The effect of climate change on avian offspring production: A global meta-analysis.

Climate change affects timing of reproduction in many bird species, but few studies have investigated its influence on annual reproductive output. Here, we assess changes in the annual production of young by female breeders in 201 populations of 104 bird species (N = 745,962 clutches) covering all continents between 1970 and 2019. Overall, average offspring production has declined in recent decades, but considerable differences were found among species and populations. A total of 56.7% of populations showed a declining trend in offspring production (significant in 17.4%), whereas 43.3% exhibited an increase (significant in 10.4%). The results show that climatic changes affect offspring production through compounded effects on ecological and life history traits of species. Migratory and larger-bodied species experienced reduced offspring production with increasing temperatures during the chick-rearing period, whereas smaller-bodied, sedentary species tended to produce more offspring. Likewise, multi-brooded species showed increased breeding success with increasing temperatures, whereas rising temperatures were unrelated to reproductive success in single-brooded species. Our study suggests that rapid declines in size of bird populations reported by many studies from different parts of the world are driven only to a small degree by changes in the production of young.

Increasingly cautious sampling, not the black colouration of unpalatable prey, is used by fish in avoidance learning.

The efficiency of aposematic colouration of prey is based on the innate bias or facilitation of avoidance learning of predators. In many toxic bufonids, larvae are uniformly black, which is considered a warning signal. We compared fish predation on normal (black) and 'transient albino' (greyish) common toad Bufo bufo tadpoles that did not differ in toxicity or activity. In a two-stage experiment, each fish was presented with tadpoles of one colour in the first trial and the other colour in a subsequent trial. While tadpoles sampled by fish were typically not ingested, some died from injuries. The attack rate did not differ between tadpole phenotypes nor trials, irrespective of which phenotype was the first exposed to the fish. However, during the second trial, the sampled tadpoles, independent of colouration, were mouthed by fish for shorter periods and tadpole mortality decreased. The duration of mouthing also declined with an increasing number of attacks during subsequent trials. We conclude that in single-species prey populations, black tadpole colouration is not a warning signal as it does not accelerate predator learning about prey unprofitability. Our results indicate that with growing experience, predators sample potentially toxic prey more cautiously. This may explain why natural selection does not eliminate aposematic morphs even if predators continuously sample conspicuous prey.

A matter of proportion? Associational effects in larval anuran communities under fish predation.

In Batesian mimicry, a species lacking defences against predators benefits from mimicking the aposematic signal of a defended species, while the model may incur the costs of reduced defensive efficacy. Similar reciprocal indirect effects may emerge even when the signal is not mimicked; termed associational effects, such interactions are well known in plants sharing herbivores but have received little attention in animal studies. We investigated associational interactions in a system where unequally defended prey (chemically defended Bufo bufo and undefended Rana temporaria tadpoles), sharing general morphology but not an aposematic signal, were exposed to predation by the carp Cyprinus carpio along a gradient of relative prey abundance. In the absence of fish, the assemblage composition had no effect on the survival of Rana, while that of Bufo decreased with increasing abundance of Rana. Fish reduced the survival of tadpoles from both species. However, increased relative abundance of Bufo in the community led to enhanced survival in both Bufo and Rana. Increasing relative proportions of heterospecifics reduced metamorph mass only in Bufo, indicating greater sensitivity to interspecific competition compared to Rana; the effect was reduced in the presence of fish. Our results show that undefended non-mimetic prey enjoy reduced predation with increasing relative abundance of chemically defended prey, which in turn suffer greater mortality with an increasing proportion of the undefended species. Associational resistance/susceptibility, driven by current assemblage composition, not by selection for resemblance, can shape the dynamics of mixed communities of defended and undefended prey in the absence of mimicry.

Total Non-Consumptive Effects of Fish on Pelobates fuscus and Hyla orientalis Tadpoles in Pond Enclosure Experiments.

Most research on non-consumptive predator effects on amphibian larvae has been conducted in laboratory or mesocosm designs. Here, Pelobates fuscus and Hyla orientalis tadpoles were separately exposed to non-lethal (free-moving, but with tied mouthparts) common carp Cyprinus carpio for one week in enclosures placed in a pond densely stocked with fish. Tadpoles exposed to nonlethal fish did not differ in mortality, body mass, or, except for deeper tail fin depth in P. fuscus, morphological plasticity from controls kept in a fishless pond. Hyla orientalis tadpoles recovered from the fish treatment were subsequently enclosed until metamorphosis in either the pond with fish or the fishless pond. Metamorphs from the pond containing fish were heavier, and did not differ in survivorship or development time from their counterparts initially kept in the fish treatment and then transferred to the fishless pond or from controls kept the entire time under fish-free conditions. The lack of apparent metamorphic costs is consistent with previous experiments on anuran larvae, but the morphological defenses induced (or their absence) are not. In the fish-dominated pond, carp indirectly affected tadpole developmental responses by generating turbidity, through adverse impacts on submerged vegetation and predatory insects, and by increasing food resource (unicellular algae) levels. While the present study does not question the validity of laboratory and mesocosm experiments on the costs of non-consumptive predator effects on amphibian larvae, their outcomes cannot easily be extrapolated to ecologically complex natural habitats.

Phylogenetic Diversity of Ossification Patterns in the Avian Vertebral Column: A Review and New Data from the Domestic Pigeon and Two Species of Grebes.

Despite many decades of studies, our knowledge of skeletal development in birds is limited in many aspects. One of them is the development of the vertebral column. For many years it was widely believed that the column ossifies anteroposteriorly. However, later studies indicated that such a pattern is not universal in birds and in many groups the ossification starts in the thoracic rather than cervical region. Recent analyses suggest that two loci, located in the cervical and thoracic vertebrae, were ancestrally present in birds. However, the data on skeletal development are very scarce in the Neoaves, a clade that includes approximately 95% of extant species. We review the available information about the vertebral column development in birds and describe the ossification pattern in three neoavians, the domestic pigeon (Columba livia domestica), the great crested grebe (Podiceps cristatus) and the red-necked grebe (Podiceps grisegena). In P. cristatus, the vertebral column starts ossifying in the thoracic region. The second locus is present in the cervical vertebrae. In the pigeon, the cervical vertebrae ossify before the thoracics, but both the thoracic and cervical loci are present. Our ancestral state reconstructions confirm that both these loci were ancestrally present in birds, but the thoracic locus was later lost in psittacopasserans and at least some galloanserans.

Consequences of the size structure of fish populations for their effects on a generalist avian predator.

Size-structured interspecific interactions can shift between predation and competition, depending on ontogenetic changes in size relationships. I examined the effects of common carp (Cyprinus carpio), an omnivorous fish, on the reproductive success of the red-necked grebe (Podiceps grisegena), an avian gape-limited predator, along a fish size gradient created by stocking distinct age-cohorts in seminatural ponds. Young-of-the-year (0+) carp were an essential food source for young grebes. Only adult birds were able to consume 1-year-old (1+) fish, while 2-year-old (2+) fish attained a size refuge from grebes. Amphibian larvae were the principal alternative prey to fish, followed by macroinvertebrates, but the abundance of both dramatically decreased along the carp size gradient. Fledging success was 2.8 times greater in ponds with 0+ versus 1+ carp; in ponds with 1+ carp, chicks received on average 2.6-3 times less prey biomass from their parents, and over 1/3 of broods suffered total failure. Breeding birds avoided settling on 2+ ponds. These results show that changes in prey fish size structure can account for shifts from positive trophic effects on the avian predator to a negative impact on the predator's alternative resources. However, competition did not fully explain the decrease in grebe food resources in the presence of large fish, as carp and grebes overlapped little in diet. In experimental cages, 1+ carp totally eliminated young larvae of amphibians palatable to fish. In field conditions, breeding adults of palatable taxa avoided ponds with 1+ and older carp. Non-trophic interactions such as habitat selection by amphibians or macroinvertebrates to avoid large fish may provide an indirect mechanism strengthening the adverse bottom-up effects of fish on birds.

Win-stay/lose-switch, prospecting-based settlement strategy may not be adaptive under rapid environmental change.

Understanding animal responses to environmental change is crucial for management of ecological traps. Between-year habitat selection was investigated in red-necked grebes (Podiceps grisegena) breeding on semi-natural fish ponds, where differential stocking of fish created contrasting yet poorly predictable brood-stage food availabilities. Grebes lured to low-quality ponds were more likely to shift territories than birds nesting on high-quality ponds, and tended to move to ponds whose habitat quality had been high in the previous year, irrespective of the current quality of the new and old territories. The territory switchers typically visited their future breeding ponds during or immediately after the brood-rearing period. However, owing to rotation of fish stocks, the habitat quality of many ponds changed in the following year, and then switchers from low-quality ponds and stayers on previously high-quality ponds were ecologically trapped. Thus, although breeders were making an informed choice, their settlement decisions, based on the win-stay/lose-switch rule and prospecting a year in advance, were inappropriate in conditions of year-to-year habitat fluctuations. Effective adaptation to rapid environmental change may necessitate both learning to correctly evaluate uncertain environmental cues and abandonment of previously adaptive decision-making algorithms (here prioritizing past-year information and assuming temporal autocorrelation of habitat quality).

Habitat selection by breeding waterbirds at ponds with size-structured fish populations.

Fish may significantly affect habitat use by birds, either as their prey or as competitors. Fish communities are often distinctly size-structured, but the consequences for waterbird assemblages remain poorly understood. We examined the effects of size structure of common carp (Cyprinus carpio) cohorts together with other biotic and abiotic pond characteristics on the distribution of breeding waterbirds in a seminatural system of monocultured ponds, where three fish age classes were separately stocked. Fish age corresponded to a distinct fish size gradient. Fish age and total biomass, macroinvertebrate and amphibian abundance, and emergent vegetation best explained the differences in bird density between ponds. Abundance of animal prey other than fish (aquatic macroinvertebrates and larval amphibians) decreased with increasing carp age in the ponds. Densities of ducks and smaller grebes were strongly negatively associated with fish age/size gradient. The largest of the grebes, the piscivorous great crested grebe (Podiceps cristatus), was the only species that preferred ponds with medium-sized fish and was positively associated with total fish biomass. Habitat selection by bitterns and most rallids was instead strongly influenced by the relative amount of emergent vegetation cover in the ponds. Our results show that fish size structure may be an important cue for breeding habitat choice and a factor affording an opportunity for niche diversification in avian communities.

Numbers, neighbors, and hungry predators: What makes chemically defended aposematic prey susceptible to predation?

Many chemically defended aposematic species are characterized by relatively low toxin levels, which enables predators to include them in their diets under certain circumstances. Knowledge of the conditions governing the survival of such prey animals-especially in the context of the co-occurrence of similar but undefended prey, which may result in mimicry-like interactions-is crucial for understanding the initial evolution of aposematism. In a one-month outdoor experiment using fish (the common carp Cyprinus carpio) as predators, we examined the survival of moderately defended aposematic tadpole prey (the European common toad Bufo bufo) with varying absolute densities in single-species prey systems or varying relative densities in two-species prey systems containing morphologically similar but undefended prey (the European common frog Rana temporaria). The density effects were investigated in conjunction with the hunger levels of the predator, which were manipulated by means of the addition of alternative (nontadpole) food. The survival of the B. bufo tadpoles was promoted by increasing their absolute density in the single-species prey systems, increasing their relative density in the two-species prey systems, and providing ample alternative food for the predator. Hungry predators eliminated all R. temporaria individuals regardless of their proportion in the prey community; in treatments with ample alternative food, high relative B. bufo density supported R. temporaria survival. The results demonstrated that moderately defended prey did benefit from high population densities (both absolute and relative), even under long-term predation pressure. However, the physiological state of the predator was a crucial factor in the survival of moderately defended prey. While the availability of alternative prey in general should promote the spread and maintenance of aposematism, the results indicated that the resemblance between the co-occurring defended and undefended prey may impose mortality costs on the defended model species, even in the absence of actual mimicry.

A novel strategy to escape a poor habitat: red-necked grebes transfer flightless young to other ponds.

Animals confronted with the threat of the death of their offspring may exhibit unusual and risk-prone behaviours. Grebes (Podicipediformes) are water birds which cannot effectively walk, thus unfledged young are assumed to be unable to depart from their natal ponds by land. We provide evidence that red-necked grebes Podiceps grisegena, breeding on ponds with scarce food resources, transferred their flightless young (2-4 weeks old) to other, unconnected ponds by land or air. Although a large proportion of breeding grebes in the study area nested on food-poor fish ponds acting as ecological traps, where they suffered significant brood losses, brood movements to new ponds accounted for only 3.3% of such breeding attempts. The infrequency of this strategy may be explained by the lack of suitable territories in close proximity and the high risk of predation or fatal injury. The means of chick transfer remains unclear; the chicks may have followed or been carried by parents shuffling across the pond levees; alternatively, parents may have carried the young on their backs in flight. Our findings indicate that red-necked grebes assess the current level of resources available for chicks and may adopt novel and risky strategies to escape total brood failure.