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11.
  • Bergström, Anders, 1969- (författare)
  • Oviposition strategies in butterflies and consequences for conservation
  • 2005
  • Doktorsavhandling (övrigt vetenskapligt/konstnärligt)abstract
    • Oviposition strategies are an important component of life history evolution in insects. Oviposition is a complex behaviour, where females must locate suitable habitat patches, locate oviposition sites, evaluate the quality, and finally decide how many eggs to lay. In this thesis, I explore some factors that may influence oviposition strategies in female butterflies. It discusses oviposition strategies from large to fine scales and ends with a discussion on implications for conservation. In paper I we tested for local adaptations in oviposition preference and larval performance in Polyommatus icarus. The result suggests that this species is evolutionary conservative in its utilization of the tested host plants. In the second paper we investigated frequency-dependent host plant choice in Polygonia c-album and showed that females spent significantly more time, and laid more eggs, in patches with high frequency of the preferred host. The influence of nectar sources on host plant choice in P. icarus was tested in paper III, females preferred ovipositing on host plants with flowers and after feeding. In paper IV we investigated oviposition “mistakes” in P. c-album and found differences in discrimination of hosts and non-hosts. Further, we showed that P. c-album butterflies are capable of responding to host quality by increasing clutch sizes on a high ranked host (paper V). Finally, in paper VI I explored how female oviposition behaviour can influence habitat management for a threatened butterfly species, Parnassius mnemosyne, by doing an observational field study on oviposition site preferences. This thesis highlights oviposition as a stepwise process and show that female discrimination is important at a multitude of levels with implications for oviposition strategies.
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12.
  • Bergström, Anders, et al. (författare)
  • Putting more eggs in the best basket : clutch size regulation in the comma butterfly
  • 2006
  • Ingår i: Ecological Entomology. - : Wiley. - 0307-6946 .- 1365-2311. ; 31:3, s. 255-260
  • Tidskriftsartikel (refereegranskat)abstract
    • 1. Many studies have identified different factors influencing clutchsize regulation, primarily within various groups of insects. One prediction is that ovipositing females should increase clutch size with host quality. However, in many studies it is not clear whether ovipositing females are responding to host quality or quantity. 2. Females of the polyphagous comma butterfly, allowed to oviposit on two hosts differing greatly in quality: the preferred host, stinging nettle (Polygonia c-album (L.), wereUrtica dioica L.), and the low-ranked host, birch (Betula pubescens  Ehrh). Ovipositing females were observed visually and clutch sizes were recorded. The experiment was repeated in three different years; in total, 938 observations of oviposition events were made. 3. In all three years, females ovipositing on (median 1.6–1.85) compared with females ovipositing on 1.0–1.3) three years were pooled. 4. Thus, on better hosts. It is suggested that the proximate mechanism is likely to be a response to the same stimuli used for female ranking of host plants in the preference hierarchy. U. dioica laid larger clutchesB. pubescens (median. The difference was significant in two out of three years and when allP. c-album females exhibit clutch-size regulation, with larger clutches
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13.
  • Blanckenhorn, Wolf U, et al. (författare)
  • Proximate causes of Rensch's rule : Does sexual size dimorphism in arthropods result from sex differences in development time?
  • 2007
  • Ingår i: American Naturalist. ; 169:2, s. 245-257
  • Tidskriftsartikel (refereegranskat)abstract
    • A prominent interspecific pattern of sexual size dimorphism (SSD) is Rensch's rule, according to which male body size is more variable or evolutionarily divergent than female body size. Assuming equal growth rates of males and females, SSD would be entirely mediated, and Rensch's rule proximately caused, by sexual differences in development times, or sexual bimaturism (SBM), with the larger sex developing for a proportionately longer time. Only a subset of the seven arthropod groups investigated in this study exhibits Rensch's rule. Furthermore, we found only a weak positive relationship between SSD and SBM overall, suggesting that growth rate differences between the sexes are more important than development time differences in proximately mediating SSD in a wide but by no means comprehensive range of arthropod taxa. Except when protandry is of selective advantage ( as in many butterflies, Hymenoptera, and spiders), male development time was equal to ( in water striders and beetles) or even longer than ( in drosophilid and sepsid flies) that of females. Because all taxa show female-biased SSD, this implies faster growth of females in general, a pattern markedly different from that of primates and birds (analyzed here for comparison). We discuss three potential explanations for this pattern based on life-history trade-offs and sexual selection.
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14.
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15.
  • Braga, Mariana P., et al. (författare)
  • Bayesian Inference of Ancestral Host-Parasite Interactions under a Phylogenetic Model of Host Repertoire Evolution
  • 2020
  • Ingår i: Systematic Biology. - : Oxford University Press (OUP). - 1063-5157 .- 1076-836X. ; 69:6, s. 1149-1162
  • Tidskriftsartikel (refereegranskat)abstract
    • Intimate ecological interactions, such as those between parasites and their hosts, may persist over long time spans, coupling the evolutionary histories of the lineages involved. Most methods that reconstruct the coevolutionary history of such interactions make the simplifying assumption that parasites have a single host. Many methods also focus on congruence between host and parasite phylogenies, using cospeciation as the null model. However, there is an increasing body of evidence suggesting that the host ranges of parasites are more complex: that host ranges often include more than one host and evolve via gains and losses of hosts rather than through cospeciation alone. Here, we develop a Bayesian approach for inferring coevolutionary history based on a model accommodating these complexities. Specifically, a parasite is assumed to have a host repertoire, which includes both potential hosts and one or more actual hosts. Over time, potential hosts can be added or lost, and potential hosts can develop into actual hosts or vice versa. Thus, host colonization is modeled as a two-step process that may potentially be influenced by host relatedness. We first explore the statistical behavior of our model by simulating evolution of host-parasite interactions under a range of parameter values. We then use our approach, implemented in the program RevBayes, to infer the coevolutionary history between 34 Nymphalini butterfly species and 25 angiosperm families. Our analysis suggests that host relatedness among angiosperm families influences how easily Nymphalini lineages gain new hosts.
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16.
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17.
  • Braga, Mariana Pires, 1988- (författare)
  • Evolution of host repertoires and the diversification of butterflies
  • 2019
  • Doktorsavhandling (övrigt vetenskapligt/konstnärligt)abstract
    • All herbivorous insects are specialized to some extent to their host plants, but the level of specialization varies greatly. Insect-plant coevolution is often invoked to explain the large diversity of herbivorous insects, but the role of specialization during diversification is still controversial. Although well-studied, our understanding of the evolution of species interactions is still improving, and recent theoretical developments have highlighted the role of generalization (via colonization of new hosts) on diversification. In this thesis, various approaches are combined for a detailed study of the origins of macroevolutionary patterns of host use and butterfly diversity. Chapter I provides a mechanistic basis for such patterns through simulations of lineages evolved in silico. By separating the effects of the number of hosts used by a parasite lineage and the diversity of resources they encompass, we found that resource diversity, rather than host range per se, was the main driver of parasite species richness in both simulated and empirical systems. In Chapter II, we combined network and phylogenetic analyses to quantify support for the two main hypothesized drivers of diversification of herbivorous insects. Based on analyses of two butterfly families, Nymphalidae and Pieridae, we found that variability in host use is essential for diversification, while radiation following the colonization of a new host is rare but can produce high diversity. We then reconciled the two alternative hypotheses into a unified process of host-associated diversification where continuous probing of new hosts and retention of the ability to use hosts colonized in the past are the main factors shaping butterfly-plant networks. While network analysis is a powerful tool for investigating patterns of interaction, other methods are necessary to directly test the mechanisms generating the observed patterns. Therefore, in Chapter III we describe a model of host repertoire evolution we developed for Bayesian inference of evolution of host-parasite interactions. The approach was validated with both simulated and empirical data sets. Finally, in Chapter IV we used the method described in Chapter III to explicitly test the predictions made in Chapter II about the evolution of butterfly-plant networks. We found direct evidence for the role of expansion of fundamental host repertoire and phylogenetic conservatism as important drivers of host repertoire evolution. Thus, using three different approaches, we found overall support for the idea that variation in host use accumulated over evolutionary time is essential for butterfly diversification.
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18.
  • Braga, Mariana P., et al. (författare)
  • Host use dynamics in a heterogeneous fitness landscape generates oscillations in host range and diversification
  • 2018
  • Ingår i: Evolution. - : Wiley. - 0014-3820 .- 1558-5646. ; 72:9, s. 1773-1783
  • Tidskriftsartikel (refereegranskat)abstract
    • Colonization of novel hosts is thought to play an important role in parasite diversification, yet little consensus has been achieved about the macroevolutionary consequences of changes in host use. Here, we offer a mechanistic basis for the origins of parasite diversity by simulating lineages evolved in silico. We describe an individual-based model in which (i) parasites undergo sexual reproduction limited by genetic proximity, (ii) hosts are uniformly distributed along a one-dimensional resource gradient, and (iii) host use is determined by the interaction between the phenotype of the parasite and a heterogeneous fitness landscape. We found two main effects of host use on the evolution of a parasite lineage. First, the colonization of a novel host allowed parasites to explore new areas of the resource space, increasing phenotypic and genotypic variation. Second, hosts produced heterogeneity in the parasite fitness landscape, which led to reproductive isolation and therefore, speciation. As a validation of the model, we analyzed empirical data from Nymphalidae butterflies and their host plants. We then assessed the number of hosts used by parasite lineages and the diversity of resources they encompass. In both simulated and empirical systems, host diversity emerged as the main predictor of parasite species richness.
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19.
  • Braga, Mariana P., et al. (författare)
  • Phylogenetic reconstruction of ancestral ecological networks through time for pierid butterflies and their host plants
  • 2021
  • Ingår i: Ecology Letters. - : Wiley. - 1461-023X .- 1461-0248. ; 24:10, s. 2134-2145
  • Tidskriftsartikel (refereegranskat)abstract
    • The study of herbivorous insects underpins much of the theory that concerns the evolution of species interactions. In particular, Pieridae butterflies and their host plants have served as a model system for studying evolutionary arms races. To learn more about the coevolution of these two clades, we reconstructed ancestral ecological networks using stochastic mappings that were generated by a phylogenetic model of host-repertoire evolution. We then measured if, when, and how two ecologically important structural features of the ancestral networks (modularity and nestedness) evolved over time. Our study shows that as pierids gained new hosts and formed new modules, a subset of them retained or recolonised the ancestral host(s), preserving connectivity to the original modules. Together, host-range expansions and recolonisations promoted a phase transition in network structure. Our results demonstrate the power of combining network analysis with Bayesian inference of host-repertoire evolution to understand changes in complex species interactions over time.
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20.
  • Braga, Mariana P., et al. (författare)
  • Unifying host-associated diversification processes using butterfly-plant networks
  • 2018
  • Ingår i: Nature Communications. - : Springer Science and Business Media LLC. - 2041-1723. ; 9
  • Tidskriftsartikel (refereegranskat)abstract
    • Explaining the exceptional diversity of herbivorous insects is an old problem in evolutionary ecology. Here we focus on the two prominent hypothesised drivers of their diversification, radiations after major host switch or variability in host use due to continuous probing of new hosts. Unfortunately, current methods cannot distinguish between these hypotheses, causing controversy in the literature. Here we present an approach combining network and phylogenetic analyses, which directly quantifies support for these opposing hypotheses. After demonstrating that each hypothesis produces divergent network structures, we then investigate the contribution of each to diversification in two butterfly families: Pieridae and Nymphalidae. Overall, we find that variability in host use is essential for butterfly diversification, while radiations following colonisation of a new host are rare but can produce high diversity. Beyond providing an important reconciliation of alternative hypotheses for butterfly diversification, our approach has potential to test many other hypotheses in evolutionary biology.
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