The evolution of exceptional diversity in parental care and fertilization modes in ray-finned fishes.

Among vertebrates, ray-finned fishes (Actinopterygii) display the highest diversity in parental care, and their diversification has been hypothesized to be related to phylogenetic changes in fertilization modes. Using the most comprehensive, sex-specific data from 7600 species of 62 extant orders of ray-finned fishes, we inferred ancestral states and transitions among care types and caring episodes (i.e. the stage of offspring development). Our work has uncovered three novel findings. First, transitions among different care types (i.e. male-only care, female-only care, biparental care and no care) are common, and the frequencies of these transitions show unusually diverse patterns concerning fertilization modes (external, or internal via oviduct, mouth or brood pouch). Second, both oviduct and mouth fertilization select for female-biased care, whereas fertilization in a brood pouch selects for male-biased care. Importantly, internal fertilization without parental care is extremely unstable phylogenetically. Third, we show that egg care in both sexes is associated with nest building (which is male-biased) and fry care (which is female-biased). Taken together, the aquatic environment, which supports considerable flexibility in care, facilitated the diversification of parenting behavior, creating the evolutionary bases for more comprehensive parenting to protect offspring in semiterrestrial or terrestrial environments.

Harem size should be measured by more than the sum of its parts: Phenology-based measurements reveal joint effects of intrinsic and extrinsic factors on a polygamous herbivore under non-stationary climatic conditions.

Social behaviour is thought to be a major component of survival, reproduction, and resilience of populations. Thus, it is a key component in management and conservation of wild populations. In polygynous breeding species, group size influences the reproductive success of males and females, and hence it is essential to understand the environmental and demographic factors that shape the phenology of group size within populations. Here, we investigate harem size and its determinants using a 15-year dataset of annual harem size phenology-based metrics from a reintroduced population of wild Przewalski horses in Hortobágy National Park, Hungary. From the initial reintroduction of 21 animals in 1997, the population grew to 174 animals in 2012. During that same period, the number of harems increased from three to 23. Despite the 8-fold increase in population size, harem sizes remained stable, and variability among harems within years decreased. The annual phenological cycle of harem size was not consistent over the 15-year period, and the associated annual phenology-based metrics varied differently over the years. The best predictors of our phenology-based harem size metrics were adult sex ratio, annual adult mortality and annual mean number of harems, with some evidence that mean age of harem stallions and drought severity were contributing factors. Our findings reveal that complex interactions between demography, climate, and harem size can emerge in social animals. Taken together, our results demonstrate that intrinsic population processes can regulate group size even in the presence of non-stationary climatic conditions during periods of growth in human-introduced, semi-free ranging animal populations.

Opportunistic citizen science data reveals habitat selection of lesser black-backed gulls in Central Europe.

This study is the first attempt to find a relation between the opportunistic observation data of an easily identifiable but locally rare bird, in our case the lesser black-backed gull (Larus fuscus fuscus), and the Corine Land Cover habitat map by applying a resource selection function model to reveal the habitat selection of the species. We provide a scientific analysis using the valuable citizen science dataset sourced from the www.birding.hu website, collected by hundreds of volunteers over a span of more than a decade in Hungary. Birds selected for landfills, river and lake habitats and showed a significant but much smaller attraction towards urban areas, while they selected against arable lands. We found that in the case of rare and moderately common birds, we can obtain a representative picture of the habitat selection of the species, even if the data are collected by non-standardized means.

Evolution of reproductive modes in sharks and rays.

The ecological and life history drivers of the diversification of reproductive modes in early vertebrates are not fully understood. Sharks, rays and chimaeras (group Chondrichthyes) have an unusually diverse variety of reproductive modes and are thus an ideal group to test the factors driving the evolution of reproductive complexity. Here, using 960 species representing all major Chondrichthyes taxa, we reconstruct the evolution of their reproduction modes and investigate the ecological and life history predictors of reproduction. We show that the ancestral Chondrichthyes state was egg-laying and find multiple independent transitions between egg-laying and live-bearing via an intermediate state of yolk-only live-bearing. Using phylogenetically informed analysis, we also show that live-bearing species have larger body size and larger offspring than egg-laying species. In addition, live-bearing species are distributed over shallow to intermediate depths, while egg-layers are typically found in deeper waters. This suggests that live-bearing is more closely associated with pelagic, rather than demersal habitats. Taken together, using a basal vertebrate group as a model, we demonstrat how reproductive mode co-evolves with environmental conditions and life-history traits.

Better safe than sorry - Understanding the attitude and habits of drivers can help mitigating animal-vehicle collisions.

The rapidly growing global road networks put serious pressures on terrestrial ecosystems and increase the number and severity of human-wildlife conflicts, which in most cases manifest in animal-vehicle collisions (AVCs). AVCs pose serious problems both for biodiversity conservation and traffic safety: each year, millions of vertebrates are roadkilled globally and the related economic damage is also substantial. For a comprehensive understanding of factors influencing AVC it is essential to explore the human factor, that is, the habits and attitude of drivers; however, to date, comprehensive surveys are lacking on this topic. Here we addressed this knowledge gap and surveyed the habits of drivers and their experience and attitude towards AVCs by a comprehensive questionnaire covering a large geographical area and involving a large number of respondents (1942 completed questionnaires). We aimed to reveal how driving habits affect the chance of AVC, and explored the attitude of the drivers regarding AVC. We found that the number of lifetime AVC cases was higher for male drivers, for those who drove longer distances per year, had more driven years, used country roads or drove large vehicles. Our results showed that almost half of the drivers surveyed had experienced at least one AVC in their lifetime. Drivers' attitudes towards the importance of nature conservation or traffic safety in the aspect of AVC, and fear of collision showed a significant correlation with experienced AVC cases. Drivers' opinions indicated that the most trusted and desired AVC prevention measures were physical objects such as fences and wildlife crossings. Our research provides guidelines for developing targeted initiatives in the future to increase awareness about the significance of AVC and target those drivers who are most vulnerable to AVC.

Polygamy and purifying selection in birds.

Good genes theories of sexual selection predict that polygamy will be associated with more efficient removal of deleterious alleles (purifying selection), due to the alignment of sexual selection with natural selection. On the other hand, runaway selection theories expect no such alignment of natural and sexual selection, and may instead predict less efficient purifying selection in polygamous species due to higher reproductive variance. In an analysis of polymorphism data extracted from 150-bird genome assemblies, we show that polygamous species carry significantly fewer nonsynonymous polymorphisms, relative to synonymous polymorphisms, than monogamous bird species (p = .0005). We also show that this effect is independent of effective population size, consistent with the alignment of natural selection with sexual selection and "good genes" theories of sexual selection. Further analyses found no impact of polygamy on genetic diversity, while polygamy in females (polyandry) had a marginally significant impact (p = .045). We also recapitulate previous findings that smaller body mass and greater geographic range size are associated with more efficient purifying selection, more intense GC-biased gene conversion, and greater genetic diversity.

Waterbird guilds predict environmental attributes of inland saline aquatic ecosystems on multi-spatial scales.

Inland saline waters in Central Asia constitute an important part of steppe ecosystems, providing considerable ecological functions and ecosystem services. Here we aim to present a multi-spatial scale analysis of nutrient cycling and transport waterbird guilds, and the environmental attributes of saline-soda aquatic ecosystems in Kazakhstan. The density, biomass, and diversity of waterbird guilds was determined in the case of three nutrient cycling and transport guilds: a) net-importer (IM), b) importer-exporter (IMEX), c) net-exporter (EX), according to the Boros's guild classification method, and for several traditional feeding guilds: carnivorous, herbivorous, invertebrate eater, omnivorous and piscivorous. Our results revealed that waterbird guilds, as predictors represented by the complete waterbird community, are in close relationship with several (N = 12) environmental attributes of inland saline waters through complex trophic linkages of waterbird populations on multi-spatial scales. The density and the biomass of the EX and IMEX guilds are strongly and positively correlated (i) with the productivity metrics of habitats (e.g., CHL, GPP), indicating their trophic position, and (ii) with water depth. We found significant correlations among guild density, biomass, diversity and environmental attributes on multi-spatial scales for IMEX and EX. Our results revealed that IMEX predicts the surrounding environment of aquatic habitats, whereas EX species are substantial environmental predictors of aquatic ecosystems. However, the diversity metric had valid models only with EX. The herbivorous and omnivorous guilds, that feed chiefly on plant materials consist mainly of IMEX and EX duck species, which were positively related to grassland coverage and the shoreline development index. As a methodological result, here we present a novel approach, the guild transport index, which has more robust relationships with environmental attributes than individual guilds, thus it provides a complex evaluation of the nutrient cycling by birds between aquatic and terrestrial environments on multi-spatial scales.

The evolution of parental care in salamanders.

Complex parenting has been proposed to contribute to the evolutionary success of vertebrates. However, the evolutionary routes to complex parenting and the role of parenting in vertebrate diversity are still contentious. Although basal vertebrates provide clues to complex reproduction, these are often understudied. Using 181 species that represent all major lineages of an early vertebrate group, the salamanders and newts (Caudata, salamanders henceforth) here we show that fertilisation mode is tied to parental care: male-only care occurs in external fertilisers, whereas female-only care exclusively occurs in internal fertilisers. Importantly, internal fertilisation opens the way to terrestrial reproduction, because fertilised females are able to deposit their eggs on land, and with maternal care provision, the eggs could potentially develop outside the aquatic environment. Taken together, our results of a semi-aquatic early vertebrate group propose that the diversity and follow-up radiation of terrestrial vertebrates are inherently associated with a complex social behaviour, parenting.

Functional composition of ant assemblages in habitat islands is driven by habitat factors and landscape composition.

Fragmented natural habitats within human-transformed landscapes play a key role in preserving biodiversity. Ants as keystone species are essential elements of terrestrial ecosystems; thus, it is important to understand the factors influencing their presence. In a large-scale multi-site study, we surveyed ant assemblages using sweep netting and D-vac sampling on 158 ancient burial mounds preserving grassland habitats in agricultural landscapes in East-Hungary. We asked the following questions: (1) How do habitat factors and landscape composition affect species richness and functional diversity of ants? (2) Which ant traits are affected by habitat factors and landscape composition? Despite their small sizes, mounds as permanent and relatively undisturbed landscape elements could provide safe havens for diverse ant assemblages even in transformed agricultural landscapes. The complex habitat structure of wooded mounds supported high species and functional diversity of ant assemblages. Ant species on wooded mounds had small or medium-sized colonies, enabling the co-existence of more species. The effect of landscape composition on ant assemblages was mediated by habitat factors: steep slopes buffered the negative effect of the cropland matrix and enabled higher ant diversity.

Are evolutionary transitions in sexual size dimorphism related to sex determination in reptiles?

Sex determination systems are highly variable in vertebrates, although neither the causes nor the implications of this diversity are fully understood. Theory suggests that sex determination is expected to relate to sexual size dimorphism, because environmental sex determination promotes sex-specific developmental bias in embryonic growth rates. Furthermore, selection for larger size in one sex or the other has been proposed to drive the evolution of different genetic sex determination systems. Here, we investigate whether sex determination systems relate to adult sexual size dimorphism, using 250 species of reptiles (Squamata, Testudines and Crocodylia) representing 26 families. Using phylogenetically informed analyses, we find that sexual size dimorphism is associated with sex determination: species with TSDIa sex determination (i.e. in which the proportion of female offspring increases with incubation temperature) have more female-biased size dimorphism than species with TSDII (i.e. species in which males are produced at mid temperatures). We also found a trend that species with TSD ancestors had more male-biased size dimorphism in XY sex chromosome systems than in ZW sex chromosome systems. Taken together, our results support the prediction that sexual size dimorphism is linked to sex-dependent developmental variations caused by environmental factors and also by sex chromosomes. Since the extent of size dimorphism is related to various behavioural, ecological and life-history differences between sexes, our results imply profound impacts of sex determination systems for vertebrate diversity.

Long-term temporal patterns in flight activities of a migrant diurnal butterfly.

Recent studies demonstrated that the Painted Lady (Vanessa cardui), a cosmopolitan diurnal butterfly performs long-range migration between subtropical Africa and north-western Europe, covered by individuals belonging to up to six generations. Here we analyze temporal patterns of complete annual migratory activity of the Painted Lady in Hungary, located in its Central European migratory route, almost completely unstudied before. To do so, we used field occurrence data collected between 2000 and 2019 and estimated temporal patterns in migratory activity by fitting kernel density functions on the daily mean number of individuals and observation frequency. The temporal distributions of kernel density estimates were analyzed as a function of time and key climatic predictors of the study area. We found that (i) the timing of spring arrivals has been advancing; (ii) the relative intensity of the first and last migratory peaks of the Painted Lady significantly increased during the past decades; and (iii) intensity of the last migratory peak is related to the mean temperature of the previous month, inferring that the migration is shifting to earlier dates and their volume of the migration has substantially intensified, evoking mutually nonexclusive, competing hypotheses. Our study indicates the strengthening migration activities of a southerly distributed, long-distance migrant diurnal butterfly, most probably linked to the northward shift of wintering areas induced by warming trends of the southern parts of Europe. However, the complexity of the likely processes leading to changing migratory strategies calls up for further research in both breeding and wintering areas.

Incubating parents serve as visual cues to predators in Kentish plover (Charadrius alexandrinus).

Ground-nesting birds face many challenges to reproduce successfully, with nest predation being the main cause of reproductive failure. Visual predators such as corvids and egg-eating raptors, are among the most common causes of nest failure; thus, parental strategies that reduce the risk of visual nest predation should be favored by selection. To date, most research has focused on egg crypsis without considering adult crypsis, although in natural circumstances the eggs are covered by an incubating parent most of the time. Here we use a ground-nesting shorebird, the Kentish plover (Charadrius alexandrinus) as model species to experimentally test whether decoy parents influence nest predation. Using artificial nests with a male decoy, a female decoy or no decoy, we found that the presence of a decoy increased nest predation (N = 107 nests, p < 0.001). However, no difference was found in predation rates between nests with a male versus female decoy (p > 0.05). Additionally, we found that nests in densely vegetated habitats experienced higher survival compared to nests placed in sparsely vegetated habitats. Nest camera images, predator tracks and marks left on eggs identified the brown-necked raven (Corvus ruficollis) as the main visual nest predator. Our study suggests that the presence of incubating parents may enhance nest detectability to visual predators. However, parents may reduce the predation risk by placing a nest in sites where they are covered by vegetation. Our findings highlight the importance of nest site selection not only regarding egg crypsis but also considering incubating adult camouflage.

Parental care and the evolution of terrestriality in frogs.

Frogs and toads (Anura) exhibit some of the most diverse parental strategies in vertebrates. Identifying the evolutionary origins of parenting is fundamental to understanding the relationships between sexual selection, social evolution and parental care systems of contemporary Anura. Moreover, parenting has been hypothesized to allow the invasion of terrestrial habitats by the ancestors of terrestrial vertebrates. Using comprehensive phylogenetic analyses of frogs and toads based on data from over 1000 species that represent 46 out of 55 Anura families, we test whether parental care is associated with terrestrial reproduction and several life-history traits. Here, we show that both the duration of care and offspring protection by males and females have coevolved with terrestrial reproduction. Sexual size dimorphism is also related to care, because the large male size relative to female size is associated with increased paternal care. Furthermore, increased egg size and reduced clutch volume are associated with increased care in bivariate but not in multivariate analyses, suggesting that the relationships between care, egg size and clutch volume are mediated by terrestrial reproduction. Taken together, our results suggest that parenting by males and females has coevolved, and complex parenting traits have evolved several times independently in Anura in response to breeding in terrestrial environments.

Global pattern of nest predation is disrupted by climate change in shorebirds.

Ongoing climate change is thought to disrupt trophic relationships, with consequences for complex interspecific interactions, yet the effects of climate change on species interactions are poorly understood, and such effects have not been documented at a global scale. Using a single database of 38,191 nests from 237 populations, we found that shorebirds have experienced a worldwide increase in nest predation over the past 70 years. Historically, there existed a latitudinal gradient in nest predation, with the highest rates in the tropics; however, this pattern has been recently reversed in the Northern Hemisphere, most notably in the Arctic. This increased nest predation is consistent with climate-induced shifts in predator-prey relationships.

Sex-biased breeding dispersal is predicted by social environment in birds.

Sex-biased dispersal is common in vertebrates, although the ecological and evolutionary causes of sex differences in dispersal are debated. Here, we investigate sex differences in both natal and breeding dispersal distances using a large dataset on birds including 86 species from 41 families. Using phylogenetic comparative analyses, we investigate whether sex-biased natal and breeding dispersal are associated with sexual selection, parental sex roles, adult sex ratio (ASR), or adult mortality. We show that neither the intensity of sexual selection, nor the extent of sex bias in parental care was associated with sex-biased natal or breeding dispersal. However, breeding dispersal was related to the social environment since male-biased ASRs were associated with female-biased breeding dispersal. Male-biased ASRs were associated with female-biased breeding dispersal. Sex bias in adult mortality was not consistently related to sex-biased breeding dispersal. These results may indicate that the rare sex has a stronger tendency to disperse in order to find new mating opportunities. Alternatively, higher mortality of the more dispersive sex could account for biased ASRs, although our results do not give a strong support to this explanation. Whichever is the case, our findings improve our understanding of the causes and consequences of sex-biased dispersal. Since the direction of causality is not yet known, we call for future studies to identify the causal relationships linking mortality, dispersal, and ASR.

Growth-form and spatiality driving the functional difference of native and alien aquatic plants in Europe.

Trait-based approaches are widely used in community ecology and invasion biology to unravel underlying mechanisms of vegetation dynamics. Although fundamental trade-offs between specific traits and invasibility are well described among terrestrial plants, little is known about their role and function in aquatic plant species. In this study, we examine the functional differences of aquatic alien and native plants stating that alien and native species differ in selected leaf traits. Our investigation is based on 60 taxa (21 alien and 39 native) collected from 22 freshwater units of Hungarian and Italian lowlands and highlands. Linear mixed models were used to investigate the effects of nativeness on four fundamental traits (leaf area, leaf dry matter content, specific leaf area, and leaf nitrogen content), while the influence of growth-form, altitude, and site were employed simultaneously. We found significantly higher values of leaf areas and significantly lower values of specific leaf areas for alien species if growth-form was included in the model as an additional predictor.We showed that the trait-based approach of autochthony can apply to aquatic environments similar to terrestrial ones, and leaf traits have relevance in explaining aquatic plant ecology whether traits are combined with growth-forms as a fixed factor. Our results confirm the importance of traits related to competitive ability in the process of aquatic plant invasions. Alien aquatic plants can be characterized as species producing soft leaves faster. We argue that the functional traits of alien aquatic plants are strongly growth-form dependent. Using the trait-based approach, we found reliable characteristics of aquatic plants related to species invasions, which might be used, for example, in conservation management.

Consistent avoidance of human disturbance over large geographical distances by a migratory bird.

Recent work on animal personalities has demonstrated that individuals may show consistent behaviour across situations and contexts. These studies were often carried out in one location and/or during short time intervals. Many animals, however, migrate and spend their life in several geographically distinct locations, and they may either adopt behaviours specific to the local environment or keep consistent behaviours over ecologically distinct locations. Long-distance migratory species offer excellent opportunities to test whether the animals maintain their personalities over large geographical scale, although the practical difficulties associated with these studies have hampered such tests. Here, we demonstrate for the first time consistency in disturbance tolerance behaviour in a long-distance migratory bird, using the common crane Grus grus as an ecological model species. Cranes that hatched in undisturbed habitats in Finland choose undisturbed migratory stop-over sites in Hungary, 1300-2000 km away from their breeding ground. This is remarkable, because these sites are not only separated by large distances, they also differ ecologically: the breeding sites are wooded bogs and subarctic tundra, whereas the migratory stop-over sites are temperate zone alkaline grasslands. The significance of our study goes beyond evolutionary biology and behavioural ecology: local effects on behaviour may carry over large distances, and this hitherto hidden implication of habitat selection needs to be incorporated into conservation planning.