Northward range expansion in spring-staging barnacle geese is a response to climate change and population growth, mediated by individual experience

The study was funded by the Norwegian Environment Agency, the Research Council of Norway (the projects ‘LANDRING 134716/720’, ‘AGRIGOOSE 165836’, ‘MIGRAPOP 204342’), the European Union (the project FRAGILE EVK2‐2001‐00235), the County Governor of Nordland, the Wildfowl and Wetlands Trust, the Fram Centre in Tromsø and a grant from the Netherlands Organization for Scientific Research awarded to TO (ref 019.172EN.011).All long-distance migrants must cope with changing environments, but species differ greatly in how they do so. In some species, individuals might be able to adjust by learning from individual experiences and by copying others. This could greatly speed up the process of adjustment, but evidence from the wild is scarce. Here, we investigated the processes by which a rapidly growing population of barnacle geese (Branta leucopsis) responded to strong environmental changes on spring-staging areas in Norway. One area, Helgeland, has been the traditional site. Since the mid-1990s, an increasing number of geese stage in another area 250 km further north, Vesterålen. We collected data on goose numbers and weather conditions from 1975 to 2017 to explore the extent to which the increase in population size and a warmer climate contributed to this change in staging area use. During the study period, the estimated onset of grass growth advanced on average by 0.54 days/year in each of the two areas. The total production of digestible biomass for barnacle geese during the staging period increased in Vesterålen but remained stable in Helgeland. The goose population has doubled in size during the past 25 years, with most of the growth being accommodated in Vesterålen. The observations suggest that this dramatic increase would not have happened without higher temperatures in Vesterålen. Records of individually marked geese indicate that from the initial years of colonization onwards, especially young geese tended to switch to Vesterålen, thereby predominating in the flocks at Vesterålen. Older birds had a lower probability of switching to Vesterålen, but over the years, the probability increased for all ages. Our findings suggest that barnacle geese integrate socially learned behaviour with adjustments to individual experiences, allowing the population to respond rapidly and accurately to global change.Publisher PDFPeer reviewe

Contrasting consequences of climate change for migratory geese:Predation, density dependence and carryover effects offset benefits of high-arctic warming

Climate change is most rapid in the Arctic, posing both benefits and challenges for migratory herbivores. However, population-dynamic responses to climate change are generally difficult to predict, due to concurrent changes in other trophic levels. Migratory species are also exposed to contrasting climate trends and density regimes over the annual cycle. Thus, determining how climate change impacts their population dynamics requires an understanding of how weather directly or indirectly (through trophic interactions and carryover effects) affects reproduction and survival across migratory stages, while accounting for density dependence. Here, we analyse the overall implications of climate change for a local non-hunted population of high-arctic Svalbard barnacle geese, Branta leucopsis, using 28 years of individual-based data. By identifying the main drivers of reproductive stages (egg production, hatching and fledging) and age-specific survival rates, we quantify their impact on population growth. Recent climate change in Svalbard enhanced egg production and hatching success through positive effects of advanced spring onset (snow melt) and warmer summers (i.e. earlier vegetation green-up) respectively. Contrastingly, there was a strong temporal decline in fledging probability due to increased local abundance of the Arctic fox, the main predator. While weather during the non-breeding season influenced geese through a positive effect of temperature (UK wintering grounds) on adult survival and a positive carryover effect of rainfall (spring stopover site in Norway) on egg production, these covariates showed no temporal trends. However, density-dependent effects occurred throughout the annual cycle, and the steadily increasing total flyway population size caused negative trends in overwinter survival and carryover effects on egg production. The combination of density-dependent processes and direct and indirect climate change effects across life history stages appeared to stabilize local population size. Our study emphasizes the need for holistic approaches when studying population-dynamic responses to global change in migratory species.</p

Predation Danger Can Explain Changes in Timing of Migration: The Case of the Barnacle Goose

Understanding stopover decisions of long-distance migratory birds is crucial for conservation and management of these species along their migratory flyway. Recently, an increasing number of Barnacle geese breeding in the Russian Arctic have delayed their departure from their wintering site in the Netherlands by approximately one month and have reduced their staging duration at stopover sites in the Baltic accordingly. Consequently, this extended stay increases agricultural damage in the Netherlands. Using a dynamic state variable approach we explored three hypotheses about the underlying causes of these changes in migratory behavior, possibly related to changes in (i) onset of spring, (ii) potential intake rates and (iii) predation danger at wintering and stopover sites. Our simulations showed that the observed advance in onset of spring contradicts the observed delay of departure, whereas both increased predation danger and decreased intake rates in the Baltic can explain the delay. Decreased intake rates are expected as a result of increased competition for food in the growing Barnacle goose population. However, the effect of predation danger in the model was particularly strong, and we hypothesize that Barnacle geese avoid Baltic stopover sites as a response to the rapidly increasing number of avian predators in the area. Therefore, danger should be considered as an important factor influencing Barnacle goose migratory behavior, and receive more attention in empirical studies

Above- and below-ground vertebrate herbivory may each favour a different subordinate species in an aquatic plant community

At least two distinct trade-offs are thought to facilitate higher diversity in productive plant communities under herbivory. Higher investment in defence and enhanced colonization potential may both correlate with decreased competitive ability in plants. Herbivory may thus promote coexistence of plant species exhibiting divergent life history strategies. How different seasonally tied herbivore assemblages simultaneously affect plant community composition and diversity is, however, largely unknown. Two contrasting types of herbivory can be distinguished in the aquatic vegetation of the shallow lake Lauwersmeer. In summer, predominantly above-ground tissues are eaten, whereas in winter, waterfowl forage on below-ground plant propagules. In a 4-year exclosure study we experimentally separated above-ground herbivory by waterfowl and large fish in summer from below-ground herbivory by Bewick’s swans in winter. We measured the individual and combined effects of both herbivory periods on the composition of the three-species aquatic plant community. Herbivory effect sizes varied considerably from year to year. In 2 years herbivore exclusion in summer reinforced dominance of Potamogeton pectinatus with a concomitant decrease in Potamogeton pusillus, whereas no strong, unequivocal effect was observed in the other 2 years. Winter exclusion, on the other hand, had a negative effect on Zannichellia palustris, but the effect size differed considerably between years. We suggest that the colonization ability of Z. palustris may have enabled this species to be more abundant after reduction of P. pectinatus tuber densities by swans. Evenness decreased due to herbivore exclusion in summer. We conclude that seasonally tied above- and below-ground herbivory may each stimulate different components of a macrophyte community as they each favoured a different subordinate plant species

Defensive responses by a social caterpillar are tailored to different predators and change with larval instar and group size

Gregariousness in animals is widely accepted as a behavioral adaptation for protection from predation. However, predation risk and the effectiveness of a prey’s defense can be a function of several other factors, including predator species and prey size or age. The objective of this study was to determine if the gregarious habit of Malacosoma disstria caterpillars is advantageous against invertebrate natural enemies, and whether it is through dilution or cooperative defenses. We also examined the effects of larval growth and group size on the rate and success of attacks. Caterpillars of M. disstria responded with predator-specific behaviors, which led to increased survival. Evasive behaviors were used against stinkbugs, while thrashing by fourth instar caterpillars and holding on to the silk mat by second instar caterpillars was most efficient against spider attacks. Collective head flicking and biting by groups of both second and fourth instar caterpillars were observed when attacked by parasitoids. Increased larval size decreased the average number of attacks by spiders but increased the number of attacks by both stinkbugs and parasitoids. However, increased body size decreased the success rate of attacks by all three natural enemies and increased handling time for both predators. Larger group sizes did not influence the number of attacks from predators but increased the number of attacks and the number of successful attacks from parasitoids. In all cases, individual risk was lower in larger groups. Caterpillars showed collective defenses against parasitoids but not against the walking predators. These results show that caterpillars use different tactics against different natural enemies. Overall, these tactics are both more diverse and more effective in fourth instar than in second instar caterpillars, confirming that growth reduces predation risk. We also show that grouping benefits caterpillars through dilution of risk, and, in the case of parasitoids, through group defenses. The decreased tendency to aggregate in the last larval instar may therefore be linked to decreasing predation risk

Influence of Snowmelt Timing on the Diet Quality of Pyrenean Rock Ptarmigan (Lagopus muta pyrenaica): Implications for Reproductive Success

The Pyrenean rock ptarmigan (Lagopus muta pyrenaica) is the southernmost subspecies of the species in Europe and is considered threatened as a consequence of changes in landscape, human pressure, climate change, and low genetic diversity. Previous studies have shown a relationship between the date of snowmelt and reproductive success in the Pyrenean ptarmigan. It is well established that birds laying early in the breeding season have higher reproductive success, but the specific mechanism for this relationship is debated. We present an explicative model of the relationship between snowmelt date and breeding success mediated by food quality for grouse in alpine environments. From microhistological analyses of 121 faecal samples collected during three years in the Canigou Massif (Eastern Pyrenees), and the assessment of the chemical composition of the main dietary components, we estimated the potential quality of individual diets. Potential dietary quality was correlated with free-urate faecal N, a proxy of the digestible protein content ingested by ptarmigan, and both were correlated with phenological stage of consumed plants, which in turn depends on snowmelt date. Our findings suggest that the average snowmelt date is subject to a strong interannual variability influencing laying date. In years of early snowmelt, hens benefit from a longer period of high quality food resources potentially leading to a higher breeding success. On the contrary, in years of late snowmelt, hens begin their breeding period in poorer nutrient condition because the peaks of protein content of their main food items are delayed with respect to laying date, hence reducing breeding performance. We discuss the possible mismatch between breeding and snowmelt timing

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 repro- ductive 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