The consequences of larval aggregation in the butterfly Chlosyne lacinia

1. Females of Chlosyne lacinia (Geyer) (Lepidoptera: Nymphalidae, Melitaenae), the bordered patch butterfly, clump eggs in a few large clusters on their host plant, Helianthus annuus. Resulting larvae form sibling aggregations to at least the third instar.
2. The effect of group size on survival and development of C. lacinia larvae was tested experimentally in the field. Larvae developed faster and survived better in larger groups.
3. The effects of various predator guilds (ground-dwelling arthropods, aerial arthropods and avian predators) on survival of larvae was then tested while controlling group size. Ground-dwelling arthropods, mainly fire ants Solenopsis xyloni , reduced larval survival greatly but other solitary invertebrate and avian predators did not alter survival. Group defences and aposematism of C. lacinia larvae are probably ineffective against predatory ants that attack en masse and recruit other colony members.
4. In laboratory experiments, two possible mechanisms underlying faster development of larvae in larger groups were tested: (i) overcoming the physical toughness of host plant leaves, and (ii) social stimulus to feed. Results support the physical toughness hypothesis but not the social stimulus hypothesis.
5. Feeding in large groups by C. lacinia larvae confers multiple advantages, including protection from solitary predators and increased feeding efficiency because grouped, early-instar larvae can initiate feeding wounds on tough sunflower leaves. These advantages of larval gregariousness, coupled with reduced desiccation at the egg stage, apparently outweigh disadvantages of aggregation, such as interference and exploitative competition among larvae.     

Population biology of the checkerspot butterfly,Euphydryas chalcedona structure of the Jasper Ridge colony

Summary Euphydryas chalcedona at Jasper Ridge consists of two independently varying major demographic units, similar in scale but not in structure to those of Euphydryas editha. In contrast, however, Jasper Ridge E. chalcedona represent just a portion of a deme-while E. editha occurs as two partially connected demes. The life history and general ecology of E. chalcedona is described and compared with those of its intensively-studied congener. Various problems of mark-release-recapture estimation of population size are discussed     

The role of adult feeding in egg production and population dynamics of the checkerspot butterfly Euphydryas editha

Carbohydrate intake increases longevity, body weight maintenance and egg production in female Euphydryas editha. Amino acid intake leads to heavier eggs, larvae from which are more likely to survive. Females fed nectar produce more eggs in later masses than females which are not fed. During years of normal and below normal precipitation, larvae emerging from these later eggs are unlikely to reach obligatory size for diapause before their food dries up. On Jasper Ridge, where mortality is density-independent, nectar plays an important role increasing production of late egg masses during years of greater than normal rainfall when larvae from these masses are likely to reach diapause. The resulting large population increases, though infrequent, are probably important in maintaining population sizes large enough to reduce the chances of extinction during dry years.     

Mimicry in North American checkerspot butterflies: Euphydryas phaeton and Chlosyne harrisii (Nymphalidae)

ABSTRACT.

• 1 An examination of the phenetics, ecology and distribution of Chlosyne harrisii suggested that it was a potential mimic of the unpalatable Euphydryas phaeton.
• 2 The wing undersurfaces of the adults are very similar and the larvae are virtually identical. The geographic range of both butterflies is the northeastern United States and both are inhabitants of wet meadow habitats where they may fly together. The life histories of both species are closely synchronized.
• 3 A series of feeding experiments using blue-jays as predators showed first, that C.harrisii is palatable, and second, suggested that the birds could not distinguish between C.harrisii and E.phaeton on the basis of visual cues. Thus, Charrisii seems to be an effective Batesian mimic of E.phaeton.

     

The inheritance and population genetics of sex ratio in the butterfly Acraea encedon

In West and East Africa the butterfly, Acraea encedon , occurs in well-defined populations that are often predominantly female. Breeding the butterfly in the laboratory revealed the presence of an all-female strain, which is inherited directly through the female parent. It is probable that the inheritance is controlled by a Y-linked gene, causing meiotic drive in the Y chromosome, but the possibility of cytoplasmic inheritance has not been ruled out. A simple model for the population genetics of a predominantly female population indicates that such a population should rapidly become extinct due to the spreading of the all-female strain, but in most field populations studied extinction does not occur as quickly as predicted, if at all. Three factors which could enable populations to avoid extinction are investigated: suppressing systems, frequency-dependent mating preference and the sequence of emergence of the sexes in normal broods. No positive evidence has yet been found for the existence of a gene or genes capable of suppressing the sex ratio aberration, and no frequency-dependent mating preference was found, but an argument is presented which shows that the sequence of emergence in normal broods could be partly responsible for the maintenance of stable equilibria in predominantly female populations. Attempts to upset the sex ratio in the normal strain by making crosses between widely separated populations were not successful.     

Comparative population genetics of a mimicry locus among hybridizing Heliconius butterfly species

The comimetic Heliconius butterfly species pair, H. erato and H. melpomene, appear to use a conserved Mendelian switch locus to generate their matching red wing patterns. Here we investigate whether H. cydno and H. pachinus, species closely related to H. melpomene, use this same switch locus to generate their highly divergent red and brown color pattern elements. Using an F2 intercross between H. cydno and H. pachinus, we first map the genomic positions of two novel red/brown wing pattern elements; the G locus, which controls the presence of red vs brown at the base of the ventral wings, and the Br locus, which controls the presence vs absence of a brown oval pattern on the ventral hind wing. The results reveal that the G locus is tightly linked to markers in the genomic interval that controls red wing pattern elements of H. erato and H. melpomene. Br is on the same linkage group but approximately 26 cM away. Next, we analyze fine-scale patterns of genetic differentiation and linkage disequilibrium throughout the G locus candidate interval in H. cydno, H. pachinus and H. melpomene, and find evidence for elevated differentiation between H. cydno and H. pachinus, but no localized signature of association. Overall, these results indicate that the G locus maps to the same interval as the locus controlling red patterning in H. melpomene and H. erato. This, in turn, suggests that the genes controlling red pattern elements may be homologous across Heliconius, supporting the hypothesis that Heliconius butterflies use a limited suite of conserved genetic switch loci to generate both convergent and divergent wing patterns.     

Rapid assessment of butterfly diversity in a montane landscape

We present the results of a rapid assessment of butterfly diversity in the 754 ha Beaver Meadows study area in Rocky Mountain National Park, Larimer County, Colorado. We measured butterfly species richness and relative abundance as part of a landscape-scale investigation of diversity patterns involving several groups of organisms. A stratified random sampling design was used to include replication in both rare and common vegetation types. We recorded 49 butterfly species from the twenty-four 0.1 ha plots that were sampled four times during June, July, and August 1996. Butterfly species richness, diversity, and uniqueness were highest in quaking aspen (Populus tremuloides Michaux) groves and wet meadows, which occupy only a small proportion of the studied landscape. This result supports the suggestion that aspen areas represent hotspots of biological diversity in this montane landscape. Patterns of butterfly species richness were positively correlated with total vascular plant species richness (r = 0.69; P < 0.001), and native plant species richness (r = 0.64; P < 0.001). However, exotic plant species richness (r = 0.70; P < 0.001) and the cover of exotic plant species (r = 0.70; P < 0.001) were the best predictors of butterfly species richness.     

Extinction,reduction, stability and increase: The responses of checkerspot butterfly (Euphydryas) populations to the California drought

Summary The California drought of 1975–77 has been correlated with unusual size changes in populations of two species of Euphydryas butterflies. Several populations became extinct, some were dramatically reduced, others remained stable and at least one increased. These differences in population dynamic response are not concordant with predictions made earlier that populations with heavy density-dependent mortality should be more stable in the face of drought than unregulated populations. The different responses are related to the fine details of the relationships between the insects and their host plants, relationships which are variable between populations. Revised predictions are given in the light of better knowledge of the variability and complexity of these insect-host relationships. The diversity of responses underlines the dangers of generalizing about the ecology of a taxonomic species. The extinctions support the view that such events are frequent and significant in the biology of populations.     

Population differences in larval hostplant use in the checkerspot butterfly, Euphydryas chalcedona

Larvae from two populations of Euphydryas chalcedona Doubleday & Hewitson (Nymphalidae) were reared on their own hostplant and that of the other population, in both pre-diapause and post-diapause instars. One population, Chico, uses Penstemon breviflorus Lindl. (Scrophulariaceae), and the other, Echo Lake, uses P. newberryi Gray. Growth rate and survival were determined for pre-diapause and post-diapause larvae from both populations on both plant species; and digestive efficiencies were calculated during the prediapause instars. The results showed that larvae from the two populations differed in their responses to the two plant species. Pre-diapause larvae from Chico performed equally well on both plant species—survival and digestive indices were not significantly different for two Penstemon species. In contrast, pre-diapause larvae from Echo Lake performed significantly worse on the non-hostplant—growth and survival were significantly lower on the non-host, P. breviflorus. In addition, comparison of digestive efficiencies for the two plants showed that larvae from Echo Lake digested P. breviflorus better than P. newberryi, but were significantly less able to convert P. breviflorus to body mass. In the post-diapause instars, larvae from Chico grew faster on the host than on the non-host. Larvae from Echo Lake grew quite slowly on both plant species and significantly more of the Echo Lake larvae returned to diapause instead of completing development.

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Résumé Des chenilles de deux populations d'E. chalcedona ont été élevées sur leur propre plante-hôte et sur celle de l'autre population, aux stades avant et après diapause. Les deux populations s'alimentent sur différentes espèces de Penstemon (Scrophulariaceae), et une population—Echo Lake—est monophage sur P. newberry, tandis que l'autre—Chico—utilise d'abord P. breviflorus, mais les chenilles après diapause sont trouvées sur au moins deux autres espèces de plantes. Les taux de croissance et de survie ont été déterminés pour des chenilles avant et après diapause pour les deux populations sur les deux plantes; les efficacités digestives ont été calculées sur les chenilles avant diapause.Les résultats ont montré que les chenilles des deux populations différaient par leur degré de spécialisation digestive sur leur plante hôte normale: les chenilles de Chico ont utilisé aussi bien les deux plantes, tandis que celles d'Echo Lake le faisaient significativement moins bien sur la plante non-hôte, par suite de l'inaptitude à la digérer. Ainsi la population oligophage est alimentairement moins spécialisée et plus capable de se débrouiller avec une plante non-hôte. Après diapause, les chenilles de Chico s'alimentaient significativement mieux sur plante hôte que non-hôte, ce qui était le cas aussi pour la population monophage. Dans l'ensemble, les chenilles de la population monophage semblaient moins capables de se débrouiller dans des conditions défavorables ou moins avantageuses.

     

Natural,not urban,barriers define population structure for a coastal endemic butterfly

Habitat loss and fragmentation are the major causes of biodiversity loss, and, increasingly, habitat is fragmented by urbanization. Yet, the degree to which urbanization creates barriers to animal dispersal remains poorly understood. We used population genetic techniques to determine whether urbanization and/or natural landscape features are dispersal barriers to a butterfly, Atrytonopsis new species 1, throughout its range on coastal sand dunes that are increasingly threatened by development. Using AFLP markers that produced 89 polymorphic loci, we found significant population structure across the range of Atrytonopsis sp1. We found no indication that existing levels of urbanization were barriers to Atrytonopsis sp1 dispersal. Rather, two natural barriers, an ocean inlet and maritime forest, explained the genetic structure. Even in areas with long histories of urbanization, we found no significant isolation-by-distance relationship, and there was very low genetic differentiation between sampling locations. Consequently, conservation strategies for Atrytonopsis sp1, and potentially for other mobile insects that use open-structured habitats, should not focus explicitly on habitat corridors through urban areas, but rather should seek to preserve and restore as much habitat as possible across the butterfly’s range.     

The genetics and evo-devo of butterfly wing patterns

Understanding how the spectacular diversity of colour patterns on butterfly wings is shaped by natural selection, and how particular pattern elements are generated, has been the focus of both evolutionary and developmental biologists. The growing field of evolutionary developmental biology has now begun to provide a link between genetic variation and the phenotypes that are produced by developmental processes and that are sorted by natural selection. Butterfly wing patterns are set to become one of the few examples of morphological diversity to be studied successfully at many levels of biological organization, and thus to yield a more complete picture of adaptive morphological evolution.     

The evolutionary genetics of egg size plasticity in a butterfly

Abstract The evolution of phenotypic plasticity requires that it is adaptive, genetically determined, and exhibits sufficient genetic variation. For the tropical butterfly Bicyclus anynana there is evidence that temperature-mediated plasticity in egg size is an adaptation to predictable seasonal change. Here we set out to investigate heritability in egg size and genetic variation in the plastic response to temperature in this species, using a half-sib breeding design. Egg size of individual females was first measured at a high temperature 4 days after eclosion. Females were then transferred to a low temperature and egg size was measured after acclimation periods of 6 and 12 days respectively. Overall, additive genetic variance explained only 3-11% of the total phenotypic variance, whereas maternal effects were more pronounced. Genotype-environment interactions and cross-environmental correlations of less than unity suggest that there is potential for short-term evolutionary change. Our findings strengthen the support for the adaptive nature of temperature-mediated plasticity in egg size.     

The fragmented population structure of the Sardinian chalk hill blue butterfly (Lepidoptera, Lycaenidae)

The endemic Sardinian chalk hill blue butterfly, Polyommatus coridon gennargenti, is considered vulnerable to extinction because of its low genetic variation and restricted distribution. The species also has a fragmented distribution, which follows the patchy distribution pattern of its larval host-plant. A preliminary investigation of the population structure of P. coridon gennargenti was carried out on a small network of four local populations by means of capture–recapture methods. Estimated population sizes and movement rates among the four adjacent local populations suggest that this taxon has a metapopulation structure composed of loosely connected small local populations. Natural fragmentation, isolation, and traditional land use contribute to the vulnerability of P. coridon gennargenti to extinction. Low effective population sizes and restricted movement between habitat patches lead to inbreeding and an increased vulnerability to extinction of this island population.     

Ecological determinants of food plant choice in the checkerspot butterfly Euphydryas editha in Colorado

Summary The larvae of Euphydryas editha in Gunnison County, Colorado, feed on Castilleja linariifolia. Two related plants, C. chromosa and Penstemon strictus, occur in the same area and are equally nutritious, but are not eaten. Restriction to C. linariifolia appears to be a case of ecological monophagy — survival on the other two species is less likely not because of their biochemical make-up but because of their ecological characteristics, primarily their phenologies.     

Understanding the population dynamics of a rare, polyvoltine butterfly

Understanding the effects of endogenous and exogenous factors on population dynamics is essential for assessing the viability of populations, setting recovery goals for endangered species, and evaluating management options. Invertebrates are particularly difficult to monitor and few long-term datasets are available for these species. Additionally, limited resources make it necessary to perform monitoring as efficiently as possible. Here, I use the bivoltine Karner blue butterfly Lycaeides melissa samuelis to demonstrate how analyzing the effects of density-dependent factors and weather on separate life stages can be utilized to understand monitoring data, assess populations and identify critical life-history parameters. My first step was to compare the use of peak numbers as an index of population size with estimates obtained from a more data intensive methodology. Peak numbers proved to be an effective index, and so I utilized this index to analyze 10 and 13 years of monitoring data at two Karner blue butterfly sites in New York, USA. I modeled the effects of weather and density dependence on two distinct population growth rates ( λ ) per year. Analysis with Akaike's Information Criteria indicated that both sites were primarily influenced by density dependence during the summer period and by weather conditions during the winter period. Large population declines occurred in the winter period and were a result of the previous year's dry summer and cool spring weather. I conclude that recovery goals for this endangered species should include a second brood-carrying capacity, mean winter growth rate and multiple sites with independent populations. This study represents a rare, long-term study on the population dynamics of a polyvoltine species. Understanding the population dynamics of polyvoltine species, such as the Karner blue butterfly, will assist in the conservation of many invertebrate and small mammal species.     

The population ecology of a natural population of the pierid butterfly Colias alexandra

Summary Key factor analysis techniques were used to examine factors determining the abundance of a population of non-pest Colias. The number of individuals entering each successive life stage in the sample population are summarized in life tables for 1975 to 1979. Survivorship to the adult is a relatively consistent proportionality (-x=1.2%, S.D.=1.14; 1975–1979). Factors resulting in reduced natality and, less importantly, mortality during larval diapause determine the population trends for C. alexandra. Egg mortality, pre-diapause larval mortality and postdiapause mortality contribute little to these trends. Possible key sources contributing to reduced natality are examined. Mortality of adults (including removal by collectors), poor weather conditions during the oviposition period, unseasonal snow or drought which affect nectar sources or oviposition sites are among the factors which cause reduced natality and result in population depression.     

Dispersal, landscape occupancy and population structure in the butterfly Melanargia galathea

Dispersal is a key ecological process linking metapopulation dynamics in the landscape to distribution patterns at larger spatial scales. In this study, we investigated the landscape occupancy and genetic population structure of a butterfly species, Melanargia galathea. Several landscapes differing in composition and structure were sampled as well as populations at different spatial scales. We found that M. galathea occupied 91.3% of all habitat patches available within a particular landscape, probably due to a dominance of landscape scale processes such as rescue effect and recolonisation. A high level of genetic polymorphism within the sampled populations and a very low amount of genetic differentiation between populations was observed (G_st=0.034), characteristic of species with high dispersal capacity and/or high density. High dispersal rates ensured considerable gene mixing at the landscape scale while the influence of distance on dispersal success was detected at the regional and continental scales by a significant amount of isolation by distance. We also found that, at the landscape scale, the dispersal of this butterfly species was influenced by the spatial distribution of its habitat patches.

Zusammenfassung

Die Ausbreitung ist ein ökologischer Schlüsselprozess, der die Metapopulationsdynamik in einer Landschaft mit den Verbreitungsmustern auf einer größeren räumlichen Skala verbindet. In dieser Studie untersuchten wir die Verbreitungsmuster und die genetische Populationsstruktur einer Schmetterlingsart, Melanargia galathea. Es wurden sowohl einige Landschaften beprobt, die sich in der Zusammensetzung und der Struktur unterschieden, als auch Populationen auf verschiedenen räumlichen Skalen. Wir fanden, dass M. galathea 91.3% aller verfügbaren Habitatflecken in einer bestimmten Landschaft besetzte, möglicherweise aufgrund der Dominanz von Landschaftsskalenprozessen wie “Rescue-Effekt” und Rekolonisation. Es wurde ein hoher genetischer Polymorphismuslevel und ein sehr geringer Umfang genetischer Differenzierung innerhalb der beprobten Populationen beobachtet (G_st=0.034), was charakteristisch für Arten mit einer großen Ausbreitungsfähigkeit und/oder hoher Dichte ist. Hohe Ausbreitungsraten sicherten eine beträchtliche Genvermischung auf der Landschaftsskala, während der Einfluss der Entfernung auf den Verbreitungserfolg auf der regionalen und kontinentalen Skala durch einen signifikanten Anteil der Isolation aufgrund von Entfernung festgestellt wurde. Wir fanden auch, dass auf der Landschaftsskala die Ausbreitung dieses Schmetterlings durch die räumliche Verteilung der Habitatflecken beeinflusst wurde.     

Thermal influences on oviposition in the montane butterfly Euphydryas gillettii

Summary Euphydryas gillettii in a montane meadow in Wyoming oviposits in clusters on the highest large leaves of its larval host, the shrub Lonicera involucrata. Egg masses are found on the underside of leaves which predominantly face the southeast and thereby intercept morning sun. Measurements of leaf temperatures show that, during the cool, clear mornings, these leaves are warmer than leaves of the opposite orientation. Manipulative experiments demonstrate that eggs on these slightly warmer microsites hatch significantly faster than eggs on leaves of other orientations. Because of the rapid onset of winter, an increased developmental rate is likely to correlate directly with increased fitness. The proximal mechanism yielding such a result may be that females choose oviposition sites in the shade.