Simultaneous GPS-tracking of parents reveals a similar parental investment within pairs, but no immediate co-adjustment on a trip-to-trip basis.

BACKGROUND: Parental care benefits the offspring, but comes at a cost for each parent, which in biparental species gives rise to a conflict between partners regarding the within-pair distribution of care. Pair members could avoid exploitation by efficiently keeping track of each other's efforts and coordinating their efforts. Parents may, therefore, space their presence at the nest, which could also allow for permanent protection of the offspring. Additionally, they may respond to their partner's previous investment by co-adjusting their efforts on a trip-to-trip basis, resulting in overall similar parental activities within pairs. METHODS: We investigated the coordination of parental care measured as nest attendance and foraging effort in the Lesser black-backed gull (Larus fuscus), a species with long nest bouts that performs extended foraging trips out of sight of their partner. This was achieved by GPS-tracking both pair members simultaneously during the entire chick rearing period. RESULTS: We found that the timing of foraging trips (and hence nest attendance) was coordinated within gull pairs, as individuals left the colony only after their partner had returned. Parents did not match their partner's investment by actively co-adjusting their foraging efforts on a trip-by-trip basis. Yet, pair members were similar in their temporal and energetic investments during chick rearing. CONCLUSION: Balanced investment levels over a longer time frame suggest that a coordination of effort may not require permanent co-adjustment of the levels of care on a trip-to-trip basis, but may instead rather take place at an earlier stage in the reproductive attempt, or over integrated longer time intervals. Identifying the drivers and underlying processes of coordination will be one of the next necessary steps to fully understand parental cooperation in long-lived species.

Breeding habitat loss reveals limited foraging flexibility and increases foraging effort in a colonial breeding seabird.

BACKGROUND: Habitat loss can force animals to relocate to new areas, where they would need to adjust to an unfamiliar resource landscape and find new breeding sites. Relocation may be costly and could compromise reproduction. METHODS: Here, we explored how the Lesser black-backed gull (Larus fuscus), a colonial breeding seabird species with a wide ecological niche, responds to the loss of its breeding habitat. We investigated how individuals adjusted their foraging behaviour after relocating to another colony due to breeding site destruction, and whether there were any reproductive consequences in the first years after relocation. To this end, we compared offspring growth between resident individuals and individuals that recently relocated to the same colony due to breeding habitat loss. Using GPS-tracking, we further investigated the foraging behaviour of resident individuals in both colonies, as well as that of relocated individuals, as enhanced foraging effort could represent a potential driver of reproductive costs. RESULTS: We found negative consequences of relocation for offspring development, which were apparent when brood demand was experimentally increased. Recently relocated gulls travelled further distances for foraging than residents, as they often visited more distant foraging sites used by residents breeding in their natal colony as well as new areas outside the home range of the residents in the colony where they settled. CONCLUSIONS: Our results imply that relocated individuals did not yet optimally adapt to the new food landscape, which was unexpected, given the social information on foraging locations that may have been available from resident neighbours in their new breeding colony. Even though the short-term reproductive costs were comparatively low, we show that generalist species, such as the Lesser black-backed gull, may be more vulnerable to habitat loss than expected. Long term studies are needed to investigate how long individuals are affected by their relocation in order to better assess potential population effects of (breeding) habitat loss.

Recently-adopted foraging strategies constrain early chick development in a coastal breeding gull.

Human-mediated food sources offer possibilities for novel foraging strategies by opportunistic species. Yet, relative costs and benefits of alternative foraging strategies vary with the abundance, accessibility, predictability and nutritional value of anthropogenic food sources. The extent to which such strategies may ultimately alter fitness, can have important consequences for long-term population dynamics. Here, we studied the relationships between parental diet and early development in free-ranging, cross-fostered chicks and in captive-held, hand-raised chicks of Lesser Black-backed Gulls (Larus fuscus) breeding along the Belgian coast. This traditionally marine and intertidal foraging species is now increasingly taking advantage of human activities by foraging on terrestrial food sources in agricultural and urban environments. In accordance with such behavior, the proportion of terrestrial food in the diet of free-ranging chicks ranged between 4% and 80%, and consistent stable isotope signatures between age classes indicated that this variation was mainly due to between-parent variation in feeding strategies. A stronger terrestrial food signature in free-ranging chicks corresponded with slower chick development. However, no consistent differences in chick development were found when contrasting terrestrial and marine diets were provided ad libitum to hand-raised chicks. Results of this study hence suggest that terrestrial diets may lower reproductive success due to limitations in food quantity, rather than quality. Recent foraging niche expansion toward terrestrial resources may thus constitute a suboptimal alternative strategy to marine foraging for breeding Lesser Black-backed Gulls during the chick-rearing period.

Time and energy costs of different foraging choices in an avian generalist species.

BACKGROUND: Animals can obtain a higher foraging yield by optimizing energy expenditure or minimizing time costs. In this study, we assessed how individual variation in the relative use of marine and terrestrial foraging habitats relates to differences in the energy and time investments of an avian generalistic feeder (the Lesser Black-backed Gull, Larus fuscus), and how this changes during the course of the chick-rearing period. METHODS: We analyzed 5 years of GPS tracking data collected at the colony of Zeebrugge (Belgium). Cost proxies for energy expenditure (overall dynamic body acceleration) and time costs (trip durations and time spent away from the colony), together with trip frequency, were analyzed against the relative use of the marine and terrestrial habitats. RESULTS: The marine habitat was most often used by males and outside weekends, when fisheries are active. Marine trips implied higher energetic costs and lower time investments. As chicks became older, terrestrial trips became more prevalent, and trip frequency reached a peak towards 20 days after hatching of the first egg. Over a full chick rearing period, energy costs varied widely between individuals, but no trends were found across the marine foraging gradient. Conversely, a higher use of marine foraging implied lower overall amounts of time spent away from the colony. CONCLUSIONS: Foraging habitat choice was related to overall time costs incurred by gulls, but not to energy costs. The effect of chick age on foraging habitat choice and effort may be driven by energy expenditure constraints on the amount of marine foraging that can be performed. If time is less constraining to them, Lesser Black-backed Gulls may meet the increasing chick demand for food by switching from high to low energy demanding foraging strategies.

Biodiversity effects on ecosystem functioning respond unimodally to environmental stress.

Understanding how biodiversity (B) affects ecosystem functioning (EF) is essential for assessing the consequences of ongoing biodiversity changes. An increasing number of studies, however, show that environmental conditions affect the shape of BEF relationships. Here, we first use a game-theoretic community model to reveal that a unimodal response of the BEF slope can be expected along environmental stress gradients, but also how the ecological mechanisms underlying this response may vary depending on how stress affects species interactions. Next, we analysed a global dataset of 44 experiments that crossed biodiversity with environmental conditions. Confirming our main model prediction, the effect of biodiversity on ecosystem functioning tends to be greater at intermediate levels of environmental stress, but varies among studies corresponding to differences in stress-effects on species interactions. Together, these results suggest that increases in stress from ongoing global environmental changes may amplify the consequences of biodiversity changes.

High-resolution GPS tracking reveals sex differences in migratory behaviour and stopover habitat use in the Lesser Black-backed Gull Larus fuscus.

Sex-, size- or age-dependent variation in migration strategies in birds is generally expected to reflect differences in competitive abilities. Theoretical and empirical studies thereby focus on differences in wintering areas, by which individuals may benefit from avoiding food competition during winter or ensuring an early return and access to prime nesting sites in spring. Here, we use GPS tracking to assess sex- and size-related variation in the spatial behaviour of adult Lesser Black-backed Gulls (Larus fuscus) throughout their annual cycle. We did not find sex- or size-dependent differences in wintering area or the timing of spring migration. Instead, sexual differences occurred prior to, and during, autumn migration, when females strongly focussed on agricultural areas. Females exhibited a more protracted autumn migration strategy, hence spent more time on stopover sites and arrived 15 days later at their wintering areas, than males. This shift in habitat use and protracted autumn migration coincided with the timing of moult, which overlaps with chick rearing and migration. Our results suggest that this overlap between energy-demanding activities may lead females to perform a more prolonged autumn migration, which results in spatiotemporal differences in foraging habitat use between the sexes.

Shifts of community composition and population density substantially affect ecosystem function despite invariant richness.

There has been considerable focus on the impacts of environmental change on ecosystem function arising from changes in species richness. However, environmental change may affect ecosystem function without affecting richness, most notably by affecting population densities and community composition. Using a theoretical model, we find that, despite invariant richness, (1) small environmental effects may already lead to a collapse of function; (2) competitive strength may be a less important determinant of ecosystem function change than the selectivity of the environmental change driver and (3) effects on ecosystem function increase when effects on composition are larger. We also present a complementary statistical analysis of 13 data sets of phytoplankton and periphyton communities exposed to chemical stressors and show that effects on primary production under invariant richness ranged from -75% to +10%. We conclude that environmental protection goals relying on measures of richness could underestimate ecological impacts of environmental change.

Mixture toxicity in the marine environment: Model development and evidence for synergism at environmental concentrations.

Little is known about the effect of metal mixtures on marine organisms, especially after exposure to environmentally realistic concentrations. This information is, however, required to evaluate the need to include mixtures in future environmental risk assessment procedures. We assessed the effect of copper (Cu)-Nickel (Ni) binary mixtures on Mytilus edulis larval development using a full factorial design that included environmentally relevant metal concentrations and ratios. The reproducibility of the results was assessed by repeating this experiment 5 times. The observed mixture effects were compared with the effects predicted with the concentration addition model. Deviations from the concentration addition model were estimated using a Markov chain Monte-Carlo algorithm. This enabled the accurate estimation of the deviations and their uncertainty. The results demonstrated reproducibly that the type of interaction-synergism or antagonism-mainly depended on the Ni concentration. Antagonism was observed at high Ni concentrations, whereas synergism occurred at Ni concentrations as low as 4.9 µg Ni/L. This low (and realistic) Ni concentration was 1% of the median effective concentration (EC50) of Ni or 57% of the Ni predicted-no-effect concentration (PNEC) in the European Union environmental risk assessment. It is concluded that results from mixture studies should not be extrapolated to concentrations or ratios other than those investigated and that significant mixture interactions can occur at environmentally realistic concentrations. This should be accounted for in (marine) environmental risk assessment of metals. Environ Toxicol Chem 2017;36:3471-3479. © 2017 SETAC.

The Consequences of Nonrandomness in Species-Sensitivity in Relation to Functional Traits for Ecosystem-Level Effects of Chemicals.

Estimating ecosystem-level effects from single-species bioassays is a major challenge in environmental risk assessment. Most extrapolation procedures are based on the implicit assumption that species sensitivities are random with regard to their functional traits. Here, we explore how nonrandomness in species sensitivities affects how species-level and ecosystem level effects of chemical exposure correspond. The effect of a correlation between the trait value under control conditions and the sensitivity of the trait to chemical stress is studied for two traits (per capita growth rate and monoculture yield) under constant and temporary exposure. Theoretical model predictions are thereby validated against a 3-week microcosm experiment, in which eight marine diatoms systems with different correlations between trait values and sensitivities were temporary (1 week) or constantly (3 weeks) exposed to two concentrations of the herbicide atrazine (100 and 250 µg L-1). Negative correlations increased the reduction in ecosystem functioning (productivity) by atrazine for both traits. However, correlations in the per capita growth rate affected productivity only shortly following changes in environmental conditions, whereas correlations in the monoculture yield affected productivity throughout exposure. Correlations between species sensitivities and functional trait values can thus help to identify when ecosystem-level effects are likely to exceed species-level effects.

Biodiversity increases functional and compositional resistance, but decreases resilience in phytoplankton communities.

There is now ample evidence that biodiversity stabilizes aggregated ecosystem functions, such as primary production, in changing environments. In primary producer systems, this stabilizing effect is found to be driven by higher functional resistance (i.e., reduced changes in functions by environmental changes) rather than through higher functional resilience (i.e., rapid recovery following environmental changes) in more diverse systems. The stability of aggregated ecosystem functions directly depends on changes in species composition and by consequence their functional contributions to ecosystem functions. Still, it remains only theoretically explored how biodiversity can stabilize ecosystem functions by affecting compositional stability. Here, we demonstrate how biodiversity effects on compositional stability drive biodiversity effects on functional stability in diatom communities. In a microcosm experiment, we exposed 39 communities of five different levels of species richness (1, 2, 4, 6, and 8 species) to three concentrations of a chemical stressor (0, 25, and 250 µg/L atrazine) for four weeks, after which all communities were transferred to atrazine-free medium for three more weeks. Biodiversity simultaneously increased, increasing functional and compositional resistance, but decreased functional and compositional resilience. These results confirm the theoretically proposed link between biodiversity effects on functional and compositional stability in primary producer systems, and provide a mechanistic underpinning for observed biodiversity-stability relationships. Finally, we discuss how higher compositional stability can be expected to become increasingly important in stabilizing ecosystem functions under field conditions when multiple environmental stressors fluctuate simultaneously.

Per capita interactions and stress tolerance drive stress-induced changes in biodiversity effects on ecosystem functions.

Environmental stress changes the relationship between biodiversity and ecosystem functions, but the underlying mechanisms are poorly understood. Because species interactions shape biodiversity-ecosystem functioning relationships, changes in per capita interactions under stress (as predicted by the stress gradient hypothesis) can be an important driver of stress-induced changes in these relationships. To test this hypothesis, we measure productivity in microalgae communities along a diversity and herbicide gradient. On the basis of additive partitioning and a mechanistic community model, we demonstrate that changes in per capita interactions do not explain effects of herbicide stress on the biodiversity-productivity relationship. Instead, assuming that the per capita interactions remain unaffected by stress, causing species densities to only change through differences in stress tolerance, suffices to predict the stress-induced changes in the biodiversity-productivity relationship and community composition. We discuss how our findings set the stage for developing theory on how environmental stress changes biodiversity effects on ecosystem functions.