Phenology effects on productivity and hatching-asynchrony of American kestrels (Falco sparverius) across a continent

Climate-driven advances in spring can result in phenological mismatch between brood rearing and prey availability and consequently cause decreased productivity in birds. How consequences of mismatch vary across species' ranges, and how individual behavior can mitigate mismatch effects is less studied. We quantified the relationship between phenological mismatch, productivity, and behavioral adaptations of American kestrels (Falco sparverius) across their breeding range in the United States and southern Canada. We obtained phenology and productivity data using nest observations from long term nest box monitoring, remote trail cameras, and community-scientist based programs. We collected data on parental incubation behavior and hatch asynchrony using trail cameras in nest boxes. Kestrels that laid eggs after the start of spring had higher rates of nest failure and fewer nestlings than earlier nesters, and effects of mismatch on productivity were most severe in the Northeast. In contrast, kestrels in the Southwest experienced a more gradual decline in productivity with seasonal mismatch. We attribute the effect of location to the growing season and temporal nesting windows (duration of nesting season). Specifically, resource availability in the Northeast is narrow and highly peaked during the breeding season, potentially resulting in shorter nesting windows. Conversely, resource curves may be more prolonged and dampened in the Southwest, and growing seasons are becoming longer with climate change, potentially resulting in longer nesting windows. We found that the onset of male incubation was negatively associated with lay date. Males from breeding pairs that laid eggs after the start of spring began incubation sooner than males from breeding pairs that laid before the start of spring. Early-onset male incubation was positively associated with hatching asynchrony, creating increased age variation in developing young. In sum, we demonstrate that American kestrels are vulnerable to phenological mismatch, and that this vulnerability varies across space. Northeastern populations could be more vulnerable to mismatch consequences, which may be one factor contributing to declines of kestrels in this region. Also, we demonstrate early onset of incubation as a potential adaptive behavior to advance average hatch date and spread out offspring demands, but it is unknown how impactful this will be in mitigating the fitness consequences of phenology mismatch.

Recent natural variability in global warming weakened phenological mismatch and selection on seasonal timing in great tits ( Parus major )

Climate change has led to phenological shifts in many species, but with large variation in magnitude among species and trophic levels. The poster child example of the resulting phenological mismatches between the phenology of predators and their prey is the great tit ( Parus major ), where this mismatch led to directional selection for earlier seasonal breeding. Natural climate variability can obscure the impacts of climate change over certain periods, weakening phenological mismatching and selection. Here, we show that selection on seasonal timing indeed weakened significantly over the past two decades as increases in late spring temperatures have slowed down. Consequently, there has been no further advancement in the date of peak caterpillar food abundance, while great tit phenology has continued to advance, thereby weakening the phenological mismatch. We thus show that the relationships between temperature, phenologies of prey and predator, and selection on predator phenology are robust, also in times of a slowdown of warming. Using projected temperatures from a large ensemble of climate simulations that take natural climate variability into account, we show that prey phenology is again projected to advance faster than great tit phenology in the coming decades, and therefore that long-term global warming will intensify phenological mismatches.

Extreme climate event promotes phenological mismatch between sexes in hibernating ground squirrels

Hibernating ground squirrels rely on a short active period for breeding and mass accrual, and are thus vulnerable to extreme climate events that affect key periods in their annual cycle. Here, we document how a heatwave in March 2012 led to a phenological mismatch between sexes in Richardson’s ground squirrels (Urocitellus richardsonii). Females emerged from hibernation and commenced breeding earlier in 2012 relative to average female emergence. Although males had descended testes and pigmented scrota, it appeared that not all males were physiologically prepared to breed since 58.6% of males had non-motile sperm when breeding commenced. Body condition, relative testes size, and the relative size of accessory glands were significant predictors of sperm motility. Males with non-motile sperm had smaller accessory glands than males with motile sperm. There was no decrease in the number of juveniles that emerged in 2012 or female yearlings recruited in 2013, nor did juveniles emerge later than other years. The impact of this heatwave on male ground squirrels emphasizes the importance of assessing the consequences of climate change on the breeding success of hibernating species in both sexes, since the different sensitivity to external cues for emergence led to a mismatch in timing under this event.

Climate Effects on Vertical Forest Phenology of Fagus sylvatica L., Sensed by Sentinel-2, Time Lapse Camera, and Visual Ground Observations

Contemporary climate change leads to earlier spring phenological events in Europe. In forests, in which overstory strongly regulates the microclimate beneath, it is not clear if further change equally shifts the timing of leaf unfolding for the over- and understory of main deciduous forest species, such as Fagus sylvatica L. (European beech). Furthermore, it is not known yet how this vertical phenological (mis)match—the phenological difference between overstory and understory—affects the remotely sensed satellite signal. To investigate this, we disentangled the start of season (SOS) of overstory F.sylvatica foliage from understory F. sylvatica foliage in forests, within nine quadrants of 5.8 × 5.8 km, stratified over a temperature gradient of 2.5 °C in Bavaria, southeast Germany, in the spring seasons of 2019 and 2020 using time lapse cameras and visual ground observations. We explained SOS dates and vertical phenological (mis)match by canopy temperature and compared these to Sentinel-2 derived SOS in response to canopy temperature. We found that overstory SOS advanced with higher mean April canopy temperature (visual ground observations: −2.86 days per °C; cameras: −2.57 days per °C). However, understory SOS was not significantly affected by canopy temperature. This led to an increase of vertical phenological mismatch with increased canopy temperature (visual ground observations: +3.90 days per °C; cameras: +2.52 days per °C). These results matched Sentinel-2-derived SOS responses, as pixels of higher canopy height advanced more by increased canopy temperature than pixels of lower canopy height. The results may indicate that, with further climate change, spring phenology of F. sylvatica overstory will advance more than F. sylvatica understory, leading to increased vertical phenological mismatch in temperate deciduous forests. This may have major ecological effects, but also methodological consequences for the field of remote sensing, as what the signal senses highly depends on the pixel mean canopy height and the vertical (mis)match.

Warming Arctic summers unlikely to increase productivity of shorebirds through renesting

Climate change in the Arctic is leading to earlier summers, creating a phenological mismatch between the hatching of insectivorous birds and the availability of their invertebrate prey. While phenological mismatch would presumably lower the survival of chicks, climate change is also leading to longer, warmer summers that may increase the annual productivity of birds by allowing adults to lay nests over a longer period of time, replace more nests that fail, and provide physiological relief to chicks (i.e., warmer temperatures that reduce thermoregulatory costs). However, there is little information on how these competing ecological processes will ultimately impact the demography of bird populations. In 2008 and 2009, we investigated the survival of chicks from initial and experimentally-induced replacement nests of arcticola Dunlin (Calidris alpina) breeding near Utqiaġvik, Alaska. We monitored survival of 66 broods from 41 initial and 25 replacement nests. Based on the average hatch date of each group, chick survival (up to age 15 days) from replacement nests (Ŝi = 0.10; 95% CI = 0.02–0.22) was substantially lower than initial nests (Ŝi = 0.67; 95% CI = 0.48–0.81). Daily survival rates were greater for older chicks, chicks from earlier-laid clutches, and during periods of greater invertebrate availability. As temperature was less important to daily survival rates of shorebird chicks than invertebrate availability, our results indicate that any physiological relief experienced by chicks will likely be overshadowed by the need for adequate food. Furthermore, the processes creating a phenological mismatch between hatching of shorebird young and invertebrate emergence ensures that warmer, longer breeding seasons will not translate into abundant food throughout the longer summers. Thus, despite having a greater opportunity to nest later (and potentially replace nests), young from these late-hatching broods will likely not have sufficient food to survive. Collectively, these results indicate that warmer, longer summers in the Arctic are unlikely to increase annual recruitment rates, and thus unable to compensate for low adult survival, which is typically limited by factors away from the Arctic-breeding grounds.

Shooting blanks: extreme climate event promotes phenological mismatch between sexes in hibernating ground squirrels

Hibernating ground squirrels rely on a short active period for breeding and mass accrual, and are thus vulnerable to extreme climate events that affect key periods in their annual cycle. Here, we observed how a heatwave in March 2012 led to a phenological mismatch between sexes in Richardson’s ground squirrels (Urocitellus richardsonii). Females emerged from hibernation and commenced breeding earlier in 2012 relative to average female emergence. Despite external indicators suggesting that males were prepared for breeding, it appeared that not all males were physiologically prepared since 58.6% of males had non-motile sperm when breeding commenced. We found that males with non-motile sperm had smaller accessory glands than males with motile sperm. Body condition, relative testes size, and the relative size of accessory glands were significant predictors of sperm motility. There was no difference in litter size among years, nor a decrease in the number of juveniles emerged in 2012 or female yearlings recruited in 2013. The impact of this heatwave on male ground squirrels emphasizes the importance of assessing the consequences of climate change on breeding success of hibernating species in both sexes, since the different cues for emergence led to a mismatch in timing under this event.

Plasticity in timing of avian breeding in response to spring temperature differs between early and late nesting species

Plasticity for breeding dates may influence population vulnerability to climate change via phenological mismatch between an organism’s life cycle requirements and resource availability in occupied environments. Some life history traits may constrain plasticity, however there have been remarkably few comparisons of how closely-related species, differing in key traits, respond to common phenology gradients. We compared population- and individual-level plasticity in clutch initiation dates (CID) in response to spring temperature among five duck species with early- to late-season nesting life histories. Plasticity was strongest in females of the earliest breeding species (common goldeneye [Bucephala clangula], mallard [Anas platyrhynchos], and gadwall [Mareca strepera]), whereas late-nesting lesser scaup (Aythya affinis) and white-winged scoter (Melanitta fusca deglandi) did not respond. These results contrast with previous work in other bird families that suggested late-breeders are generally more flexible. Nevertheless, late-breeding species exhibited annual variation in mean CID, suggesting response to other environmental factors unrelated to spring temperature. Goldeneye and gadwall females varied in their strength of individual plasticity (‘individual × environment’ interactions) and goldeneye and scoter females showed evidence of interannual repeatability of CID. Fitness consequences of CID plasticity in response to spring phenology, including trophic mechanisms and population consequences, warrant investigation.

Phenological Response in the Trophic Levels to Climate Change in Korea

The response of the phenological events of individual species to climate change is not isolated, but is connected through interaction with other species at the same or adjacent trophic level. Using long-term phenological data observed since 1976 in Korea, whose temperature has risen more steeply than the average global temperature, this study conducted phenological analysis (differ-ences in the phenology of groups, differences in phenological shifts due to climate change, differ-ences in phenological sensitivity to climate by groups, and the change of phenological day differ-ences among interacting groups). The phenological shift of the producer group (plants) was found to be negative in all researched species, which means that it blooms quickly over the years. The regression slope of consumers (primary consumers and secondary consumers) was generally posi-tive which means that the phenological events of these species tended to be later during the study period. The inter-regional deviation of phenological events was not large for any plant except for plum tree and Black locust. In addition, regional variations in high trophic levels of secondary consumers tended to be greater than that of producers and primary consumers. Among the studied species, plum was the most sensitive to temperature, and when the temperature rose by 1 °C, the flowering time of plum decreased by 7.20 days. As a result of checking the day differences in the phenological events of the interacting species, the phenological events of species were reversed, and butterflies have appeared earlier than plum, Korean forsythia, and Korean rosebay since 1990. Using long-term data from Korea, this study investigated differences in phenological reactions among trophic groups. There is a possibility of a phenological mismatch between trophic groups in the future if global warming continues due to differences in sensitivity to climate and phenological shifts between trophic levels.