| 1. |
- Esperk, Toomas, et al.
(författare)
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Achieving high sexual dimorphism : insects add instars
- 2007
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Ingår i: Ecological Entomology. ; 32, s. 243-256
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Tidskriftsartikel (refereegranskat)abstract
- 1. In arthropods, the evolution of sexual size dimorphism (SSD) may be constrained by a physiological limit on growth within each particular larval instar. A high SSD could, however, be attained if the larvae of the larger sex pass through a higher number of larval instars.2. Based on a survey of published case studies, the present review shows that sex-related difference in the number of instars is a widespread phenomenon among insects. In the great majority of species with a sexually dimorphic instar number, females develop through a higher number of instars than males.3. Female-biased sexual dimorphism in final sizes in species with sexually dimorphic instar number was found to considerably exceed a previously estimated median value of SSD for insects in general. This suggests a causal connection between high female-biased SSD, and additional instars in females. Adding an extra instar to larval development allows an insect to increase its adult size at the expense of prolonged larval development.4. As in the case of additional instars, SSD is fully formed late in ontogeny, larval growth schedules and imaginal sizes can be optimised independently. No conflict between selective pressures operating in juvenile and adult stages is therefore expected.5. In most species considered, the number of instars also varied within the sexes. Phenotypic plasticity in instar number may thus be a precondition for a sexual difference in instar number to evolve.
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| 2. |
- Esperk, Toomas, et al.
(författare)
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Distinguishing between anticipatory and responsive plasticity in a seasonally polyphenic butterfly
- 2013
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Ingår i: Evolutionary Ecology. - : Springer Science and Business Media LLC. - 0269-7653 .- 1573-8477. ; 27:2, s. 315-332
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Tidskriftsartikel (refereegranskat)abstract
- Seasonal generations of short-lived organisms often differ in their morphological, behavioural and life history traits, including body size. These differences may be either due to immediate effects of seasonally variable environment on organisms (responsive plasticity) or rely on presumably adaptive responses of organisms to cues signalizing forthcoming seasonal changes (anticipatory plasticity). When directly developing individuals of insects are larger than their overwintering conspecifics, the between-generation differences are typically ascribed to responsive plasticity in larval growth. We tested this hypothesis using the papilionid butterly Iphiclides podalirius as a model species. In laboratory experiments, we demonstrated that seasonal differences in food quality could not explain the observed size difference. Similarly, the size differences are not likely to be explained by the immediate effects of ambient temperature and photoperiod on larval growth. The qualitative pattern of natural size differences between the directly developing and diapausing butterflies could be reproduced in the laboratory as a response to photoperiod, indicating anticipatory character of the response. Directly developing and diapausing individuals followed an identical growth trajectory until the end of the last larval instar, with size differences appearing just a few days before pupation. Taken together, various lines of evidence suggest that between-generation size differences in I. podalirius are not caused by immediate effects of environmental factors on larval growth. Instead, these differences rather represent anticipatory plasticity and are thus likely to have an adaptive explanation. It remains currently unclear, whether the seasonal differences in adult size per se are adaptive, or if they constitute co-product of processes related to the diapause. Our study shows that it may be feasible to distinguish between different types of plasticity on the basis of empirical data even if fitness cannot be directly measured, and contributes to the emerging view about the predominantly adaptive nature of seasonal polyphenisms in insects.
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| 3. |
- Esperk, Toomas, et al.
(författare)
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Intraspecific variability in the number of larval instars in insects
- 2007
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Ingår i: J. Economic Entomology. ; 100, s. 627-646
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Tidskriftsartikel (refereegranskat)abstract
- The number of larval instars varies widely across insect species. Although in star number is frequently considered to be invariable within species, intraspecific variability in the number of instars is not an exceptional phenomenon. However, the knowledge has remained fragmentary, and there are no recent attempts to synthesize the results of relevant studies. Based on published case studies, we show that intraspecific variability in the number of larval instars is widespread across insect taxa, occurring in most major orders, in both hemimetabolous and holometabolous insects. We give an overview of various factors that have been observed to affect the number of instars. Temperature, photoperiod, food quality and quantity, humidity, rearing density, physical condition, inheritance, and sex are the most common factors influencing the number of instars. We discuss adaptive scenarios that may provide ultimate explanations for the plasticity in instar number. The data available largely support the compensation scenario, according to which instar number increases in adverse conditions when larvae fail to reach a species-specific threshold size for metamorphosis. However, in Orthoptera and Coleoptera, there are some exceptional species in which instar number is higher in favorable conditions. In more specific cases, the adaptive value of the variability in instar number may be in reaching or maintaining the developmental stage adapted to hibernation, producing additional generations in multivoltine species, or increasing the probability of surviving in long-lasting adverse conditions.
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| 4. |
- Kivelä, Sami M., et al.
(författare)
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Comparative analysis of larval growth in Lepidoptera reveals instar-level constraints
- 2020
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Ingår i: Functional Ecology. - : Wiley. - 0269-8463 .- 1365-2435. ; 34:7, s. 1391-1403
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Tidskriftsartikel (refereegranskat)abstract
- Juvenile growth trajectories evolve via the interplay of selective pressures on age and size at maturity, and developmental constraints. In insects, the moulting cycle is a major constraint on larval growth trajectories. Surface area to volume ratio of a larva decreases during growth, so renewal of certain surfaces by moulting is likely needed for the maintenance of physiological efficiency. A null hypothesis of isometry, implied by Dyar's Rule, would mean that the relative measures of growth remain constant across moults and instars. We studied ontogenetic changes and allometry in instar-specific characteristics of larval growth in 30 lepidopteran species in a phylogenetic comparative framework. Relative instar-specific mass increments (RMI) typically, but not invariably, decreased across instars. Ontogenetic change in RMIs varied among families with little within-family variation. End-of-instar growth deceleration (GD) became stronger with increasing body size across instars. Across-instar change in GD was conserved across taxa. Ontogenetic allometry was generally non-isometric in both RMI and GD. Results indicate that detailed studies on multiple species are needed for generalizations concerning growth trajectory evolution. Developmental and physiological mechanisms affecting growth trajectory evolution show different degrees of evolutionary conservatism, which must be incorporated into models of age and size at maturation.
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| 5. |
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| 6. |
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| 7. |
- Kodandaramaiah, Ullasa, et al.
(författare)
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Phylogeography of the threatened butterfly, the woodland brown Lopinga achine (Nymphalidae Satyrinae) : implications for conservation
- 2012
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Ingår i: Journal of Insect Conservation. - : Springer Science and Business Media LLC. - 1366-638X .- 1572-9753. ; 16:2, s. 305-313
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Tidskriftsartikel (refereegranskat)abstract
- We have studied the phylogeography of the red-listed Palearctic butterfly Lopinga achine (Nymphalidae: Satyrinae) based on 1,450 base pairs of mitochondrial DNA sequences from 86 individuals representing 12 populations. Our results indicate a strong structuring of genetic variation, with among-population differences accounting for ca. 67% of the variation and almost all populations being significantly differentiated from each other. We surmise that the insular nature of populations as well as the low dispersal ability of the species has given rise to such a pattern. The genetic diversity within populations is low compared to that in other butterflies. Our results point to a scenario where the species originated in the Eastern Palearctic and expanded into Europe. Based on the analyses, we suggest that the Czech population merits the highest conservation priority. The two Swedish populations represent a distinct evolutionary lineage, and hence merit high conservation attention. The Estonian and Asian populations had the highest genetic diversity, and although we do not consider them to be under immediate threat, their genetic diversity should be conserved in the long term.
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| 8. |
- Lindman, Ly, et al.
(författare)
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Host plant relationships of an endangered butterfly, Lopinga achine (Lepidoptera: Nymphalidae) in northern Europe
- 2013
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Ingår i: Journal of Insect Conservation. - : Springer Science and Business Media LLC. - 1366-638X .- 1572-9753. ; 17:2, s. 375-383
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Tidskriftsartikel (refereegranskat)abstract
- The aim of the present study was to evaluate-in a geographic perspective-the role of host plant as a determinant of habitat quality for Lopinga achine, a satyrine butterfly endangered over much of its European range. Laboratory trials were performed to record host choices made by the ovipositing females as well as by neonate larvae. In rearing experiments, growth performance and mortality on different host plants was determined. Oviposition was found to be indiscriminate but larvae were shown to be able to choose between host plants, with the choices made broadly consistent with growth performance of the larvae on particular hosts. Nevertheless, most grasses and sedges offered were found to support larval development reasonably well. No clear superiority of the previously suggested primary host plant Carex montana could be shown. Importantly, no differences in host plant relationships were found between the populations of Sweden, western Estonia and eastern Estonia. In particular, the larvae originating from eastern Estonian populations developed on C. montana equally well even if the plant is absent from their native habitat. In the context of species conservation, one should conclude that L. achine is polyphagous enough on various grasses and sedges so that the presence of any particular host species cannot be a critical component of habitat quality. Nevertheless, some preference to broad- and soft-leaved hosts, as well as sensitivity to host wilting, may partly explain the butterfly's preference to moist forest habitats, further emphasizing the central role of habitat management in the conservation practice of this species. In turn, the absence of ecological differences between geographic populations should enable conservationists to successful transfer their experience across national boundaries.
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| 9. |
- Lundhagen, Anna, et al.
(författare)
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Are peripheral populations special? Congruent patterns in two butterfly species
- 2009
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Ingår i: Ecography. - Malden : Wiley-Blackwell. - 0906-7590 .- 1600-0587. ; 32:4, s. 591-600
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Tidskriftsartikel (refereegranskat)abstract
- Populations at range margins may be genetically different from more central ones for a number of mutually non-exclusive reasons. Specific selection pressures may operate in environments that are more marginal for the species. Genetic drift may also have a strong effect in these populations if they are small, isolated and/or have experienced significant bottlenecks during the colonisation phase. The question if peripheral populations are special, and if yes then how and why, is of obvious relevance for speciation theory, as well as for conservation biology. To evaluate the uniqueness of populations at range margins and the influence of gene flow and selection, we performed a morphometric study of two grassland butterfly species: Coenonympha arcania and C. hero (Lepidoptera: Nymphalidae). The samples were collected from Swedish populations that are peripheral and isolated from the main area of the species distributions and from populations in the Baltic states that are peripheral but connected to the main area of the species distributions. These samples were compared to those from central parts of the species distributions. The isolated populations in both species differed consistently from both peripheral and central populations in their wing size and shape. We interpret this as a result of selection caused by differences in population structure in these isolated locations, presumably favoring different dispersal propensity of these butterflies. Alternative explanations based on colonisation history, latitudinal effects, inbreeding or phenotypic plasticity appear less plausible. As a contrast, the much weaker and seemingly random among-region differences in wing patterns are more likely to be ascribed to weaker selection pressures allowing genetic drift to be influential. In conclusion, both morphological data and results from neutral genetic markers in earlier studies of the same system provide congruent evidence of both adaptation and genetic drift in the isolated Swedish populations of both species.
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| 10. |
- Maes, Dirk, et al.
(författare)
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Integrating national Red Lists for prioritising conservation actions for European butterflies
- 2019
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Ingår i: Journal of Insect Conservation. - : Springer Science and Business Media LLC. - 1366-638X .- 1572-9753. ; 23:2, s. 301-330
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Tidskriftsartikel (refereegranskat)abstract
- Red Lists are very valuable tools in nature conservation at global, continental and (sub-) national scales. In an attempt to prioritise conservation actions for European butterflies, we compiled a database with species lists and Red Lists of all European countries, including the Macaronesian archipelagos (Azores, Madeira and Canary Islands). In total, we compiled national species lists for 42 countries and national Red Lists for 34 of these. The most species-rich countries in Europe are Italy, Russia and France with more than 250 species each. Endemic species are mainly found on the Macaronesian archipelagos and on the Mediterranean islands. By attributing numerical values proportionate to the threat statuses in the different national Red List categories, we calculated a mean Red List value for every country (cRLV) and a weighted Red List value for every species (wsRLV) using the square root of the country’s area as a weighting factor. Countries with the highest cRLV were industrialised (NW) European countries such as the Netherlands, Belgium, the Czech Republic and Denmark, whereas large Mediterranean countries such as Spain and Italy had the lowest cRLV. Species for which a Red List assessment was available in at least two European countries and with a relatively high wsRLV (≥ 50) are Colias myrmidone, Pseudochazara orestes, Tomares nogelii, Colias chrysotheme and Coenonympha oedippus. We compared these wsRLVs with the species statuses on the European Red List to identify possible mismatches. We discuss how this complementary method can help to prioritise butterfly conservation on the continental and/or the (sub-)national scale.
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