High juvenile mortality during migration in a declining population of a long‐distance migratory raptor

Many populations of long‐distance migrants are declining and there is increasing evidence that declines may be caused by factors operating outside the breeding season. Among the four vulture species breeding in the western Palaearctic, the species showing the steepest population decline, the Egyptian Vulture Neophron percnopterus, is a long‐distance migrant wintering in Africa. However, the flyways and wintering areas of the species are only known for some populations, and without knowledge of where mortality occurs, effective conservation management is not possible. We tracked 19 juvenile Egyptian Vultures from the declining breeding population on the Balkan Peninsula between 2010 and 2014 to estimate survival and identify important migratory routes and wintering areas for this species. Mortality during the first autumn migration was high (monthly survival probability 0.75) but mortality during migration was exclusively associated with suboptimal navigation. All birds from western breeding areas and three birds from central and eastern breeding areas attempted to fly south over the Mediterranean Sea, but only one in 10 birds survived this route, probably due to stronger tailwind. All eight birds using the migratory route via Turkey and the Middle East successfully completed their first autumn migration. Of 14 individual and environmental variables examined to explain why juvenile birds did or did not successfully complete their first migration, the natal origin of the bird was the most influential. We speculate that in a declining population with fewer experienced adults, an increasing proportion of juvenile birds are forced to migrate without conspecific guidance, leading to high mortality as a consequence of following sub‐optimal migratory routes. Juvenile Egyptian Vultures wintered across a vast range of the Sahel and eastern Africa, and had large movement ranges with core use areas at intermediate elevations in savannah, cropland or desert. Two birds were shot in Africa, where several significant threats exist for vultures at continental scales. Given the broad distribution of the birds and threats, effective conservation in Africa will be challenging and will require long‐term investment. We recommend that in the short term, more efficient conservation could target narrow migration corridors in southern Turkey and the Middle East, and known congregation sites in African wintering areas.     

“Closer‐to‐home” strategy benefits juvenile survival in a long‐distance migratory bird

Human‐induced changes in the climate and environment that occur at an unprecedented speed are challenging the existence of migratory species. Faced with these new challenges, species with diverse and flexible migratory behaviors may suffer less from population decline, as they may be better at responding to these changes by altering their migratory behavior. At the individual level, variations in migratory behavior may lead to differences in fitness and subsequently influence the population's demographic dynamics. Using lifetime GPS bio‐logging data from 169 white storks (Ciconia ciconia), we explore whether the recently shortened migration distance of storks affects their survival during different stages of their juvenile life. We also explore how other variations in migratory decisions (i.e., time, destination), movement activity (measured using overall body dynamic acceleration), and early life conditions influence juvenile survival. We observed that their first autumn migration was the riskiest period for juvenile white storks. Individuals that migrated shorter distances and fledged earlier experienced lower mortality risks. In addition, higher movement activity and overwintering “closer‐to‐home” (with 84.21% of the tracked individuals stayed Europe or North Africa) were associated with higher survival. Our study shows how avian migrants can change life history decisions over only a few decades, and thus it helps us to understand and predict how migrants respond to the rapidly changing world.     

Finish with a sprint: Evidence for time‐selected last leg of migration in a long‐distance migratory songbird

Under time‐selected migration, birds should choose a strategy for outcompeting rivals over securing access to prime resources at the final destination. Thus, migration can be viewed as a race among individuals where winners are arriving first when conditions are suitable. The sprint migration hypothesis predicts that individuals shift from maximum sustained speed to a final burst of sprint to shorten the transition from migration to breeding (Alerstam, 2006). In this study, we test the hypothesis of a final sprint migration in a long‐distance Afro‐Palearctic migrant, the collared flycatcher Ficedula albicollis, during autumn and spring, and compare migration strategies between the seasons. In both seasons, collared flycatchers evidently exhibited sprint migration by increasing their overall speed over the last leg of migration after the Sahara crossing. This phenomenon was more pronounced in spring, contributing to overall faster spring migration and possibly highlighting higher importance for early arrival at the breeding grounds. In both seasons and particularly in spring, late departing individuals flew at a faster rate, partially being able to catch up with their early departing conspecifics. Differential fueling strategies may play an important role in determining migration speed, especially during the early stages of the migration, and might explain the observed differences in migration speeds between late and early departing individuals. Our findings suggest competition for early arrival at the breeding and at the nonbreeding destinations alike. Sprint migration might be an appropriate strategy to gain advantage over conspecifics and settle in prime territories as well as to cope with the increasingly earlier springs at high latitudes.     

Repeatability of individual migration routes,wintering sites,and timing in a long‐distance migrant bird

Migratory birds are often faithful to wintering (nonbreeding) sites, and also migration timing is usually remarkably consistent, that is, highly repeatable. Spatiotemporal repeatability can be of advantage for multiple reasons, including familiarity with local resources and predators as well as avoiding the costs of finding a new place, for example, nesting grounds. However, when the environment is variable in space and time, variable site selection and timing might be more rewarding. To date, studies on spatial and temporal repeatability in short‐lived long‐distance migrants are scarce, most notably of first‐time and subsequent migrations. Here, we investigated repeatability in autumn migration directions, wintering sites, and annual migration timing in Hoopoes (Upupa epops), a long‐distance migrant, using repeated tracks of adult and first‐time migrants. Even though autumn migration directions were mostly the same, individual wintering sites often changed from year to year with distances between wintering sites exceeding 1,000 km. The timing of migration was repeatable within an individual during autumn, but not during spring migration. We suggest that Hoopoes respond to variable environmental conditions such as north–south shifts in rainfall during winter and differing onset of the food availability during spring migration.     

Spatial and temporal variation in population trends in a long‐distance migratory bird

Aim Over the past three decades, evidence has been growing that many Afro‐Palaearctic migratory bird populations have suffered sustained and severe declines. As causes of these declines exist across both the breeding and non‐breeding season, identifying potential drivers of population change is complex. In order to explore the roles of changes in regional and local environmental conditions on population change, we examine spatial and temporal variation in population trajectories of one of Europe’s most abundant Afro‐Palaearctic summer migrants, the willow warbler, Phylloscopus trochilus. Location Britain and Ireland. Methods We use national survey data from Britain and Ireland (BBS: BTO/RSPB/JNCC Breeding Bird Survey and CBS: BWI/NPWS/Heritage Council Countryside Breeding Survey) from 1994 to 2006 to model the spatial and temporal variation in willow warbler population trends. Results Across Britain and Ireland, population trends follow a gradient from sharp declines in the south and east of England to shallow declines and/or slight increases in parts of north and west England, across Scotland and Ireland. Decreasing the spatial scale of analysis reveals variation in both the rate and spatial extent of population change within central England and the majority of Scotland. The rates of population change also vary temporally; declines in the south of England are shallower now than at the start of the time series, whereas populations further north in Britain have undergone periods of increase and decline. Main conclusion These patterns suggest that regional‐scale drivers, such as changing climatic conditions, and local‐scale processes, such as habitat change, are interacting to produce spatially variable population trends. We discuss the potential mechanisms underlying these interactions and the challenges in addressing such changes at scales relevant to migratory species.     

Rainfall at African wintering grounds predicts age‐specific probability of haemosporidian infection in a migratory passerine bird

In migratory species breeding in temperate zones and wintering in tropical areas, the prevalence of blood parasites may be affected by migratory strategies and winter habitat choice. We explored whether African winter habitat was linked to the probability of haemosporidian infection in the House Martin Delichon urbicum breeding in Spain, and tested for potential differences between age‐classes. As a proxy for winter habitat features, we analysed stable isotope (δ²H, δ¹³C and δ¹⁵N) values of winter‐grown feathers moulted in tropical Africa. Rainfall at the African winter grounds was related to the probability of being infected with haemosporidians and this effect differed among age‐classes. We found that haemosporidian prevalence was similar for young and experienced birds wintering in habitats of higher rainfall (²H‐depleted), whereas there were great differences in winter habitats of lower rainfall (²H‐enriched), with young having a much higher prevalence compared with experienced birds. Likewise, experienced birds wintering in habitats of higher rainfall had a higher probability of haemosporidian infection compared with experienced birds wintering in habitats of lower rainfall. By contrast, young birds wintering in habitats of lower rainfall had a higher probability of haemosporidian infection compared with young birds wintering in habitats of higher rainfall. These outcomes highlight the interaction of age with haemosporidian infection in the migratory ecology of the House Martin, which may drive carry‐over effects in this long‐distance aerial insectivore.     

Can long‐distance migratory birds adjust to the advancement of spring by shortening migration distance? The response of the pied flycatcher to latitudinal photoperiodic variation

Many organisms use day length as a cue for synchronizing their life cycles with seasonal changes in environmental productivity. Under rapid climate change, however, responses to day length may become maladaptive, and photo‐responsive organisms may only be able to evade increasingly unsuitable habitats if they can accommodate to a wide range of photoperiodic conditions. A previous experiment showed that the pied flycatcher, Ficedula hypoleuca, a Palaearctic‐Afrotropical migratory bird, would strongly advance the timing of spring migration and reproductive maturation if it shifted its wintering area from sub‐Saharan Africa to the Mediterranean region. However, it is unknown whether this marked response to latitudinal variation in photoperiodic conditions is continuous over the entire range of potential wintering areas, and if a shortening of migration distance would be an effective mechanism to adjust the timing of migration to rapidly changing climatic conditions. Here, we experimentally show that a moderate northward displacement of the pied flycatcher's current wintering grounds by 10° would result in a clear advancement of the termination of prenuptial moult and the initiation of spring migratory activity and gonadal growth. However, we found no further advancement under conditions simulating higher wintering latitudes, suggesting the existence of a critical photoperiodic threshold or a steep gradual response within a narrow geographical range between 10° and 20° northern latitude. Because habitat conditions in this area are deteriorating rapidly, the potential for pied flycatchers to adjust their life cycle to changing climatic conditions by shortening the migration distance may be limited in the future.     

Disentangling the effects of environmental conditions on wintering and breeding grounds on age‐specific survival rates in a trans‐Saharan migratory raptor

Migratory species are subject to environmental variability occurring on breeding and wintering grounds. Estimating the relative contribution of environmental factors experienced sequentially during breeding and wintering, and their potential interaction, to the variation of survival is crucial to predict population viability of migratory species. Here we investigated this issue for the Montagu's harrier Circus pygargus, a trans‐Saharan migrant. We analysed capture–recapture data from a 29‐year long monitoring of wing‐tagged offspring and adults at two study sites in France (Rochefort‐RO and Maine‐et‐Loire‐ML). The study period covers a climatic shift occurring in the Sahel with increasing rainfall following a period of droughts (Sahel greening). We found that harriers’ adult survival in RO (between 1988 and 2005) varied over time and was sensitive to the interaction between the amount of rainfall in the Sahel and the annual mean breeding success, two proxies of prey availability. The occurrence of adverse conditions on breeding and wintering grounds in the same year decreased survival from 0.70–0.77 to 0.48 ± 0.05. Juvenile survival in RO was slightly more sensitive to conditions in Europe than in the Sahel. Unexpectedly, lower survival rates were found in years with higher mean breeding success, suggesting compensatory density feedbacks may operate. By contrast, adult survival in ML, monitored between 1999 and 2017, was higher compared to RO (0.76 ± 0.03 versus 0.66 ± 0.02), remained constant and unaffected by any proxy of prey availability. This difference seems consistent with the fact that harriers in ML experienced better and especially less variable environmental conditions during breeding and wintering seasons compared to RO. Overall, we showed that survival of a migratory bird is sensitive to the level of variability in environmental conditions and that adverse conditions on wintering grounds can amplify the negative effects of conditions during the previous breeding season on birds’ survival.     

Short‐ and long‐term costs of reproduction in a migratory songbird

Costs of reproduction represent a common life‐history trade‐off. Critical to understanding these costs in migratory species is the ability to track individuals across successive stages of the annual cycle. We assessed the effects of total number of offspring fledged and date of breeding completion on pre‐migratory body condition, the schedule of moult and annual survival in a migratory songbird, the Savannah Sparrow Passerculus sandwichensis. Between 2008 and 2010, moult was delayed for individuals that finished breeding later in the breeding period and resulted in reduced lean tissue mass during the pre‐migratory period, suggesting an indirect trade‐off between the timing of breeding completion and condition just prior to migration. Lean tissue mass decreased as the number of offspring fledged increased in 2009, a particularly cool and wet year, illustrating a direct trade‐off between reproductive effort and condition just prior to migration in years when weather is poor. However, using a 17‐year dataset from the same population, we found that parents that fledged young late in the breeding period had the highest survival and that number of offspring fledged did not affect survival, suggesting that individuals do not experience long‐term trade‐offs between reproduction and survival. Taken together, our results suggest that adult Savannah Sparrows pay short‐term costs of reproduction, but that longer‐term costs are mitigated by individual quality, perhaps through individual variation in resource acquisition.     

Around the Mediterranean: an extreme example of loop migration in a long‐distance migratory passerine

An important issue in migration research is how small‐bodied passerines pass over vast geographical barriers; in European–African avian migration, these are represented by the Mediterranean Sea and the Sahara Desert. Eastern (passing eastern Mediterranean), central (passing Apennine Peninsula) and western (via western Mediterranean) major migration flyways are distinguished for European migratory birds. The autumn and spring migration routes may differ (loop migration) and there could be a certain level of individual flexibility in how individuals navigate themselves during a single migration cycle. We used light‐level loggers to map migration routes of barn swallows Hirundo rustica breeding in the centre of a wide putative contact zone between the northeastern and southernwestern European populations that differ in migration flyways utilised and wintering grounds. Our data documented high variation in migration patterns and wintering sites of tracked birds (n = 19 individuals) from a single breeding colony, with evidence for loop migration in all but one of the tracked swallows. In general, two migratory strategies were distinguished. In the first, birds wintering in a belt stretching from southcentral to southern Africa that used an eastern route for both the spring and autumn migration, then shifted their spring migration eastwards (anti‐clockwise loops, n = 12). In the second, birds used an eastern or central route to their wintering grounds in central Africa, shifting the spring migration route westward (clockwise loops, n = 7). In addition, we observed an extremely wide clockwise loop migration encompassing the entire Mediterranean, with one individual utilising both the eastern (autumn) and western (spring) migratory flyway during a single annual migration cycle. Further investigation is needed to ascertain whether clockwise migratory loops encircling the entire Mediterranean also occur other small long‐distance passerine species.     

Quantifying full phenological event distributions reveals simultaneous advances,temporal stability and delays in spring and autumn migration timing in long‐distance migratory birds

Phenological changes in key seasonally expressed life‐history traits occurring across periods of climatic and environmental change can cause temporal mismatches between interacting species, and thereby impact population and community dynamics. However, studies quantifying long‐term phenological changes have commonly only measured variation occurring in spring, measured as the first or mean dates on which focal traits or events were observed. Few studies have considered seasonally paired events spanning spring and autumn or tested the key assumption that single convenient metrics accurately capture entire event distributions. We used 60 years (1955–2014) of daily bird migration census data from Fair Isle, Scotland, to comprehensively quantify the degree to which the full distributions of spring and autumn migration timing of 13 species of long‐distance migratory bird changed across a period of substantial climatic and environmental change. In most species, mean spring and autumn migration dates changed little. However, the early migration phase (≤10th percentile date) commonly got earlier, while the late migration phase (≥90th percentile date) commonly got later. Consequently, species' total migration durations typically lengthened across years. Spring and autumn migration phenologies were not consistently correlated within or between years within species and hence were not tightly coupled. Furthermore, different metrics quantifying different aspects of migration phenology within seasons were not strongly cross‐correlated, meaning that no single metric adequately described the full pattern of phenological change. These analyses therefore reveal complex patterns of simultaneous advancement, temporal stability and delay in spring and autumn migration phenologies, altering species' life‐history structures. Additionally, they demonstrate that this complexity is only revealed if multiple metrics encompassing entire seasonal event distributions, rather than single metrics, are used to quantify phenological change. Existing evidence of long‐term phenological changes detected using only one or two metrics should consequently be interpreted cautiously because divergent changes occurring simultaneously could potentially have remained undetected.     

Carry‐over effects of conditions at the wintering grounds on breeding plumage signals in a migratory bird: roles of phenotypic plasticity and selection

To understand the consequences of ever‐changing environment on the dynamics of phenotypic traits, distinguishing between selection processes and individual plasticity is crucial. We examined individual consistency/plasticity in several male secondary sexual traits expressed during the breeding season (white wing and forehead patch size, UV reflectance of white wing patch and dorsal melanin coloration) in a migratory pied flycatcher (Ficedula hypoleuca) population over an 11‐year period. Furthermore, we studied carry‐over effects of three environmental variables (NAO, a climatic index; NDVI, a vegetation index; and rainfall) at the wintering grounds (during prebreeding moult) on the expression of these breeding plumage traits of pied flycatcher males at individual and population levels. Whereas NAO correlates negatively with moisture in West Africa, NDVI correlates positively with primary production. Forehead patch size and melanin coloration were highly consistent within individuals among years, whereas the consistency of the other two traits was moderate. Wing patch size decreased with higher NAO and increased with higher rainfall and NDVI at the individual level. Interestingly, small‐patched males suffered lower survival during high NAO winters than large‐patched males, and vice versa during low NAO winters. These counteracting processes meant that the individual‐level change was masked at the population level where no relationship was found. Our results provide a good example of how variation in the phenotypic composition of a natural population can be a result of both environment‐dependent individual plasticity and short‐term microevolution. Moreover, when plasticity and viability selection operate simultaneously, their impacts on population composition may not be evident.     

Low and annually variable migratory connectivity in a long‐distance migrant: Whinchats Saxicola rubetra may show a bet‐hedging strategy

The spatial scale of non‐breeding areas used by long‐distance migrant animals can vary from specific, relatively small non‐breeding areas for each independent breeding population (high connectivity) to a distribution over a large non‐breeding area with mixing of breeding populations (low connectivity). Measuring variation in the degree of connectivity and how it arises is crucial to predict how migratory animals can respond to global habitat and climate change because low connectivity is likely to be an adaptation to environmental uncertainty. Here, we assess whether use of non‐breeding areas in a long‐distance migrant may be stochastic by measuring the degree of connectivity, and whether it is annually variable. Twenty‐nine wintering Whinchats tagged with geolocators over 2 years within 40 km² in central Nigeria were found to be breeding over 2.55 million km² (26% of the land area of Europe), without an asymptote being approached in the relationship between area and sample size. Ranges differed in size between years by 1.51 million km² and only 15% of the total breeding range across both years overlapped (8% overlap between years when only first‐year birds were considered), well above the range size difference and below the proportion of overlap that would be predicted from two equivalent groups breeding at random locations within the observed range. Mean distance between breeding locations (i.e. migratory spread) differed significantly between years (604 ± 18 km in 2013 and 869 ± 33 km in 2014). The results showed very low and variable connectivity that was reasonably robust to the errors and assumptions inherent in the use of geolocators, but with the caveat of having only ranges of 2 years to compare, and the sensitivity of range to the breeding locations of a small number of individuals. However, if representative, the results suggest the scope for between‐year variation (cohort effects) to determine migrant distribution on a large scale. Furthermore, for species with similarly low connectivity, we would predict breeding population trends to reflect average conditions across large non‐breeding areas: thus, as large areas of Africa become subject to habitat loss, migrant populations throughout Europe will decline.     

Density‐dependent survival and recruitment in a long‐distance Palaearctic migrant,the Sand Martin Riparia riparia

Long‐term studies can provide powerful insights into the relative importance of different demographic and environmental factors determining avian population dynamics. Here we use 23 years of capture–mark–recapture data (1981–2003) to estimate recruitment and survival rates for a Sand Martin Riparia riparia population in Cheshire, NW England. Inter‐annual variation in recruitment and adult survival was positively related to rainfall in the sub‐Saharan wintering grounds, but unrelated to weather conditions on the breeding grounds. After allowing for the effects of African rainfall, both demographic rates were negatively density‐dependent: adult survival was related to the size of the western European Sand Martin population (probably reflecting competition for resources in the shared wintering grounds) while recruitment was related to the size of the local study population in Cheshire (potentially reflecting competition for nesting sites or food). Local population size was more sensitive to variation in adult survival than to variation in recruitment, and an increase in population size after 1995 was driven mainly by the impact of more favourable conditions in the African wintering grounds on survival rates of adults. Overwinter survival in this long‐distance Palaearctic migrant is determined partly by the amount of suitable wetland foraging habitat in the sub‐Saharan wintering grounds (which is limited by the extent of summer rainfall) and partly by the number of birds exploiting that habitat.     

Foraging conditions ‘at the end of the world’ in the context of long‐distance migration and population declines in red knots

The long‐distance migrant red knot (Calidris canutus ssp. rufa– Scolopacidae) alternates between the northern and southern ends of the New World, one of the longest yearly migrations of any bird and paradoxically overflying apparently suitable habitat at lower latitudes. This subspecies is sharply declining, with a major mortality event following 2000, attributed to commercial overharvesting of food resources at its Delaware Bay (USA) stop‐over site. A full understanding of this peculiar migrant requires an assessment of the foraging conditions at its southern hemisphere wintering sites. Here, for a major wintering site in Argentinean Tierra del Fuego (Río Grande), we describe and compare food abundance, diet and intake rates during January–February in 1995, 2000 and 2008. The two main prey types were the burrowing clam Darina solenoides and three species of epibenthic mussels Mytilidae. In the year 2000, food availability and intake rate were higher than those recorded at other sites used by knots anywhere else in the world, contributing to the explanation of why red knots carry out this impressive migration. Intake rate in 2008 on the two main prey types was dramatically reduced as a result of birds eating smaller prey and strongly increased human disturbance; the same year we also found a high prevalence of a digenean parasite in Darina. We suggest that during the strongly enhanced winter mortality in 2000, knots did not yet face ecological problems in their southernmost wintering area, consistent with the previous evidence that problems at northern stop‐overs negatively affected their numbers. However, in 2008 the ecological conditions at Río Grande were such that they would have facilitated a further decline, emphasizing the importance of a hemispheric approach to research and management.     

Changes in species interactions across a 2.5 km elevation gradient: effects on plant migration in response to climate change

Predicted climate change in the Andes will require plant species to migrate upslope to avoid extinction. Central to predictions of species responses to climate change is an understanding of species distributions along environmental gradients. Environmental gradients are frequently modelled as abiotic, but biotic interactions can play important roles in setting species distributions, abundances, and life history traits. Biotic interactions also have the potential to influence species responses to climate change, yet they remain mostly unquantified. An important interaction long studied in tropical forests is postdispersal seed predation which has been shown to affect the population dynamics, community structure, and diversity of plant species in time and space. This paper presents a comparative seed predation study of 24 species of tropical trees across a 2.5 km elevation gradient in the Peruvian Andes and quantifies seed predation variation across the elevational gradient. We then use demographic modelling to assess effects of the observed variation in seed predation on population growth rates in response to observed increasing temperatures in the area. We found marked variation among species in total seed predation depending on the major seed predator of the species and consistent changes in seed predation across the gradient. There was a significant increase in seed survival with increasing elevation, a trend that appears to be driven by regulation of seed predators via top–down forces in the lowlands giving way to bottom–up (productivity) regulation at mid‐ to high elevations, resulting in a ninefold increase in effective fecundity for trees at high elevations. This potential increase in seed crop size strongly affects modelled plant population growth and seed dispersal distances, increasing population migration potential in the face of climate change. These results also indicate that species interactions can have effects on par with climate in species responses to global change.     

Divergent long‐term trends and interannual variation in ecosystem resource use efficiencies of a southern boreal old black spruce forest 1999–2017

Long‐term trends in ecosystem resource use efficiencies (RUEs) and their controlling factors are key pieces of information for understanding how an ecosystem responds to climate change. We used continuous eddy covariance and microclimate data over the period 1999–2017 from a 120‐year‐old black spruce stand in central Saskatchewan, Canada, to assess interannual variability, long‐term trends, and key controlling factors of gross ecosystem production (GEP) and the RUEs of carbon (CUE = net primary production [NPP]/GEP), light (LUE = GEP/absorbed photosynthetic radiation [APAR]), and water (WUE = GEP/evapotranspiration [E]). At this site, annual GEP has shown an increasing trend over the 19 years (p < 0.01), which may be attributed to rising atmospheric CO₂ concentration. Interannual variability in GEP, aside from its increasing trend, was most strongly related to spring temperatures. Associated with the significant increase in annual GEP were relatively small changes in NPP, APAR, and E, so that annual CUE showed a decreasing trend and annual LUE and WUE showed increasing trends over the 19 years. The long‐term trends in the RUEs were related to the increasing CO₂ concentration. Further analysis of detrended RUEs showed that their interannual variation was impacted most strongly by air temperature. Two‐factor linear models combining CO₂ concentration and air temperature performed well (R²~0.60) in simulating annual RUEs. LUE and WUE were positively correlated both annually and seasonally, while LUE and CUE were mostly negatively correlated. Our results showed divergent long‐term trends among CUE, LUE, and WUE and highlighted the need to account for the combined effects of climatic controls and the ‘CO₂ fertilization effect’ on long‐term variations in RUEs. Since most RUE‐based models rely primarily on one resource limitation, the observed patterns of relative change among the three RUEs may have important implications for RUE‐based modeling of C fluxes.