| 1. |
- Berger, David, et al.
(författare)
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Ecological Constraints on Female Fitness in a Phytophagous Insect
- 2012
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Ingår i: American Naturalist. - : University of Chicago Press. - 0003-0147 .- 1537-5323. ; 180:4, s. 464-480
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Tidskriftsartikel (refereegranskat)abstract
- Although understanding female reproduction is crucial for population demography, determining how and to what relative extent it is constrained by different ecological factors is complicated by difficulties in studying the links between individual behavior, life history, and fitness in nature. We present data on females in a natural population of the butterfly Leptidea sinapis. These data were combined with climate records and laboratory estimates of life-history parameters to predict the relative impact of different ecological constraints on female fitness in the wild. Using simulation models, we partitioned effects of male courtship, host plant availability, and temperature on female fitness. Results of these models indicate that temperature is the most constraining factor on female fitness, followed by host plant availability; the short-term negative effects of male courtship that were detected in the field study were less important in models predicting female reproductive success over the entire life span. In the simulations, females with more reproductive reserves were more limited by the ecological variables. Reproductive physiology and egg-laying behavior were therefore predicted to be co-optimized but reach different optima for females of different body sizes; this prediction is supported by the empirical data. This study thus highlights the need for studying behavioral and life-history variation in orchestration to achieve a more complete picture of both demographic and evolutionary processes in naturally variable and unpredictable environments.
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| 2. |
- Berger, David, et al.
(författare)
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Intraspecific variation in body size and the rate of reproduction in female insects- adaptive allometry or biophysical constraint?
- 2012
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Ingår i: Journal of Animal Ecology. - : Wiley. - 0021-8790 .- 1365-2656. ; 81:6, s. 1244-1258
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Tidskriftsartikel (refereegranskat)abstract
- 1. A high rate of reproduction may be costly if ecological factors limit immediate reproductive output as a fast metabolism compromises own future survival. Individuals with more reserves need more time and opportunity to realize their reproductive potential. Theory therefore predicts that the reproductive rate, defined as the investment in early reproduction in proportion to total potential, should decrease with body size within species. 2. However, metabolic constraints on body size- and temperature-dependent biological rates may impede biophysical adaptation. Furthermore, the sequential manner resources that are allocated to somatic vs. reproductive tissue during ontogeny may, when juveniles develop in unpredictable environments, further contribute to non-adaptive variation in adult reproductive rates. 3. With a model on female egg laying in insects, we demonstrate how variation in body reserves is predicted to affect reproductive rate under different ecological scenarios. Small females always have higher reproductive rates but shorter lifespans. However, incorporation of female host selectivity leads to more similar reproductive rates among female size classes, and oviposition behaviour is predicted to co-evolve with reproductive rate, resulting in small females being more selective in their choice and gaining relatively more from it. 4. We fed simulations with data on the butterfly Pararge aegeria to compare model predictions with reproductive rates of wild butterflies. However, simulated reproductive allometry was a poor predictor of that observed. Instead, reproductive rates were better explained as a product of metabolic constraints on rates of egg maturation, and an empirically derived positive allometry between reproductive potential and size. However, fitness is insensitive to moderate deviations in reproductive rate when oviposition behaviour is allowed to co-evolve in the simulations, suggesting that behavioural compensation may mitigate putative metabolic and developmental constraints. 5. More work is needed to understand how physiology and development together with compensatory behaviours interact in shaping reproductive allometry. Empirical studies should evaluate adaptive hypotheses against proper null hypotheses, including prediction from metabolic theory, preferentially by studying reproductive physiology in combination with behaviour. Conversely, inferences of constraint explanations on reproductive rates must take into consideration that adaptive scenarios may predict similar allometric exponents.
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| 3. |
- Friberg, Magne, et al.
(författare)
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Asymmetric life-history decision-making in butterfly larvae
- 2011
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Ingår i: Oecologia. - : Springer Science and Business Media LLC. - 0029-8549 .- 1432-1939. ; 165:2, s. 301-310
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Tidskriftsartikel (refereegranskat)abstract
- In temperate environments, insects appearing in several generations in the growth season typically have to decide during the larval period whether to develop into adulthood, or to postpone adult emergence until next season by entering a species-specific diapause stage. This decision is typically guided by environmental cues experienced during development. An early decision makes it possible to adjust growth rate, which would allow the growing larva to respond to time stress involved in direct development, whereas a last-minute decision would instead allow the larva to use up-to-date information about which developmental pathway is the most favourable under the current circumstances. We study the timing of the larval pathway decision-making between entering pupal winter diapause and direct development in three distantly related butterflies (Pieris napi, Araschnia levana and Pararge aegeria). We pinpoint the timing of the larval diapause decision by transferring larvae from first to last instars from long daylength (inducing direct development) to short daylength conditions (inducing diapause), and vice versa. Results show that the pathway decision is typically made in the late instars in all three species, and that the ability to switch developmental pathway late in juvenile life is conditional; larvae more freely switched from diapause to direct development than in the opposite direction. We contend that this asymmetry is influenced by the additional physiological preparations needed to survive the long and cold winter period, and that the reluctance to make a late decision to enter diapause has the potential to be a general trait among temperate insects.
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| 4. |
- Friberg, Magne, et al.
(författare)
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Autumn mass change and winter mass loss differ between migratory and nonmigratory butterflies
- 2023
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Ingår i: Animal Behaviour. - 0003-3472 .- 1095-8282. ; 204, s. 67-75
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Tidskriftsartikel (refereegranskat)abstract
- Poikilotherms have two major alternative strategies to deal with the inhospitable conditions of temperate winters: hibernation or migration. The vast majority of insects spend winter in a speciesspecific diapause resting stage, while others remain reproductively active after having migrated to more hospitable environments in the autumn. The butterfly clade Nymphalini offers an interesting possibility for comparisons among species adopting different strategies. Whereas most genera in this clade have evolved adult diapause, species in the genus Vanessa engage in an annual multigeneration migration cycle, leaving northern latitudes during autumn. Here, we compared two species of diapausing Aglais butterflies, A. io and A. urtica, two species of the migratory genus Vanessa, V. atalanta and V. cardui, and two morphs of Polygonia c-album, the diapausing winter morph and the direct developing hutchinsoni morph. We tested how these different species differ in resource acquisition strategy during early adult life, how the acquisition of resources affects survival in cold winter conditions and how A. io, Aglais urticae and V. atalanta differ in resting metabolism under cold conditions. The butterflies set for adult diapause (1) acquired more mass during early adult life and (2) lost less mass and had a lower resting metabolism under cold conditions. In addition, (3) the life span under cold conditions was positively related to the weight increase during early adult life in the diapausing species, but not in the migratory species. Our laboratory results demonstrate how different developmental pathways, such as diapause or direct development, can be studied by measuring mass change in living butterflies. Our expectation that migratory species would increase in mass like species set for overwintering were not met, perhaps because such studies should ideally be performed under natural conditions. (c) 2023 The Author(s). Published by Elsevier Ltd on behalf of The Association for the Study of Animal Behaviour. This is an open access article under the CC BY license (http://creativecommons.org/licenses/ by/4.0/).
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| 5. |
- Friberg, Magne, et al.
(författare)
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Butterflies and plants : preference/performance studies in relation to plant size and the use of intact plants vs. cuttings
- 2016
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Ingår i: Entomologia Experimentalis et Applicata. - : Wiley. - 0013-8703 .- 1570-7458. ; 160:3, s. 201-208
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Tidskriftsartikel (refereegranskat)abstract
- Plants have evolved a number of defences to ameliorate herbivore attacks including chemicals induced by mechanical wounding. Such changes in plant chemical composition are potential confounding factors in experiments on plant-insect interactions, which often present cuttings of potential host plants to phytophagous insects. In particular, this could affect studies of female egg-laying preference and larval performance, because the same plant chemicals that deter certain generalist insects can elevate attacks from more specialized insects. Furthermore, plant cuttings are by definition smaller than intact plants, and any female host size preference could thus affect experiments using plant cuttings. We first assessed female preference and larval performance of a specialist herbivore, Pieris napi (L.) (Lepidoptera: Pieridae, Pierini), confronted with either intact plants or leaf-cuttings of four Brassicaceae host plants, Alliaria petiolata (Bieb.) Cavara & Grande, Barbarea vulgaris (L.) WT Aiton, Berteroa incana (L.) DC., and Brassica napus (L.). Egg and larval survival did not differ between intact plants and leaf-cuttings, whereas larval growth was slightly, but significantly, faster on leaf-cuttings. Females, however, significantly preferred to lay eggs on intact plants of all four hosts, although the preference hierarchy for the intact plants was largely mirrored by that for leaf-cuttings. We then tested the female preference for different size-classes of intact B. napus plants. Small individuals received more eggs than larger individuals, and follow-up experiments showed that this difference was largely generated by a strong female preference for cotyledon leaves; there was no significant difference in female preference for large and small individuals when both carried cotyledons, and females landing on cotyledons were more likely to oviposit compared to when landing on a true leaf. Our study concludes that plant cuttings can serve as adequate proxies for live plants for preference/performance studies, but that experimentalists should be aware of the variation imposed both by plant handling and plant phenology for female oviposition preference.
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| 6. |
- Friberg, Magne, et al.
(författare)
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Decoupling of female host plant preference and offspring performance in relative specialist and generalist butterflies
- 2015
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Ingår i: Oecologia. - : Springer Science and Business Media LLC. - 0029-8549 .- 1432-1939. ; 178:4, s. 1181-1192
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Tidskriftsartikel (refereegranskat)abstract
- The preference-performance hypothesis posits that the host plant range of plant-feeding insects is ultimately limited by larval costs associated with feeding on multiple resources, and that female egg-laying preferences evolve in response to these costs. The trade-off of either using few host plant species and being a strong competitor on them due to effective utilization or using a wide host plant range but being a poor competitor is further predicted to result in host plant specialization. This follows under the hypothesis that both females and offspring are ultimately favoured by utilizing only the most suitable host(s). We develop an experimental approach to identify such trade-offs, i.e. larval costs associated with being a host generalist, and apply a suite of experiments to two sympatric and syntopic populations of the closely related butterflies Pieris napi and Pieris rapae. These butterflies show variation in their level of host specialization, which allowed comparisons between more and less specialized species and between families within species. Our results show that, first, the link between female host preference and offspring performance was not significantly stronger in the specialist compared to the generalist species. Second, the offspring of the host plant specialist did not outperform the offspring of the generalist on the former's most preferred host plant species. Finally, the more generalized species, or families within species, did not show higher survival or consistently higher growth rates than the specialists on the less preferred plants. Thus, the preference and performance traits appear to evolve as largely separated units.
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| 7. |
- Friberg, Magne, et al.
(författare)
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Female mate choice determines reproductive isolation between sympatric butterflies
- 2008
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Ingår i: Behavioral Ecology and Sociobiology. - Berlin : Springer Science and Business Media LLC. - 0340-5443 .- 1432-0762. ; 62:6, s. 873-886
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Tidskriftsartikel (refereegranskat)abstract
- Animal courtship rituals are important for species recognition, and a variety of cues might be utilized to recognize conspecific mates. In this paper, we investigate different species-recognition mechanisms between two sympatric butterfly sister species: the wood white (Leptidea sinapis) and Real's wood white (Leptidea reali). We show that males of both species frequently court heterospecific females both under laboratory and field conditions. The long-lasting elaborate courtships impose energetic costs, since the second courtship of males that were introduced to two subsequent conspecific females lasted on average only one fourth as long as the first courtship. In this paper, we demonstrate that premating reproductive isolation is dependent on female unwillingness to accept heterospecific mates. We studied female and male courtship behavior, chemical signaling, and the morphology of the sexually dimorphic antennae, one of the few male traits visible for females during courtship. We found no differences in ultraviolet (UV) reflectance and only small differences in longer wavelengths and brightness, significant between-species differences, but strongly overlapping distributions of male L. sinapis and L. reali antennal morphology and chemical signals and minor differences in courtship behavior. The lack of clear-cut between-species differences further explains the lack of male species recognition, and the overall similarity might have caused the long-lasting elaborate courtships, if females need prolonged male courtships to distinguish between con- and heterospecific suitors.
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| 8. |
- Friberg, Magne, et al.
(författare)
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Generation-dependent female choice: behavioral polyphenism in a bivoltine butterfly
- 2007
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Ingår i: Behavioral Ecology. - : Oxford University Press (OUP). - 1045-2249 .- 1465-7279. ; 18:4, s. 758-763
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Tidskriftsartikel (refereegranskat)abstract
- Climatic and biotic circumstances vary as seasons shift, and different cohorts of multivoltine species are likely subjected to different selection regimes. The bivoltine butterfly Leptidea reali (Re´al’s wood white; Lepidoptera: Pieridae) appears during May and June in central Sweden and has a partial second generation in late July. We manipulated both generations to appear simultaneously and performed laboratory mating experiments that showed the presence of a behavioral polyphenism in mating propensity, which is induced during the developmental stages. Females of the summer generation expressed higher mating propensities than spring generation females. Spring females showed an increase in mating propensity with increasing age, whereas summer females accepted most matings already when they were only 1 or 2 days old. It is likely that larval time constraints, a lower abundance of males and a lower risk of accepting a male of their univoltine sister species Leptidea sinapis (wood white), have relaxed selection on mate discrimination among summer generation females. A major challenge for future research is to further investigate the developmental pathways causing the polyphenism and the adaptive implications of cohort dependent behaviors.
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| 9. |
- Friberg, Magne, et al.
(författare)
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Genetic differentiation and phylogeographic patterns in European populations of Leptidea sinapis and L. reali
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Annan publikation (refereegranskat)abstract
- The direction of a coevolutionary interaction can differ between local populations as in the butterflies Leptidea sinapis and L. reali. The morphologically virtually identical sister-species have partitioned their niches differently in different parts of their distribution by shifting habitat specialist and generalist roles between different sympatric areas. Hence, a species that is a generalist in some areas can be a local specialist in others, and vice versa. We have sequenced the mitochondrial COI gene from specimens collected across Europe in order to (i) describe the between-species variation over a large area, (ii) identify possible glacial refugia and re-colonisation routes to obtain a phylogeographic hypothesis for explaining the geographic mosaic of niche separation and (iii) apply a population genetic approach to determine the level of intraspecific genetic differentiation. The results show evidence for species distinctiveness throughout Europe. Only small variation was found in L. reali, whereas the haplotype network of L. sinapis showed a deep division into two haplotype families of which one was restricted to Spain and the other was widespread over the continent (including Spain). The widespread haplotype family was divided into two common variants, one eastern and one western, each being surrounded by rare haplotypes. The both deep and shallow genetic differentiation implies that L. sinapis might have been divided into different refugia during several glaciations. Both species showed significant genetic differentiation in pairwise ФST, and as habitat generalist populations could differ significantly from other habitat generalist populations but not from habitat specialist populations, we conclude that this study supports that the geographic mosaic of niche separation is caused by local processes rather than common ancestry of local habitat generalists or specialists within each species
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| 10. |
- Friberg, Magne, et al.
(författare)
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Habitat choice precedes host plant choice - niche separation in a species pair of a generalist and a specialist butterfly
- 2008
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Ingår i: Oikos. - Lund : Department of Ecology, Lund university. ; 117:9, s. 1337-1344
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Tidskriftsartikel (refereegranskat)abstract
- The sister species Leptidea reali and L. sinapis have partitioned their niches differently in different parts of their sympatric distribution. In Spain and France L. sinapis is a widespread generalist whereas L. reali is specialized on high altitude open areas. Interestingly, the reverse is true in Ireland and the Czech Republic where L. reali is widespread and L. sinapis specialized on meadows. In Sweden, L. reali is a habitat specialist confined to meadows, whereas L. sinapis is a habitat generalist also inhabiting forests. Ultimately, the geographic mosaic of niche separation is the result of local processes in each contact zone or a secondary effect of the host plant distribution, if L. sinapis and L. reali prefer different legume host plants. Hence, in Sweden L. sinapis might utilize the forest habitat either due to a wider habitat preference or due to a wider host plant preference than L. reali. Studies of wild butterflies showed that L. sinapis laid 26% of their eggs on forest-associated legumes compared to 6% in L. reali, although laboratory experiments showed that both species had virtually identical host plant preferences strongly preferring the meadow-associated legume Lathyrus pratensis. Furthermore, flight duration tests in a variety of temperatures demonstrated a between-species difference; L. sinapis females reached their flight optimum at a lower temperature than L. reali females. The lower L. sinapis flight temperature optimum is most probably a secondary effect due to habitat-specific selection, and therefore a consequence rather than the cause of the habitat partitioning. The finding that habitat choice precedes host plant choice suggests that the European geographic mosaic of niche separation, with L. sinapis and L. reali shifting habitat specialist/generalist roles, is not caused by rigid between-species differences in a related niche parameter, but instead is a result of local processes within each secondary contact zone.
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