Patterns and processes of nestedness in a Great Basin butterfly community

We examined nestedness and potential mechanisms causing that distributional pattern in resident butterfly communities of the Toiyabe Range, a mountain range in the central Great Basin of western North America. We tested whether life history characteristics, including habitat use and vagility, affected the relative degree of nestedness or mean species incidence. We also tested whether nestedness at the level of individual species was independent of life history. Relationships between distributional patterns and habitat use, particularly in ecologically sensitive riparian areas, are relevant to ongoing conservation planning in the Great Basin. The distributional pattern of the 68 resident butterfly species in 19 Toiyabe Range canyons was significantly nested, as was the distribution of all functional subgroups that we tested. Life history affected neither relative nestedness of species groups nor mean species incidence. More than 80% of the individual butterfly species that inhabit the Toiyabe Range had distributions that were more nested than expected. Colonization does not appear to have played an important role in determining the composition of butterfly communities in Toiyabe Range canyons. Likewise, selective dispersal has probably played a minor role in producing nested distributions of Toiyabe Range butterflies. Our results suggest either that impacts to riparian areas are not jeopardizing species viability, or that highly sensitive butterfly species have already been extirpated from the Toiyabe Range. Received: 15 February 1998 / Accepted: 19 December 1998     

The genetic consequence of differing ecological demands of a generalist and a specialist butterfly species

Many studies aim at testing the impact of recent fragmentation on the genetic diversity and connectivity of populations, while some species do exist naturally in fragmented landscapes because of their habitat requirements. Therefore, it is important to look at the genetic signatures of species occurring in naturally fragmented landscapes in order to disentangle the effect of fragmentation from the effect of habitat requirements. We selected two Nymphalid butterflies for this purpose. While Melanargia galathea is a common butterfly in flower-rich meadows, Melitaea aurelia is closely connected to calcareous grasslands, thus being historically fragmented due to its ecological demands. For the analysis of the genetic response on these opposed patterns, we analysed 18 allozyme loci for 789 individuals (399 individuals of M. galathea and 390 individuals of M. aurelia) in a western German study region with adjacent areas in Luxemburg and northeastern France. Both species showed similarly low genetic differentiations among local populations (M. galathea: F _ST 3.3%; M. aurelia: F _ST 3.6%), both combined with a moderate level of inbreeding. Isolation-by-distance analysis revealed a significant correlation for both species with similar amounts of explained variances (M. galathea: r ² = 27.8%; M. aurelia: r ² = 28.5%). Most parameters of genetic diversity were higher in M. galathea than in M. aurelia, but the latter species had a considerably higher amount of rare or locally restricted genes; the differing ecological demands are thus reflected in these differences. Both species thus seem to be genetically well suited to their respective ecological requirements. In the light of conservation genetics, we deduce that highly fragmented populations are not necessarily prone to extinction. The extinction risk might be linked to the life history of an organism and its population genetic structure.     

Extending conventional priors for testing general hypotheses in linear models

We consider that observations come from a general normal linearmodel and that it is desirable to test a simplifying null hypothesisabout the parameters. We approach this problem from an objectiveBayesian, model-selection perspective. Crucial ingredients forthis approach are ‘proper objective priors’ to beused for deriving the Bayes factors. Jeffreys-Zellner-Siow priorshave good properties for testing null hypotheses defined byspecific values of the parameters in full-rank linear models.We extend these priors to deal with general hypotheses in generallinear models, not necessarily of full rank. The resulting priors,which we call ‘conventional priors’, are expressedas a generalization of recently introduced ‘partiallyinformative distributions’. The corresponding Bayes factorsare fully automatic, easily computed and very reasonable. Themethodology is illustrated for the change-point problem andthe equality of treatments effects problem. We compare the conventionalpriors derived for these problems with other objective Bayesianproposals like the intrinsic priors. It is concluded that bothpriors behave similarly although interesting subtle differencesarise. We adapt the conventional priors to deal with nonnestedmodel selection as well as multiple-model comparison. Finally,we briefly address a generalization of conventional priors tononnormal scenarios.     

Key traits in a threatened butterfly and its common sibling: implications for conservation

We ask here which traits predispose one species to extreme rarity and possible extinction while a sympatric sibling is geographically widespread. With background knowledge on the level of habitat specialization of the two species, the population structure and movement of the localized and threatened Orachrysops ariadne were compared to those of the common and highly sympatric O. subravus, using mark-release-recapture. Of a total of 290 marked O. ariadne individuals 42.8% were recaptured, while of 631 O. subravus individuals 49.3% were recaptured. The Jolly-Seber model was used to estimate daily population numbers (N _i ), survival rates (ϕ _i ), recruitment rates (B _i ), proportion of marked animals in the total population (∝_i), and the number of marked animals at risk (M _i ). O. ariadne is a remarkably rare animal, averaging only 10 individuals ha⁻¹ within its small, remaining colonies. Average residence times of male adults were generally similar in both species, being just over 5 days. O. ariadne is a strong and rapid back and forth flier, covering mean recapture distances of 157 m, almost twice that of O. subravus, principally in search of scarce nectar sources. In short, the rarity of O. ariadne is not so much to do with behaviour, survivorship or longevity, but rather with limited availability of the specialized habitat patches for both larvae and adults, and, in particular, the extreme scarcity of the host plant. Evidence suggests that there has been very high selection pressure on the key trait of strong flight as a compensation for going down the apparently highly risky path of extreme microhabitat specialization. Of concern for conservation of this rare species is that these rare habitat patches have become increasingly isolated through transformation of the surrounding landscape. Reduction of the barrier effects of agroforestry through creation of linkages between colonies is recommended, especially as O. ariadne is such a strong flier. Such corridors are indeed now being implemented.     

Maternal body size as an evolutionary constraint on egg size in a butterfly

Genetic and developmental constraints have often been invoked to explain patterns of existing morphologies. Yet, empirical tests addressing this issue directly are still scarce. We here set out to investigate the importance of maternal body size as an evolutionary constraint on egg size in the tropical butterfly Bicyclus anynana, employing an artificial two-trait selection experiment on simultaneous changes in body and egg size (synergistic and antagonistic selection). Selection on maternal body size and egg size was successful in both the synergistic and the antagonistic selection direction. Yet, responses to selection and realized heritabilities varied across selection regimes: the most extreme values for pupal mass were found in the synergistic selection directions, whereas in the antagonistic selection direction realized heritabilities were low and nonsignificant in three of four cases. In contrast, for egg size the highest values were obtained in the lines selected for low pupal mass. Thus, selection on body size yielded a stronger correlated response in egg size than vice versa, which is likely to bias (i.e., constrain), if weakly, evolutionary change in body size. However, it seems questionable whether this will prevent evolution toward novel phenotypes, given enough time and that natural selection is strong. Correlated responses to selection were overall weak. Egg and larval development times tended to be associated with changes in maternal size, whereas variation in pupal development times weakly tended to follow variation in egg size. Lifetime fecundity was similar across selection regimes, except for females simultaneously selected for large body mass and small egg size, exhibiting increased fecundity. Multiple regressions showed that lifetime fecundity and concomitantly reproductive investment were primarily determined by longevity, as expected for an income breeder, whereas egg size was primarily determined by pupal mass. Evidence for a phenotypic trade-off between egg size and number was weak.     

Postglacial recolonization at a snail's pace (Trochulus villosus): confronting competing refugia hypotheses using model selection

The localization of Last Glacial Maximum (LGM) refugia is crucial information to understand a species' history and predict its reaction to future climate changes. However, many phylogeographical studies often lack sampling designs intensive enough to precisely localize these refugia. The hairy land snail Trochulus villosus has a small range centred on Switzerland, which could be intensively covered by sampling 455 individuals from 52 populations. Based on mitochondrial DNA sequences (COI and 16S), we identified two divergent lineages with distinct geographical distributions. Bayesian skyline plots suggested that both lineages expanded at the end of the LGM. To find where the origin populations were located, we applied the principles of ancestral character reconstruction and identified a candidate refugium for each mtDNA lineage: the French Jura and Central Switzerland, both ice-free during the LGM. Additional refugia, however, could not be excluded, as suggested by the microsatellite analysis of a population subset. Modelling the LGM niche of T. villosus , we showed that suitable climatic conditions were expected in the inferred refugia, but potentially also in the nunataks of the alpine ice shield. In a model selection approach, we compared several alternative recolonization scenarios by estimating the Akaike information criterion for their respective maximum-likelihood migration rates. The 'two refugia' scenario received by far the best support given the distribution of genetic diversity in T. villosus populations. Provided that fine-scale sampling designs and various analytical approaches are combined, it is possible to refine our necessary understanding of species responses to environmental changes.     

Seed dispersal patterns in a temperate forest during a mast event: performance of alternative dispersal kernels

Seed dispersal patterns were studied in a north-western Spain temperate forest community to assess the performances of alternative dispersal kernels during two years with ecologically contrasting scenarios; a non-mast year, and a mast year of the dominant canopy species, beech Fagus sylvatica. Dispersal kernels were fitted under a Bayesian modeling framework. Both simple and mixture kernels were considered for the five more abundant tree species (Corylus avellana, Crataegus monogyna, F. sylvatica, Ilex aquifolium and Taxus baccata). Mixture kernels provided a better fit for almost all species, and the log-normal performed best for T. baccata. No relationship between dispersal syndromes and the best dispersal kernel function emerged. However, we found temporal changes in the shape of the dispersal kernels that seemed to be related to variation in relative fruit production among species and the resulting changes in the responses of dispersal vectors. This reveals a potential role for disperser-mediated indirect effects in terms of introducing temporal variation in species spread. In this sense, our results highlight the need to consider single species seed dispersal as a community process. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Monandry and polyandry as alternative lifestyles in a butterfly

Butterflies show considerable variability in female mating frequency, ranging from monandrous species to females mating several timesin their lifetime. Degree of polyandry also varies within species,with some females only mating once and others mating multiply.Previous studies have shown that one reason for female multiplemating is to obtain nutritious male donations that both increasethe longevity of females and result in higher lifetime fecundity.Despite the presence of male nutrient donations, some femalesof the green-veined white butterfly (Pieridae: Pieris napi)never mate more than once. In this study, we examined thisapparent paradox. We assessed to what degree polyandry is undergenetic control by a full-sib analysis, and we also estimatedthe broad sense heritability of female lifetime fecundity in singly mated females. Both polyandry and lifetime fecundityhave a genetic component. However, degree of polyandry appearsto be traded off against reduced longevity when denied theopportunity to mate more than once. It is possible that femaleP. napi display different reproductive strategies, with somefemales relying on male donations to realize their potentialfecundity and others relying on their own resources for egg production. In nature, polyandrous females may be preventedfrom mating multiply due to unfavorable weather. We discussthe possibility that the trade-off between degree of polyandryand life span when singly mated may affect the maintenanceof genetic variability in female mating frequency in this species.Possible reasons for these different reproductive strategiesare discussed.     

The fitness consequences of kin-biased dispersal in a cooperatively breeding bird

Cooperative alliances among kin may not only lead to indirect fitness benefits for group-living species, but can also provide direct benefits through access to mates or higher social rank. However, the immigrant sex in most species loses any potential benefits of living with kin unless immigrants disperse together or recruit relatives into the group in subsequent years. To look for evidence of small subgroups of related immigrants within social groups (kin substructure), we used microsatellites to assess relatedness between immigrant females of the cooperatively breeding superb starling, Lamprotornis superbus. We determined how timing of immigration led to kin subgroup formation and if being part of one influenced female fitness. Although mean relatedness in groups was higher for males than females, 26% of immigrant females were part of a kin subgroup with a sister. These immigrant sibships formed through kin recruitment across years more often than through coalitions immigrating together in the same year. Furthermore, females were more likely to breed when part of a kin subgroup than when alone, suggesting that female siblings form alliances that may positively influence their fitness. Ultimately, kin substructure should be considered when determining the role of relatedness in the evolution of animal societies.     

Contest outcome in a territorial butterfly: the role of motivation

In many butterfly species, males compete over areas advantageous for encountering females. Rules for contest settlement are, however, largely unknown and neither morphological nor physiological traits can reliably predict the contest outcome. Here, we test the hypothesis that contests are settled in accordance with a motivation asymmetry. We staged contests between males of Pararge aegeria and after removing the resident, the non-resident was allowed (i) either to interact with a non-receptive female for 30 min (n = 30) or (ii) to spend 30 min alone in the cage (n = 30), after which the initial resident was reintroduced. The results show that males that had interacted with a female had a higher probability of becoming dominant and reversing contest outcome. Moreover, males that were faster to take over a vacant territory when the resident was removed were more likely to become dominant. Here, we show for the first time, to our knowledge, that frequent encounters with a mated female can increase male motivation to persist in a territorial contest in a butterfly. Further, we suggest that variation in intrinsic motivation reflects male eagerness to take over vacant territory. This study indicates that variation in resource value and motivational asymmetries are important for settling contests in butterflies.     

Generation-dependent female choice: behavioral polyphenism in a bivoltine butterfly

Climatic and biotic circumstances vary as seasons shift, anddifferent cohorts of multivoltine species are likely subjectedto different selection regimes. The bivoltine butterfly Leptideareali (Réal's wood white; Lepidoptera: Pieridae) appearsduring May and June in central Sweden and has a partial secondgeneration in late July. We manipulated both generations toappear simultaneously and performed laboratory mating experimentsthat showed the presence of a behavioral polyphenism in matingpropensity, which is induced during the developmental stages.Females of the summer generation expressed higher mating propensitiesthan spring generation females. Spring females showed an increasein mating propensity with increasing age, whereas summer femalesaccepted most matings already when they were only 1 or 2 daysold. It is likely that larval time constraints, a lower abundanceof males and a lower risk of accepting a male of their univoltinesister species Leptidea sinapis (wood white), have relaxed selectionon mate discrimination among summer generation females. A majorchallenge for future research is to further investigate thedevelopmental pathways causing the polyphenism and the adaptiveimplications of cohort-dependent behaviors.     

Condition-dependent dispersal of a patchily distributed riparian ground beetle in response to disturbance

In common with many habitat elements of riverine landscapes, exposed riverine sediments (ERS) are highly disturbed, naturally patchy and regularly distributed, whose specialists are strongly adapted to flood disturbance and loss of habitat due to succession. Investigations of dispersal in ERS habitats therefore provide an important contrast to the unnaturally fragmented, stable systems usually studied. The present investigation analysed the three interdependent stages of dispersal: (1) emigration, (2) inter-patch movement and (3) immigration of a common ERS specialised beetle, Bembidion atrocaeruleum (Stephens 1828) (Coleoptera, Carabidae), in a relatively unmodified section of river, using mark–resight methods. Dispersal was correlated with estimates of local population size and density, water level and patch quality in order to test for condition-dependent dispersal cues. Flood inundation of habitat was found to increase strongly the overall rate of dispersal, and the rate of emigration was significantly higher from patches that were heavily trampled by cattle. Strongly declining numbers of dispersers with distance suggested low dispersal rates during periods of low water level. Dispersal in response to habitat degradation by cattle trampling would likely lead to a higher overall population fitness than a random dispersal strategy. Dispersal distances were probably adapted to the underlying habitat landscape distribution, high-flow dispersal cues and ready means of long-distance dispersal through hydrochory. Species whose dispersal is adapted to the natural habitat distribution of riverine landscapes are likely to be strongly negatively affected by reduced flood frequency and intensity and habitat fragmentation through flow regulation or channelisation.Electronic Supplementary Material Supplementary material is available to authorised users in the online version of this article at .     

A generalized diffusion model for growth and dispersal in a population

A reaction-diffusion model is presented in which spatial structure is maintained by means of a diffusive mechanism more general than classical Fickian diffusion. This generalized diffusion takes into account the diffusive gradient (or gradient energy) necessary to maintain a pattern even in a single diffusing species. The approach is based on a Landau-Ginzburg free energy model. A problem involving simple logistic kinetics is fully analyzed, and a nonlinear stability analysis based on a multi-scale perturbation method shows bifurcation to non-uniform states.Part of this work was done while at the Mathematical Institute, Oxford University as a Senior Visiting Fellow supported by the Science Research Council of Great Britain under grant GR/B31378     

Maintenance of high pollen dispersal in Eucalyptus wandoo, a dominant tree of the fragmented agricultural region in Western Australia

Pollen dispersal was investigated in five remnant populations of Eucalyptus wandoo, a dominant insect-pollinated tree in the fragmented agricultural region of southern Western Australia. Paternity analysis using six microsatellite loci identified a pollen source for 45% of seedlings, and the remainder were assumed to have arisen from pollen sources outside the stands. Outcrossing was variable, ranging from 52 to 89%, and long distance pollen dispersal was observed in all populations with up to 65% of pollen sourced from outside the populations over distances of at least 1 km. Modelling dispersal functions for pollination events within the two larger populations showed little difference between the four two-parameter models tested and indicated a fat-tailed dispersal curve. Similarity of direct and indirect historical estimates of gene flow indicates maintenance of gene flow at levels experienced prior to fragmentation. The study revealed extensive long distance pollen dispersal in remnant patches of trees within a fragmented agricultural landscape in the southern temperate region and highlighted the role of remnant patches in maintaining genetic connectivity at the landscape scale.     

Decoupling of reproductive rates and parental expenditure in a polyandrous butterfly

Current theory postulates that the operational sex ratio (OSR)determines the relative degree of mating competition in thetwo sexes and is in turn influenced by a sexual difference inthe potential reproductive rate (PRR) denned as 1/time out,where time out is the time an individual must spend recoveringfrom a bout of mating activity and/or caring for offspring.In bushcricket mating systems where males provide females witha nuptial gift, relative energy expenditure in offspring influencesthe PRR of males and females and underlies a diet-mediated shiftin the OSR. Here we investigated if there is a similar positiverelationship between relative parental nutrient expenditurein offspring and PRR in the polyandrous butterfly Pieris napi,where female fecundity is strongly dependent on male nuptialgifts at mating. By varying the amount of nutrients femalesreceive at mating and relating this to number of offspring produced,we show that male P. napi have, on average, a nutrient expenditurein offspring equaling that of females. In spite of this, themale reproductive rate is 8–13 times higher than thatof females. Hence the relative degree of parental expenditurein offspring is largely decoupled from the degree of matingcompetition in P. napi. Two alternative explanations are advancedto account for the difference between the butterfly and thebushcricket mating systems.     

Genetic diversity and dispersal of Phragmites australis in a small river system

Even though the reed, Phragmites australis, is an extensively studied wetland species, little is known about reproduction and dispersal modes within and among reed populations at the scale of small river systems. Using microsatellite analysis of 189 individuals from three adjacent river catchments in the Czech Republic, we elucidated the role of the river corridors in the dispersal of P. australis. Using Bayesian clustering of individuals, we found that 19% of clusters were distributed only along one river, which implied dispersal by water (or by wind) along river corridors, whereas 38% of clusters were widely distributed and were likely the product of wind long-distance dispersal among rivers. Intensive exchange of propagules among river systems is further demonstrated by only 6% of total variance being attributed to the variance among rivers in the AMOVA-analysis. Spatial autocorrelation analysis revealed a decreasing pattern up to 5–10 km and no clear pattern over longer distances. This gives an evidence for pollen and seed dispersal at short distances (up to 1 km), whereas most likely only seed dispersal at longer distances up to 10 km. We found five multilocus genotypes distributed in two different populations. The distances between populations with the same genotype ranged from 0.5 to 10.8 km. This can be interpreted as long-distance vegetative dispersal.     

Butterfly edge effects are predicted by a simple model in a complex landscape

Edge responses have been studied for decades and form a critical component of our understanding of how organisms respond to landscape structure and habitat fragmentation. Until recently, however, the lack of a general, conceptual framework has made it difficult to make sense of the patterns and variability reported in the edge literature. We present a test of an edge effects model which predicts that organisms should avoid edges with less-preferred habitat, show increased abundance near edges with preferred habitat or habitat containing complementary resources, and show no response to edges with similar-quality habitat that offers only supplementary resources. We tested the predictions of this model against observations of the edge responses of 15 butterfly species at 12 different edge types within a complex, desert riparian landscape. Observations matched model predictions more than would be expected by chance for the 211 species/edge combinations tested over 3 years of study. In cases where positive or negative edge responses were predicted, observed responses matched those predictions 70% of the time. While the model tends to underpredict neutral results, it was rare that an observed edge response contradicted that predicted by the model. This study also supported the two primary ecological mechanisms underlying the model, although not equally. We detected a positive relationship between habitat preferences and the slope of the observed edge response, suggesting that this basic life history trait underlies edge effects and influences their magnitude. Empirical evidence also suggested the presence of complementary resources underlies positive edge responses, but only when completely confined to the adjacent habitat. This multi-species test of a general edge effects model at multiple edge types shows that resource-based mechanisms can explain many edge responses and that a modest knowledge of life history attributes and resource availability is sufficient for predicting and understanding many edge responses in complex landscapes. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.