Seasonal timing of life stages often co-evolves with other ecological traits. Among temperate Lepidoptera, moths flying in early spring or late autumn have been suggested to share a suite of correlated traits – including female flightlessness, spring-feeding larvae, woody host plants, and forest habitats – collectively described as the “winter moth syndrome”. Using a newly available complete phylogeny of northern European Geometridae, we applied phylogenetic comparative methods to reveal the ecological correlates of female flightlessness and, more generally, “winter flight”. Further, we aimed to identify traits that predispose lineages to the evolution of such phenotypes. Our analyses confirm that adult phenology is the strongest predictor of female flightlessness, whereas other traits show weaker or inconsistent associations once phylogeny is accounted for. Sister clades of flightless taxa were significantly more likely to have spring-feeding larvae and, in younger lineages, to have larger body sizes and adult activity shifted towards colder months, suggesting that phenology is among the key preconditions for the loss of flight. We propose that the key adaptive shift underlying the evolution of winter flight and female flightlessness is the abandonment of selective oviposition, with wing reduction evolving secondarily in response to predation and abiotic conditions characteristic of the extreme timing of adult flight. Nevertheless, while the traits facilitating the evolution of female flightlessness in winter moths appear reasonably well understood, evaluating various selective pressures potentially responsible for this evolutionary shift per se still remains a challenge.

Inter-annual variation in insect abundance and seasonal phenology can be related to species traits such as body size, larval diet, overwintering stage, and colour variation. We sampled noctuid moths in the canopy of a forest in Western Poland using flight-interception traps during two vegetative seasons. We calculated inter-annual variation for 31 species, and the date of peak abundance and length of the flight season for 18 species. We found that for the 23 species for which we had phylogenetic information, larger moths and those with adult colour variation showed less inter-annual abundance variation, which corroborates the results of previous studies. We found no indication that phenological traits are associated with the tested species traits. However, species that can feed on the dominant broad-leaved tree in which the traps were placed (Quercus petraea) tended to have a later date of peak abundance than other species. To draw more robust conclusions, future research should encompass a longer time span and a broader range of species.

Understanding the movement patterns of organisms is crucial for effective biodiversity conservation in the increasingly dynamic and fragmented landscapes. Since the colonization of habitat patches relies largely on females, sex differences in movement capability must also be considered. However, obtaining direct measurements of mobility and dispersal, and sex biases in these traits, is often challenging. This underscores the importance of predicting sex‐specific estimates of mobility based on species' functional traits. Our phylogenetic comparative study aims at identifying species traits that could serve as proxies for sex bias in mobility in flying insects. We rely on a comprehensive dataset on the sex ratios of 454 moth species (Lepidoptera: Macroheterocera), captured by light traps of the Finnish national moth monitoring scheme. We first demonstrate that, in the vast majority of species, males outnumber females among the captured individuals. Our phylogenetic regression models reveal that species‐specific sex ratios correlate with traits typically predicted to be associated with sex differences in mobility. Female proportions decrease as sexual dimorphism in wingspan becomes more male‐biased and female wing loading relative to males increases. Proportions of females are also lower in larger species. Females are particularly scarce in trap samples of species in which the reproductive output of females is primarily determined by larval‐derived resources (i.e. capital breeders). These associations suggest that the observed variations in sex ratios do indeed mirror the variation in sex bias in mobility across species. Our findings highlight the potential of trait‐based approaches to identify meaningful indicators of insect mobility, including sex biases in movement patterns. The availability of such proxies facilitates predictions about how different species might respond to contemporary challenges, such as light pollution and habitat loss and fragmentation. The detected associations also advance ordination schemes of insect life histories by integrating mobility measures into relevant analyses.

ABSTRACT Substantial progress has been made to map biodiversity and its drivers across the planet at multiple scales, yet studies that quantify the evolutionary processes that underpin this biodiversity, and test their drivers at multiple scales, are comparatively rare. Studying most fish species, we quantify rates of body size evolution to test the role of fundamental salinity habitats in shaping rates of evolution at multiple scales. We also determine how four additional factors shape evolutionary rates. In up to 1710 comparisons studying over 27,000 ray-finned fish species, we compare rates of body size evolution between five salinity habits using 12 metrics. The comparisons span a molecular tree, supertrees, and ten scales of observation to test for robust patterns and reveal how patterns change with scale. Then, three approaches assess the role of three non-salinity factors on rates, and an alternative habitat scheme tests if lakes influence evolutionary rates. Rates of size evolution rarely differ consistently between salinity habitats; rate patterns are highly clade- and scale-dependent. One exception is freshwater-brackish fishes, which possess among the highest size rates of any salinity, and show higher rates than euryhaline fishes in most groupings studied at most scales, and verses marine, freshwater, and marine-brackish habitats at specific scales. Additionally, species richness had the greatest potential to predict phenotypic rates, followed by branch duration and absolute values of body size. Lacustrine environments were consistently associated with high rates of size evolution. We reveal the rate patterns that underpin global body size diversity for fishes, identifying factors that play a limited role in shaping rates of size evolution, such as salinity, and those such as species richness, age, and lake environments that consistently shape evolutionary rates across half of vertebrate diversity.

Locomotory performance is an important determinant of fitness in most animals, including flying insects. Strong selective pressures on wing morphology are therefore expected. Previous studies on wing shape in Lepidoptera have found some support for hypotheses relating wing shape to environment-specific selective pressures on aerodynamic performance. Here, we present a phylogenetic comparative study on wing shape in the lepidopteran family Geometridae, covering 374 species of the northern European fauna. We focused on 11 wing traits including aspect ratio, wing roundness, and the pointedness of the apex, as well as the ratio of forewing and hindwing areas. All measures were taken from images available on the internet, using a combination of tools available in Fiji software and R. We found that wing shape demonstrates a phylogenetically conservative pattern of evolution in Geometridae, showing similar or stronger phylogenetic signal than many of its potential predictors. Several wing traits showed statistically significant associations with predictors such as body size, phenology, and preference for forest habitats. Overall, however, all of these associations remained notably weak, with no wing shape being excluded for any value of the predictors, including body size. We conclude that, in geometrids, wing traits do not readily respond to selective pressures optimizing aerodynamic performance of the moths in different environments. Selection on wing shape may nevertheless operate through other functions of the wings, with the effectiveness of crypsis at rest being a promising candidate for further studies.

Ecological and life-history data on the Northern European macromoth (Lepidoptera: Macroheterocera) fauna is widely available and ideal for use in answering phylogeny-based research questions: for example, in comparative biology. However, phylogenetic information for such studies lags behind. Here, as a synthesis of all currently available phylogenetic information on the group, we produce a supertree of 114 Northern European macromoth genera (in four superfamilies, with Geometroidea considered separately), providing the most complete phylogenetic picture of this fauna available to date. In doing so, we assess those parts of the phylogeny that are well resolved and those that are uncertain. Furthermore, we identify those genera for which phylogenetic information is currently too poor to include in such a supertree, or entirely absent, as targets for future work. As an aid to studies involving these genera, we provide information on their likely positions within the macromoth tree. With phylogenies playing an ever more important role in the field, this supertree should be useful in informing future ecological and evolutionary studies.

Understanding how and why some groups have become more species-rich than others, and how past biogeography may have shaped their current distribution, are questions that evolutionary biologists have long attempted to answer. We investigated diversification patterns and historical biogeography of a hyperdiverse lineage of Lepidoptera, the geometrid moths, by studying its most species-rich tribe Boarmiini, which comprises ca. 200 genera and ca. known 3000 species. We inferred the evolutionary relationships of Boarmiini based on a dataset of 346 taxa, with up to eight genetic markers under a maximum likelihood approach. The monophyly of Boarmiini is strongly supported. However, the phylogenetic position of many taxa does not agree with current taxonomy, although the monophyly of most major genera within the tribe is supported after minor adjustments. Three genera are synonymized, one new combination is proposed, and four species are placed in incertae sedis within Boarmiini. Our results support the idea of a rapid initial diversification of Boarmiini, which also implies that no major taxonomic subdivisions of the group can currently be proposed. A time-calibrated tree and biogeographical analyses suggest that boarmiines appeared in Laurasia ca. 52 Mya, followed by dispersal events throughout the Australasian, African and Neotropical regions. Most of the transcontinental dispersal events occurred in the Eocene, a period of intense geological activity and rapid climate change. Diversification analyses showed a relatively constant diversification rate for all Boarmiini, except in one clade containing the species-rich genus Cleora. The present work represents a substantial contribution towards understanding the evolutionary origin of Boarmiini moths. Our results, inevitably biased by taxon sampling, highlight the difficulties with working on species-rich groups that have not received much attention outside of Europe. Specifically, poor knowledge of the natural history of geometrids (particularly in tropical clades) limits our ability to identify key innovations underlying the diversification of boarmiines.

1. 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

Specificity is one of the fundamental concepts in ecology. Host specificity of phytophagous insects has been of particular interest because of its crucial role in diversification and life-history evolution. However, the majority of tropical insects remain insufficiently explored with respect to their host-plant relations. A lack of respective data is also hindering the debate over whether higher levels of host-plant specificity prevail in tropical insects compared to temperate ones. We investigated host-plant specificity of forest geometrid moths (Lepidoptera: Geometridae) in equatorial Africa using host-plant acceptability trials with neonate larvae, with the addition of field observations. We compare our experimental data to the (well-known) host-specificity patterns of closely related temperate (hemiboreal) species. Similarly to the temperate region, there were broadly polyphagous tropical species in several clades of Geometridae utilizing hosts belonging to different plant families. Phylogenetic comparative analysis returned no significant differences in host specificity between the two regions. Our study contributes to the evidence that host-plant specificity of herbivores is not necessarily substantially higher in tropical than temperate regions.

1. Comparative studies on insect life histories are facilitated by the increasing availability of reliable phylogenies but are hampered by the scarcity of comparable data. Fortunately, morphological proxies of some life‐history traits can be measured on preserved specimens.2. This study compared values of size‐related life‐history traits among a tropical (Ugandan) and a temperate (Estonian) assemblage of geometrid moths.3. A comparative analysis based on an originally derived phylogeny revealed that tropical moths were, on average, larger than temperate ones. Tropical moths also had somewhat lower relative abdomen masses than temperate ones. This indicates that the tropical rather than the temperate moths tend to use an income (rather than capital) breeding strategy. Nevertheless, no difference was found in a related index of pro‐ovigeny. When body size was accounted for, tropical moths were found to lay smaller eggs than temperate ones.4. The differences between the two compared areas are consistent with selection on higher mobility of the moths imposed by the more diverse tropical vegetation. Relatively larger eggs of temperate moths may constitute an adaptation to overcome the presumably stronger quantitative defences of their host plants.5. Overall, however, we conclude that the differences in ecologically relevant size‐related traits are relatively low among moth assemblages of a tropical and a temperate forest region, indicating that these environments may not impose radically different selective pressures on insect life histories.

Diet breadth and the degree of capital breeding have been established as major determinants of species-specific ecology of herbivorous insects. Both of these variables are related to resource foraging and therefore can be expected to have effects on sensory capacity. However, such effects have remained poorly studied, let alone in phylogenetically explicit multi-species comparisons. We contribute to filling this gap in a study of 60 species of geometrid moths (Lepidoptera: Geometridae), using adult head measures (eye size, antennal length and shape, forehead width) as indices of sensory capacity. When controlled for body size, eyes of the more capital breeding species (i.e. those with low contribution of adult feeding to reproduction) were found to be smaller, and female antennae shorter, than in income breeders. Feathery (vs simple filiform) male antennae were more frequently present in the capital breeders and in larger species. Regarding diet breadth, generalist species were found to have relatively wider male foreheads than specialists. The results suggest that (a) breeding strategy rather than diet breadth predicts sensory capacity in geometrids, (b) capital breeding (vs income breeding) is related to low sensory capacity in females, and (c) in contrast, males of the capital breeding species have evolved towards elevated olfactory capacity.

Evading predators is a fundamental aspect of the ecology and evolution of all prey animals. In studying the influence of prey traits on predation risk, previous researchers have shown that crypsis reduces attack rates on resting prey, predation risk increases with increased prey activity, and rapid locomotion reduces attack rates and increases chances of surviving predator attacks. However, evidence for these conclusions is nearly always based on observations of selected species under artificial conditions. In nature, it remains unclear how defensive traits such as crypsis, activity levels and speed influence realized predation risk across species in a community. Whereas direct observations of predator-prey interactions in nature are rare, insight can be gained by quantifying bodily damage caused by failed predator attacks. We quantified how butterfly species traits affect predation risk in nature by determining how defensive traits correlate with wing damage caused by failed predation attempts, thereby providing the first robust multi-species comparative analysis of predator-induced bodily damage in wild animals. For 34 species of fruit-feeding butterflies in an African forest, we recorded wing damage and quantified crypsis, activity levels and flight speed. We then tested for correlations between damage parameters and species traits using comparative methods that account for measurement error. We detected considerable differences in the extent, location and symmetry of wing surface loss among species, with smaller differences between sexes. We found that males (but not females) of species that flew faster had substantially less wing surface loss. However, we found no correlation between cryptic coloration and symmetrical wing surface loss across species. In species in which males appeared to be more active than females, males had a lower proportion of symmetrical wing surface loss than females. Our results provide evidence that activity greatly influences the probability of attacks and that flying rapidly is effective for escaping pursuing predators in the wild, but we did not find evidence that cryptic species are less likely to be attacked while at rest.

Specificity (= the degree of ecological specialisation) is one of the fundamental concepts of the science of ecology. Ambiguities on how to define and measure specificity have however complicated respective research efforts. Here we propose that, in insects, a behavioural trait – adult oviposition latency in captivity without a favourable host plant – correlates with a species’ specificity in larval host use. In the absence of a suitable host, monophagous insects are expected to wait for a long time before commencing oviposition, with the long waiting time corresponding to careful host location behaviour in nature. Polyphagous insects, in contrast, should be selected for an increased oviposition rate at the expense of the quality of oviposition substrate encountered and will on average have a short latency time. Using experimentally derived data on oviposition latency, we performed a phylogenetically informed analysis based on Bayesian inference to show that this variable correlates with host specificity (larval diet breadth) in a sample of north European species of geometrid moths. A closely related index – the probability to lay any eggs on an unfavourable substrate – shows an analogous pattern. To provide an example of how these indices can be applied, we compare our sample of geometrid moths from northern Europe with a sample from equatorial Africa. A comparative analysis based on an original phylogenetic reconstruction found no differences between the two study sites in parameters of oviposition behaviour. We conclude that behavioural tests can provide information about ecological interactions when the latter can not be directly recorded. Our example study also hints at the possibility that host specificity of herbivores is not necessarily higher in a tropical region compared to a temperate one.

Research on evolutionary forces determining optimal body sizes has primarily relied on experimental evaluation of respective selective pressures. Accounting for among-species variation through application of phylogenetic comparative methods is a complementary although little used approach. It enables the direct association of body size values with particular environments. Using phylogenetically explicit comparative analyses, we show that small body size is associated with diurnal (rather than nocturnal) activity of adults among temperate species of the moth family Geometridae. The association of an exclusively adult trait with species-specific body size suggests that optimal body sizes are at least partly determined by the costs being a large adult, as opposed to the more frequently considered costs of attaining large size. It appears likely that size-selective predation by insectivorous birds is the primary factor responsible for selection against large body size in day-flying moths.

Both physiologically and ecologically based explanations have been proposed to account for among-species differences in lifespan, but they remain poorly tested. Phylogenetically explicit comparative analyses are still scarce and those that exist are biased towards homoeothermic vertebrates. Insect studies can significantly contribute as lifespan can feasibly be measured in a high number of species, and the selective forces that have shaped it may differ largely between species and from those acting on larger animals. We recorded adult lifespan in 98 species of geometrid moths. Phylogenetic comparative analyses were applied to study variation in species-specific values of lifespan and to reveal its ecological and life-history correlates. Among-species and between-gender differences in lifespan were found to be notably limited; there was also no evidence of phylogenetic signal in this trait. Larger moth species were found to live longer, with this result supporting a physiological rather than ecological explanation of this relationship. Species-specific lifespan values could not be explained by traits such as reproductive season and larval diet breadth, strengthening the evidence for the dominance of physiological determinants of longevity over ecological ones.

Predictive classifications of life histories are essential for evolutionary ecology. While attempts to apply a single approach to all organisms may be overambitious, recent advances suggest that more narrow ordination schemes can be useful. However, these schemes mostly lack easily observable proxies of the position of a species on respective axes. It has been proposed that, in insects, the degree of capital (vs. income) breeding, reflecting the importance of adult feeding for reproduction, correlates with various ecological traits at the level of among-species comparison. We sought to prove these ideas via rigorous phylogenetic comparative analyses. We used experimentally derived life-history data for 57 species of European Geometridae (Lepidoptera), and an original phylogenetic reconstruction. The degree of capital breeding was estimated based on morphological proxies, including relative abdomen size of females. Applying Brownian-motion-based comparative analyses (with an original update to include error estimates), we demonstrated the associations between the degree of capital breeding and larval diet breadth, sexual size dimorphism, and reproductive season. Ornstein-Uhlenbeck model based phylogenetic analysis suggested a causal relationship between the degree of capital breeding and diet breadth. Our study indicates that the gradation from capital to income breeding is an informative axis to ordinate life-history strategies in flying insects which are affected by the fecundity vs. mobility trade off, with the availability of easy to record proxies contributing to its predictive power in practical contexts.

Numerous studies have suggested a general relationship between the degree of host specialization and body size in herbivorous animals. In insects, smaller species are usually shown to be more specialized than larger-bodied ones. Various hypotheses have attempted to explain this pattern but rigorous proof of the body sizediet breadth relationship has been lacking, primarily because the scarceness of reliable phylogenetic information has precluded formal comparative analyses. Explicitly using phylogenetic information for a group of herbivores (geometrid moths) and their host plant range, we perform a comparative analysis to study the body sizediet breadth relationship. Considering several alternative measures of body size and diet breadth, our results convincingly demonstrate without previous methodological issuesa first for any taxona positive association between these traits, which has implications for evaluating various central aspects of the evolutionary ecology of herbivorous insects. We additionally demonstrate how the methods used in this study can be applied in assessing hypotheses to explain the body sizediet breadth relationship. By analyzing the relationship in tree-feeders alone and finding that the positive relationship remains, the result suggests that the body sizediet breadth relationship is not solely driven by the type of host plant that species feed on.

Background: The fossil record has suggested that clade growth may differ in marine and terrestrial taxa, supporting equilibrial models in the former and expansionist models in the latter. However, incomplete sampling may bias findings based on fossil data alone. To attempt to correct for such bias, we assemble phylogenetic supertrees on one of the oldest clades of insects, the Odonatoidea (dragonflies, damselflies and their extinct relatives), using MRP and MRC. We use the trees to determine when, and in what clades, changes in taxonomic richness have occurred. We then test whether equilibrial or expansionist models are supported by fossil data alone, and whether findings differ when phylogenetic information is used to infer gaps in the fossil record.Results: There is broad agreement in family-level relationships between both supertrees, though with some uncertainty along the backbone of the tree regarding dragonflies (Anisoptera). "Anisozygoptera" are shown to be paraphyletic when fossil information is taken into account. In both trees, decreases in net diversification are associated with species-poor extant families (Neopetaliidae, Hemiphlebiidae), and an upshift is associated with Calopterygidae + Polythoridae. When ghost ranges are inferred from the fossil record, many families are shown to have much earlier origination dates. In a phylogenetic context, the number of family-level lineages is shown to be up to twice as high as the fossil record alone suggests through the Cretaceous and Cenozoic, and a logistic increase in richness is detected in contrast to an exponential increase indicated by fossils alone.Conclusions: Our analysis supports the notion that taxa, which appear to have diversified exponentially using fossil data, may in fact have diversified more logistically. This in turn suggests that one of the major apparent differences between the marine and terrestrial fossil record may simply be an artifact of incomplete sampling. Our results also support previous notions that adult colouration plays an important role in odonate radiation, and that Anisozygoptera should be grouped in a single inclusive taxon with Anisoptera, separate from Zygoptera.

Given that immature and adult insects have different life styles, different target body compositions can be expected. For adults, such targets will also differ depending on life history strategy, and thus vary among the sexes, and in females depend on the degree of capital versus income breeding and ovigeny. Since these targets may in part be approximated by loss of substances upon eclosion, comparing sexual differences in such losses upon eclosion among species that differ in life history would provide insights into insect functional ecology. We studied weight loss in eclosing insects using original data on pupal and adult live weights of 38 species of Lepidoptera (mainly Geometridae) and further literature data on 15 species of Lepidoptera and six representatives of other insect orders, and applied the phylogenetic independent contrasts approach. In addition, data on live and dry weights of pupae of four species of Lepidoptera are presented. We documented that Lepidoptera typically lose a large proportion (20–80%) of their pupal weight upon adult eclosion. Sexual differences in weight loss varied between absent and strongly male biased. Most of the weight loss was water loss, and sexual differences in adult water content correlate strongly with differences in weight loss. Using feeding habits (feeds or does not feed as an adult) and female biased sexual size dimorphism as measures of degree of capital breeding, we found that the difference among the sexes in weight loss tends to be more pronounced in capital breeding species. Additionally, females of more pro-ovigenic species (large proportion of eggs mature upon emergence) tend to have higher water contents. Our results suggests that metamorphosis is generally facilitated by a high water content, while adults excrete water upon eclosion to benefit flight unless water has been allocated to eggs, or is treated as a capital resource for adult survival or future allocation to eggs.