A circumpolar study unveils a positive non-linear effect of temperature on arctic arthropod availability that may reduce the risk of warming-induced trophic mismatch for breeding shorebirds.

Seasonally abundant arthropods are a crucial food source for many migratory birds that breed in the Arctic. In cold environments, the growth and emergence of arthropods are particularly tied to temperature. Thus, the phenology of arthropods is anticipated to undergo a rapid change in response to a warming climate, potentially leading to a trophic mismatch between migratory insectivorous birds and their prey. Using data from 19 sites spanning a wide temperature gradient from the Subarctic to the High Arctic, we investigated the effects of temperature on the phenology and biomass of arthropods available to shorebirds during their short breeding season at high latitudes. We hypothesized that prolonged exposure to warmer summer temperatures would generate earlier peaks in arthropod biomass, as well as higher peak and seasonal biomass. Across the temperature gradient encompassed by our study sites (>10°C in average summer temperatures), we found a 3-day shift in average peak date for every increment of 80 cumulative thawing degree-days. Interestingly, we found a linear relationship between temperature and arthropod biomass only below temperature thresholds. Higher temperatures were associated with higher peak and seasonal biomass below 106 and 177 cumulative thawing degree-days, respectively, between June 5 and July 15. Beyond these thresholds, no relationship was observed between temperature and arthropod biomass. Our results suggest that prolonged exposure to elevated temperatures can positively influence prey availability for some arctic birds. This positive effect could, in part, stem from changes in arthropod assemblages and may reduce the risk of trophic mismatch.

Individual quality overwrites carry-over effects across the annual cycle of a long-distance migrant.

In seasonal environments, the fitness of animals depends upon the successful integration of life-history stages throughout their annual cycle. Failing to do so can lead to negative carry-over effects where individuals are transitioning into the next season in different states, consequently affecting their future performance. However, carry-over effects can be masked by individual quality when individuals vary in their efficiency at acquiring resources year after year (i.e. 'quality'), leading to cross-seasonal consistency in individual performance. Here we investigated the relative importance of carry-over effects and individual quality in determining cross-seasonal interactions and consequences for breeding success over the full annual cycle of a migratory seabird (black-legged kittiwake Rissa tridactyla). We monitored the reproduction and annual movement of kittiwakes over 13 years using geolocators to estimate their breeding success, distribution and winter energy expenditure. We combined this with an experimental approach (clutch removal experiment, 2 years) to manipulate the reproductive effort irrespective of individual quality. Piecewise path analyses showed that successful breeders reproduced earlier and were more likely to breed successfully again the following year. This positive interaction among consecutive breeding stages disappeared after controlling for individual quality, suggesting that quality was dominant in determining seasonal interactions. Moreover, controlling experimentally for individual quality revealed underlying carry-over effects that were otherwise masked by quality, with breeding costs paid in higher energy expenditure and delayed onset of reproduction. We highlight the need to combine an experimental approach along with long-term data while assessing apparent carry-over effects in wild animals, and their potential impact on fitness and population demography.

Winter Tracking Data Suggest that Migratory Seabirds Transport Per- and Polyfluoroalkyl Substances to Their Arctic Nesting Site.

Seabirds are often considered sentinel species of marine ecosystems, and their blood and eggs utilized to monitor local environmental contaminations. Most seabirds breeding in the Arctic are migratory and thus are exposed to geographically distinct sources of contamination throughout the year, including per- and polyfluoroalkyl substances (PFAS). Despite the abundance and high toxicity of PFAS, little is known about whether blood concentrations at breeding sites reliably reflect local contamination or exposure in distant wintering areas. We tested this by combining movement tracking data and PFAS analysis (nine compounds) from the blood of prelaying black-legged kittiwakes (Rissa tridactyla) nesting in Arctic Norway (Svalbard). PFAS burden before egg laying varied with the latitude of the wintering area and was negatively associated with time upon return of individuals at the Arctic nesting site. Kittiwakes (n = 64) wintering farther south carried lighter burdens of shorter-chain perfluoroalkyl carboxylates (PFCAs, C9-C12) and heavier burdens of longer chain PFCAs (C13-C14) and perfluorooctanesulfonic acid compared to those wintering farther north. Thus, blood concentrations prior to egg laying still reflected the uptake during the previous wintering stage, suggesting that migratory seabirds can act as biovectors of PFAS to Arctic nesting sites.

Predator home range size mediates indirect interactions between prey species in an arctic vertebrate community.

Indirect interactions are widespread among prey species that share a common predator, but the underlying mechanisms driving these interactions are often unclear, and our ability to predict their outcome is limited. Changes in behavioural traits that impact predator space use could be a key proximal mechanism mediating indirect interactions, but there is little empirical evidence of the causes and consequences of such behavioural-numerical response in multispecies systems. Here, we investigate the complex ecological relationships between seven prey species sharing a common predator. We used a path analysis approach on a comprehensive 9-year data set simultaneously tracking predator space use, prey densities and prey mortality rate on key species of a simplified Arctic food web. We show that high availability of a clumped and spatially predictable prey (goose eggs) leads to a twofold reduction in predator (arctic fox) home range size, which increases local predator density and strongly decreases nest survival of an incidental prey (American golden plover). On the contrary, a scattered cyclic prey with potentially lower spatial predictability (lemming) had a weaker effect on fox space use and an overall positive impact on the survival of incidental prey. These contrasting effects underline the importance of studying behavioural responses of predators in multiprey systems and to explicitly integrate behavioural-numerical responses in multispecies predator-prey models.

Do foraging ecology and contaminants interactively predict parenting hormone levels in common eider?

Global climate change is causing abiotic shifts such as higher air and ocean temperatures, and disappearing sea ice in Arctic ecosystems. These changes influence Arctic-breeding seabird foraging ecology by altering prey availability and selection, affecting individual body condition, reproductive success, and exposure to contaminants such as mercury (Hg). The cumulative effects of alterations to foraging ecology and Hg exposure may interactively alter the secretion of key reproductive hormones such as prolactin (PRL), important for parental attachment to eggs and offspring and overall reproductive success. However, more research is needed to investigate the relationships between these potential links. Using data collected from 106 incubating female common eiders (Somateria mollissima) at six Arctic and sub-Arctic colonies, we examined whether the relationship between individual foraging ecology (assessed using δ13C, δ15N) and total Hg (THg) exposure predicted PRL levels. We found a significant, complex interaction between δ13C, δ15N and THg on PRL, suggesting that individuals cumulatively foraging at lower trophic levels, in phytoplankton-dominant environments, and with the highest THg levels had the most constant significant relationship PRL levels. Cumulatively, these three interactive variables resulted in lowered PRL. Overall, results demonstrate the potential downstream and cumulative implications of environmentally induced changes in foraging ecology, in combination with THg exposure, on hormones known to influence reproductive success in seabirds. These findings are notable in the context of continuing environmental and food web changes in Arctic systems, which may make seabird populations more susceptible to ongoing stressors.

COVID19-induced reduction in human disturbance enhances fattening of an overabundant goose species.

Overabundant species can have major impacts on their habitat and induce trophic cascades within ecosystems. In North America, the overabundant greater snow goose (Anser caerulescens atlanticus) has been successfully controlled through special spring hunting regulations since 1999. Hunting is a source of mortality but also of disturbance, which affects the behavior and nutrient storage dynamics of staging snow geese. In 2020, the lockdown imposed by the COVID19 pandemic reduced hunting activity during their migratory stopover in Québec by at least 31%. This provided a unique opportunity to assess the effects of a sudden reduction in hunting disturbance on geese. We used long-term data on body mass combined with movement data from GPS-tracked birds in 2019 and 2020 to assess the effects of the 2020 lockdown on the spring body condition and behavior of greater snow geese. Body condition was higher in 2020 than in all years since the inception of spring hunting in 1999, except for 2019. However, in 2020 geese reached maximal body condition earlier during the staging period than in any other year and reduced by half time spent feeding in highly profitable but risky agricultural habitat in late spring compared to 2019. Although our study was not designed to evaluate the effects of the lockdown, the associated reduction in disturbance in 2020 supports the hypothesis that hunting-related disturbance negatively affects foraging efficiency and body condition in geese. Since spring body condition is related to subsequent breeding success, the lockdown could increase productivity in this overabundant population.

Seasonal food webs with migrations: multi-season models reveal indirect species interactions in the Canadian Arctic tundra.

Models incorporating seasonality are necessary to fully assess the impact of global warming on Arctic communities. Seasonal migrations are a key component of Arctic food webs that still elude current theories predicting a single community equilibrium. We develop a multi-season model of predator-prey dynamics using a hybrid dynamical systems framework applied to a simplified tundra food web (lemming-fox-goose-owl). Hybrid systems models can accommodate multiple equilibria, which is a basic requirement for modelling food webs whose topology changes with season. We demonstrate that our model can generate multi-annual cycling in lemming dynamics, solely from a combined effect of seasonality and state-dependent behaviour. We compare our multi-season model to a static model of the predator-prey community dynamics and study the interactions between species. Interestingly, including seasonality reveals indirect interactions between migrants and residents not captured by the static model. Further, we find that the direction and magnitude of interactions between two species are not necessarily accurate using only summer time-series. Our study demonstrates the need for the development of multi-season models and provides the tools to analyse them. Integrating seasonality in food web modelling is a vital step to improve predictions about the impacts of climate change on ecosystem functioning. This article is part of the theme issue 'The changing Arctic Ocean: consequences for biological communities, biogeochemical processes and ecosystem functioning'.

Linking pre-laying energy allocation and timing of breeding in a migratory arctic raptor.

For migratory species, acquisition and allocation of energy after arrival on the breeding grounds largely determine reproductive decisions. Few studies have investigated underlying physiological mechanisms driving variation in breeding phenology so far. We linked physiological state to individual timing of breeding in pre-laying arctic-nesting female peregrine falcons (Falco peregrinus tundrius). We captured females from two populations 2-20 days before egg-laying to assess plasma concentration of ß-hydroxybutyric acid (BUTY) and triglyceride (TRIG), two metabolites known to reflect short-term changes in fasting and fattening rate, respectively. We also assessed baseline corticosterone (CORTb), a hormone that mediates energy allocation, and the scaled mass index (SMI) as an indicator of somatic body reserves. Plasma BUTY was slightly higher during the pre-recruiting period compared to the period of rapid follicular growth, indicating a reduction in catabolism of lipid reserves before investment in follicle development. Conversely, TRIG levels increased in pre-recruiting females, and best-predicted individual variation in pre-laying interval and lay date. A marked increase in CORTb occurred concomitantly with the onset of rapid follicle growth. SMI was highly variable possibly reflecting variation in food availability or individuals at different stages. Results suggest that (1) lower rates of pre-laying fattening and/or lower mobilization rate of lipoproteins to ovarian follicles delayed laying, and (2) an elevation in pre-laying CORTb may result from, or be required to compensate for, the energetic costs of egg production. Results of this study illustrate how variation in the allocation of energy before laying can influence individual fitness-related reproductive decisions.

Thirty Gigahertz Optoelectronic Mixing in Chemical Vapor Deposited Graphene.

The remarkable properties of graphene, such as broadband optical absorption, high carrier mobility, and short photogenerated carrier lifetime, are particularly attractive for high-frequency optoelectronic devices operating at 1.55 µm telecom wavelength. Moreover, the possibility to transfer graphene on a silicon substrate using a complementary metal-oxide-semiconductor-compatible process opens the ability to integrate electronics and optics on a single cost-effective chip. Here, we report an optoelectronic mixer based on chemical vapor-deposited graphene transferred on an oxidized silicon substrate. Our device consists in a coplanar waveguide that integrates a graphene channel, passivated with an atomic layer-deposited Al2O3 film. With this new structure, 30 GHz optoelectronic mixing in commercially available graphene is demonstrated for the first time. In particular, using a 30 GHz intensity-modulated optical signal and a 29.9 GHz electrical signal, we show frequency downconversion to 100 MHz. These results open promising perspectives in the domain of optoelectronics for radar and radio-communication systems.

Energetic Physiology Mediates Individual Optimization of Breeding Phenology in a Migratory Arctic Seabird.

The influence of variation in individual state on key reproductive decisions impacting fitness is well appreciated in evolutionary ecology. Rowe et al. (1994) developed a condition-dependent individual optimization model predicting that three key factors impact the ability of migratory female birds to individually optimize breeding phenology to maximize fitness in seasonal environments: arrival condition, arrival date, and ability to gain in condition on the breeding grounds. While empirical studies have confirmed that greater arrival body mass and earlier arrival dates result in earlier laying, no study has assessed whether individual variation in energetic management of condition gain effects this key fitness-related decision. Using an 8-year data set from over 350 prebreeding female Arctic common eiders (Somateria mollissima), we tested this component of the model by examining whether individual variation in two physiological traits influencing energetic management (plasma triglycerides: physiological fattening rate; baseline corticosterone: energetic demand) predicted individual variation in breeding phenology after controlling for arrival date and body mass. As predicted by the optimization model, individuals with higher fattening rates and lower energetic demand had the earliest breeding phenology (shortest delays between arrival and laying; earliest laying dates). Our results are the first to empirically determine that individual flexibility in prebreeding energetic management influences key fitness-related reproductive decisions, suggesting that individuals have the capacity to optimally manage reproductive investment.

Feather corticosterone reveals effect of moulting conditions in the autumn on subsequent reproductive output and survival in an Arctic migratory bird.

For birds, unpredictable environments during the energetically stressful times of moulting and breeding are expected to have negative fitness effects. Detecting those effects however, might be difficult if individuals modulate their physiology and/or behaviours in ways to minimize short-term fitness costs. Corticosterone in feathers (CORTf) is thought to provide information on total baseline and stress-induced CORT levels at moulting and is an integrated measure of hypothalamic-pituitary-adrenal activity during the time feathers are grown. We predicted that CORTf levels in northern common eider females would relate to subsequent body condition, reproductive success and survival, in a population of eiders nesting in the eastern Canadian Arctic during a capricious period marked by annual avian cholera outbreaks. We collected CORTf data from feathers grown during previous moult in autumn and data on phenology of subsequent reproduction and survival for 242 eider females over 5 years. Using path analyses, we detected a direct relationship between CORTf and arrival date and body condition the following year. CORTf also had negative indirect relationships with both eider reproductive success and survival of eiders during an avian cholera outbreak. This indirect effect was dramatic with a reduction of approximately 30% in subsequent survival of eiders during an avian cholera outbreak when mean CORTf increased by 1 standard deviation. This study highlights the importance of events or processes occurring during moult on subsequent expression of life-history traits and relation to individual fitness, and shows that information from non-destructive sampling of individuals can track carry-over effects across seasons.

Pre-breeding energetic management in a mixed-strategy breeder.

Integrative biologists have long appreciated that the effective acquisition and management of energy prior to breeding should strongly influence fitness-related reproductive decisions (timing of breeding and reproductive investment). However, because of the difficulty in capturing pre-breeding individuals, and the tendency towards abandonment of reproduction after capture, we know little about the underlying mechanisms of these life-history decisions. Over 10 years, we captured free-living, arctic-breeding common eiders (Somateria mollissima) up to 3 weeks before investment in reproduction. We examined and characterized physiological parameters predicted to influence energetic management by sampling baseline plasma glucocorticoids (i.e., corticosterone), very-low-density lipoprotein (VLDL), and vitellogenin (VTG) for their respective roles in mediating energetic balance, rate of condition gain (physiological fattening rate) and reproductive investment. Baseline corticosterone increased significantly from arrival to the initiation of reproductive investment (period of rapid follicular growth; RFG), and showed a positive relationship with body mass, indicating that this hormone may stimulate foraging behaviour to facilitate both fat deposition and investment in egg production. In support of this, we found that VLDL increased throughout the pre-breeding period, peaking as predicted during RFG. Female eiders exhibited unprecedentedly high levels of VTG well before their theoretical RFG period, a potential strategy for pre-emptively depositing available protein stores into follicles while females are simultaneously fattening. This study provides some of the first data examining the temporal dynamics and interaction of the energetic mechanisms thought to be at the heart of individual variation in reproductive decisions and success in many vertebrate species.

Evolutionary kinetics of graphene formation on copper.

It has been claimed that graphene growth on copper by chemical vapor deposition is dominated by crystallization from the surface initially supersaturated with carbon adatoms, which implies that the growth is independent of hydrocarbon addition after the nucleation phase. Here, we present an alternative growth model based on our observations that oppose this claim. Our Gompertzian sigmoidal growth kinetics and secondary nucleation behavior support the postulate that the growth can be controlled by adsorption-desorption dynamics and the dispersive kinetic processes of catalytic dissociation and dehydrogenation of carbon precursors on copper.

European birds adjust their flight initiation distance to road speed limits.

Behavioural responses can help species persist in habitats modified by humans. Roads and traffic greatly affect animals' mortality not only through habitat structure modifications but also through direct mortality owing to collisions. Although species are known to differ in their sensitivity to the risk of collision, whether individuals can change their behaviour in response to this is still unknown. Here, we tested whether common European birds changed their flight initiation distances (FIDs) in response to vehicles according to road speed limit (a known factor affecting killing rates on roads) and vehicle speed. We found that FID increased with speed limit, although vehicle speed had no effect. This suggests that birds adjust their flight distance to speed limit, which may reduce collision risks and decrease mortality maximizing the time allocated to foraging behaviours. Mobility and territory size are likely to affect an individuals' ability to respond adaptively to local speed limits.

Probing Carrier Dynamics in Large-Scale MBE-Grown PtSe2 Films by Terahertz Spectroscopy.

Atomically thin platinum diselenide (PtSe2) films are promising for applications in the fields of electronics, spintronics, and photodetectors owing to their tunable electronic structure and high carrier mobility. Using terahertz (THz) spectroscopy techniques, we investigated the layer-dependent semiconducting-to-metallic phase transition and associated intrinsic carrier dynamics in large-scale PtSe2 films grown by molecular beam epitaxy. The uniformity of large-scale PtSe2 films was characterized by spatially and frequency-resolved THz-based sheet conductivity mapping. Furthermore, we use an optical-pump-THz-probe technique to study the transport dynamics of photoexcited carriers and explore light-induced intergrain carrier transport in PtSe2 films. We demonstrate large-scale THz-based mapping of the electrical properties of transition metal dichalcogenide films and show that the two noncontact THz-based approaches provide insight in the spatial and temporal properties of PtSe2 films.

Synthesis of Few-Layered Transition-Metal Dichalcogenides by Ion Implantation of Chalcogen and Metal Species into Sapphire.

The growth of transition-metal dichalcogenides (TMDCs) has been performed so far using most established thin-film growth techniques (e.g., vapor phase transport, chemical vapor deposition, molecular beam epitaxy, etc.). However, because there exists no self-limiting mechanism for the growth of TMDCs, none of these techniques allows precise control of the number of TMDC layers over large substrate areas. Here, we explore the ion implantation of the parent TMDC atoms into a chemically neutral substrate for the synthesis of TMDC films. The idea is that once all of the ion-implanted species have reacted together, the synthesis reaction stops, thereby effectively stopping growth. In other words, even if there is no self-limiting mechanism, growth stops when the nutrients are exhausted. We have co-implanted Mo and S ions into c-oriented sapphire substrates using various doses corresponding to 1- to 5-layer atom counts. We find that the subsurface region of the sapphire substrates is amorphized by the ion implantation process, at least for implanted doses of 2-layer atom counts and over. For all doses, we have observed the formation of MoS2 material inside the sapphire after postimplantation annealing between 800 and 850 °C. We report that the order of implantation (i.e., whether S or Mo is implanted first) is an important parameter. More precisely, samples for which S is implanted first tend to yield thin crystals with a large lateral extension (more than 200 nm for 5-layer doses) and mainly located at the interface between the amorphized and crystalline sapphire. When Mo is first implanted, the MoS2 crystals still predominantly appear at the amorphous-crystalline interface (which is much rougher), but they are much thicker, suggesting a different nucleation mechanism.

Manipulating individual state during migration provides evidence for carry-over effects modulated by environmental conditions.

Despite observational evidence of carry-over effects (COEs, events occurring in one season that produce residual effects on individuals the following seasons), to our knowledge no experimental studies have been carried out to explore how COEs might affect reproductive output. We simulated an environmental perturbation affecting spring-staging migrants to investigate COEs in greater snow geese (Anser caerulescens atlanticus). During three consecutive years, 2037 females captured during spring staging (approx. 3000 km south of their Arctic breeding grounds) were maintained in captivity (with or without access to food) for 0-4 days. Duration of captivity (but not food treatment) negatively affected reproductive success, probably through stress response. Reproductive success was reduced by 45-71% in 2 years, but not in a third year with unusually favourable breeding conditions. This unprecedented manipulation indicates that COEs can have a strong effect on individual reproductive success in long-distance migrants, but that this effect can be partly compensated for by good environmental conditions on the breeding ground.

Ultra-light photosensor collars to monitor Arctic lemming activity.

Background: Studying the anti-predatory behavior of mammals represents an important challenge, especially for fossorial small mammals that hide in burrows. In the Arctic, such behaviors are critical to the survival of lemmings considering that predation risks are high every summer. Because detailed information about how lemmings use burrows as hideouts is still lacking, we developed a 1.59 g photosensitive collar to record any event of a small mammal moving between a dark area (e.g., burrow) and a bright area (e.g., outside the burrow). Tests of how collars affected lemming behavior were conducted in captivity in Cambridge Bay, Nunavut, Canada, in November 2019 and field tests were conducted on Bylot Island, Nunavut, Canada, in August 2021. Results: The device was made of two chemical batteries and a printed circuit board (PCB) equipped with a photosensor and a real-time clock that recorded amplitude transient thresholds of light (lux) continuously. In accordance with ethical use of such devices, we verified that no abnormal loss of body mass was observed in captive or free-ranging lemmings, and no difference in recapture rates were observed between those with and without a collar, though we could not test this for periods longer than 108 h. Measurements of light intensities revealed consistent patterns with high lux levels at mid-day and lowest during the night. Lemmings showed clearly defined behavioral patterns alternating between periods outside and inside burrows. Despite 24-h daylight in the middle of the summer, August nighttime (i.e., 11 PM to 4 AM) lux levels were insufficient for amplitude transient thresholds to be reached. Conclusion: By taking advantage of the long periods of daylight in the Arctic, such technology is very promising as it sets new bases for passive recording of behavioral parameters and builds on the prospect of further miniaturization of batteries and PCBs.

Microwave devices: carbon nanotubes as cold cathodes.

To communicate, spacecraft and satellites rely on microwave devices, which at present are based on relatively inefficient thermionic electron sources that require heating and cannot be switched on instantaneously. Here we describe a microwave diode that uses a cold-cathode electron source consisting of carbon nanotubes and that operates at high frequency and at high current densities. Because it weighs little, responds instantaneously and has no need of heating, this miniaturized electron source should prove valuable for microwave devices used in telecommunications.

Do glucocorticoids in droppings reflect baseline level in birds captured in the wild? A case study in snow geese.

Baseline glucocorticoid (CORT) levels in plasma are increasingly used as physiological indices of the relative condition or health of individuals and populations. The major limitation is that CORT production is stimulated by the stress associated with capture and handling. Measuring fecal CORT is one way to solve this problem because elevation of fecal CORT usually does not occur before 1-12h after a stressful event in captive animals. However, the effect of capture and handling on fecal CORT levels has seldom been investigated in the wild. In a first experiment, we validated that fecal CORT levels starts to increase in droppings (a mixture of fecal and urinary material) about 1-2h following injection of CORT-release hormone (ACTH) in captive greater snow geese (Chen caerulescens atlantica). In a second experiment, we investigated whether dropping and plasma CORT were related and if the capture affected fecal CORT levels in wild birds. Baseline CORT was obtained by bleeding individuals within 4 min after capture. No relationship was found between baseline and CORT in droppings shortly after capture (<4 min). In addition, CORT levels in droppings increased linearly with time after capture and was already elevated by a factor two 40 min after capture. The different turnover time of CORT between urine and feces could explain such results. We conclude that droppings cannot provide an index of basal CORT levels in snow geese captured in the wild. Such a result contrast with previous studies conducted on habituated, captive animals. We thus recommend that use of droppings as a non-invasive technique to measure baseline CORT be restricted to non-manipulated individuals in the wild.