. Thus, a single butterfly morphological parameter termed body shape (body length/thoracic diameter ratio) can adequately predict the feeding responses of jacamars. Visually detectable traits associated with butterflies possessing chemical defences may represent a balance between the need to signal unambiguously to specialized and/or experienced predators and the need to escape attacks by generalized and/or opportunistic predators. Since the proportion of specialized predators is higher in the tropical rainforest than in other habitat types, we expect greater divergence of morphological and behavioural characteristics between palatable and unpalatable butterflies in rainforest habitats.     

Pollen flow in Psiguria warscewiczii: a comparison of Heliconius butterflies and hummingbirds

Summary Pollen dispersal by male and female Heliconius butterflies and hermit hummingbirds was examined in a natural population of the tropical vine Psiguria warscewiczii. Fluorescent dyes were used to track flowers visited by individual pollinators. Mean distance of pollen dispersal was significantly higher by males than by female Heliconius. Butterflies dispersed pollen greater distances than did hummingbirds. There were no significant differences in dispersal distance among species of Heliconius. The number of flowers or plants to which dye was dispersed was greater for Heliconius than for hummingbirds.The home range behavior and site specificity exhibited by Heliconius, together with the preference by most Heliconius species in the study site for Psiguria led us to examine the relationship between the butterflies' home range and its daily foraging among Psiguria. A Heliconius mark-recapture program was conducted simultaneously with the dye program. This study showed that individual butterflies, for which extensive recaptures were made, visited an area of Psiguria plants on any one day that corresponded to the area in which those butterflies were caught or observed. The area over which a butterfly moved was termed its home range; within home ranges relative movement by butterflies was estimated as the mean distance between sequential captures. This estimate differed significantly among species of Heliconius; males of all species moving greater distances than females.