Phenological Cues Research Articles

Lagging spawning and increasing phenological extremes jeopardize walleye (Sander vitreus) in north‐temperate lakes

AbstractThe phenology of critical biological events in aquatic ecosystems is rapidly shifting due to climate change. Growing variability in phenological cues can increase the likelihood of trophic mismatches (i.e., mismatches in the timing of peak prey and predator abundances), causing recruitment failures in important fisheries. We assessed changes in the spawning phenology of walleye (Sander vitreus) in 194 Midwest US lakes to investigate factors influencing walleye phenological responses to climate change and associated climate variability, including ice‐off timing, lake physical characteristics, and population stocking history. Ice‐off phenology shifted earlier, about three times faster than walleye spawning phenology over time. Spawning phenology deviations from historic averages increased in magnitude over time, and large deviations were associated with poor offspring survival. Our results foreshadow the risks of increasingly frequent natural recruitment failures due to mismatches between historically tightly coupled spawning and ice‐off phenology.

Adaptive photoperiod interpretation modulates phenological timing in Atlantic salmon

Photoperiod, the portion of 24-h cycle during which an organism is exposed to illumination, is an important phenological cue in many animals. However, despite its influence on critical biological processes, there remain many unknowns regarding how variations in light intensity translate into perceived photoperiod. This experiment examined how light intensity variations affect perceived photoperiod in Atlantic salmon (Salmo salar) to determine whether photoperiod interpretation is, a) fixed such that anything above a minimum detection threshold is regarded as ‘illumination’, or b) adaptive and varies with recent light exposure. To do this we compared the frequency of smoltification and sexual maturation between groups of male parr which were exposed to one of eight light regimes on a 12:12 cycling regime (12-hour day/12-hour night). The eight regimes were divided into two treatments, four with ‘High’ daytime light intensity and four with ‘Low’ daytime light intensity. The ‘High' and ‘Low' intensity treatments were each sub-divided into four groups for which the subjective ‘night’ light intensity was 100%, 10%, 1% and 0% of the daytime light intensity, with four replicate tanks of each treatment. The results show that above a minimum detection threshold, Atlantic salmon have adaptive photoperiod interpretation which varies with recent light exposure, and that adaptive photoperiod interpretation modulates the timing of the parr-smolt transformation and sexual maturation. Further, we show that photoperiod interpretation varies between closely related families. Given the influence of phenological timing on species survival, our results reveal a critical role for integration of photoperiod interpretation in attempts to understand how geographically shifting thermal niches due to climate change will affect future populations.

Primed and cued: long-term acoustic telemetry links interannual and seasonal variations in freshwater flows to the spawning migrations of Common Snook in the Florida Everglades

BackgroundSpawning migrations are a widespread phenomenon among fishes, often occurring in response to environmental conditions prompting movement into reproductive habitats (migratory cues). However, for many species, individual fish may choose not to migrate, and research suggests that conditions preceding the spawning season (migratory primers) may influence this decision. Few studies have provided empirical descriptions of these prior conditions, partly due to a lack of long-term data allowing for robust multi-year comparisons. To investigate how primers and cues interact to shape the spawning migrations of coastal fishes, we use acoustic telemetry data from Common Snook (Centropomus undecimalis) in Everglades National Park, Florida, USA. A contingent of Snook migrate between rivers and coastal spawning sites, varying annually in both the proportion of the population that migrates and the timing of migration within the spawning season. However, the specific environmental factors that serve as migratory primers and cues remain unknown.MethodsWe used eight years of acoustic telemetry data (2012–2019) from 173 tagged Common Snook to investigate how primers and cues influence migratory patterns at different temporal scales. We hypothesize that (1) interannual differences in hydrologic conditions preceding the spawning season contribute to the number of individuals migrating each year, and (2) specific environmental cues trigger the timing of migrations during the spawning season. We used GLMMs to model both the annual and seasonal migratory response in relation to flow characteristics (water level, rate of change in water level), other hydrologic/abiotic conditions (temperature, salinity), fish size, and phenological cues independent of riverine conditions (photoperiod, lunar cycle).ResultsWe found that the extent of minimum marsh water level prior to migration and fish size influence the proportion of Snook migrating each year, and that high river water level and daily rates of change serve as primary cues triggering migration timing.ConclusionOur findings illustrate how spawning migrations are shaped by environmental factors acting at different temporal scales and emphasize the importance of long-term movement data in understanding these patterns. Research providing mechanistic descriptions of conditions that promote migration and reproduction can help inform management decisions aimed at conserving ecologically and economically important species.

Host plant phenology shapes aphid abundance and interactions with ants

Phenological mismatch can occur when plants and herbivores differentially respond to changing phenological cues, such as temperature or snow melt date. This often shifts herbivore feeding to plant stages of lower quality. How herbivores respond to plant quality may be also mediated by temperature, which could lead to temperature‐by‐phenology interactions. We examined how aphid abundance and mutualism with ants were impacted by temperature and host plant phenology. In this study system, aphids Aphis asclepiadis colonize flowering stalks of the host plant, Ligusticum porteri. Like other aphids, abundance of this species is dependent on ant protection. To understand how host plant phenology and temperature affect aphid abundance, we used a multiyear observational study and a field experiment. We observed 20 host plant populations over five years (2017–2021), tracking temperature and snow melt date as well as host plant phenology and insect abundance. We found host plant and aphid phenology to differentially respond to temperature and snow melt timing. Early snow melt accelerated host plant phenology to a greater extent than aphid phenology, which was more responsive to temperature. Both the likelihood of aphid colony establishment and ant recruitment were reduced when aphids colonized host plants at post‐flowering stages. In 2019, we experimentally accelerated host plant phenology by advancing snow melt date by two weeks. We factorially combined this treatment with open top warming chambers surrounding aphid colonies. Greatest growth occurred for colonies under ambient temperatures when they occurred on host plants at the flowering stage. Altogether, our results suggest that phenological mismatch with host plants can decrease aphid abundance, and this effect is exacerbated by temperature increases and changes to the ant–aphid mutualism.

Integrating experiments to predict interactive cue effects on spring phenology with warming.

Climate change has advanced plant phenology globally 4-6 d °C-1 on average. Such shifts are some of the most reported and predictable biological impacts of rising temperatures. Yet as climate change has marched on, phenological shifts have appeared muted over recent decades - failing to match simple predictions of an advancing spring with continued warming. The main hypothesis for these changing trends is that interactions between spring phenological cues - long-documented in laboratory environments - are playing a greater role in natural environments due to climate change. Here, we argue that accurately linking shifts observed in long-term data to underlying phenological cues is slowed by biases in observational studies and limited integration of insights from laboratory studies. We synthesize seven decades of laboratory experiments to quantify how phenological cue-space has been studied and how treatments compare with shifts caused by climate change. Most studies focus on one cue, limiting our ability to make accurate predictions, but some well-studied forest species offer opportunities to advance forecasting. We outline how greater integration of controlled-environment studies with long-term data could drive a new generation of laboratory experiments, built on physiological insights, that would transform our fundamental understanding of phenology and improve predictions.

Phenological cues to breeding and the differential response of Pacific auks to variation in marine productivity

Many bird species use features of the physical environment to cue breeding activity. We show that for 2 species of Pacific auks (zooplanktivorous Cassin’s auklet Ptychoramphus aleuticus and generalist rhinoceros auklet Cerorhinca monocerata), spatio-temporal variation in marine production indicators surrounding a major breeding colony in the northeast Pacific Ocean can cue laying date. By utilizing a multi-year phenological time series and a spatio-temporal sliding-window analysis spanning November up until median lay dates in spring, chlorophyll a (chl a) concentrations in the surface ocean around the colony strongly predicted lay date. However, the response to this cue differed between species by over 2 mo, as each species was exposed to different, sequential water-masses. We show that for Cassin’s auklets, chl a levels at Triangle Island, British Columbia (Canada) in February, nearly 2 mo prior to actual laying, strongly correlated with lay date in April (average r = 0.83). At this time, the ocean environment around Triangle Island is dominated by water-masses from the south-southeast. However, for sympatric rhinoceros auklets, chl a along broad shelf-break areas in April prior to lay dates in May strongly correlated with lay date (average r = 0.76). Occurring after the spring transition, these water-masses flow primarily from the northwest. Consistent with other bird species, both auks appear to respond to information about food production, but at different spatio-temporal scales. We suggest that these different responses to environmental cues reflect species-specific differences in female migration behaviour, and an attempt by Cassin’s auklets to bet-hedge against phenological mismatches.

From the ground up: Building predictions for how climate change will affect belowground mutualisms, floral traits, and bee behavior

Climate change affects species and their interactions, resulting in novel communities and modified ecosystem processes. Through shifts in phenology and distribution, climatic change can disrupt interactions, including those between mutualists. Mutualisms influence the structure and stability of communities and can link species to a common fate. However, research on climate change has focused on pairwise mutualisms, neglecting the higher-order interactions that can arise when species interact with multiple mutualists. We explore the effects of climate change on tripartite interactions involving belowground mutualists, namely soil bacteria and fungi, flowering plants, and pollinators. We outline how climate change is predicted to affect the phenology and distribution of these belowground mutualists, emphasizing the consequent effects on host plant floral traits, plant-pollinator interactions, and bee behavior. We find evidence that warming, advanced snowmelt, and drought are likely to cause phenological and distributional shifts in soil microbes, leading to diminished mutualistic interactions with plants and symbiont switching. Consequently, shifts in flowering phenology, smaller floral displays, and lower quality floral rewards are expected, increasing foraging time and energy demands for bees and altering their floral preferences. Such costs could translate into reduced fitness and novel selection pressures for bees and flowering plants in the short term. We highlight knowledge gaps and ways forward, urging studies on microbe dispersal and phenological cues, experiments that manipulate soil microbe-host plant interactions under simulated climate change conditions, and large-scale field studies across environmental gradients, all with the goal of understanding how climate change will affect soil microbe-plant-pollinator mutualisms.

Does Phenological Plasticity Help or Hinder Range Shifts Under Climate Change?

Climate warming is predicted to shift species’ ranges as previously uninhabitable environments just beyond the leading range edges become suitable habitat and trailing range edges become increasingly unsuitable. Understanding which aspects of the environment and species traits mediate these range shifts is critical for understanding species’ possible redistributions under global change, yet we have a limited understanding of the ecological and evolutionary responses underlying population spread or extinction at species’ range edges. Within plant populations, shifts in flowering phenology have been one of the strongest and most consistent responses to climate change, and are likely to play an important role in mediating population dynamics within and beyond species’ ranges. However, the role of phenological shifts, and particularly phenological plasticity, in species’ range shifts remains relatively unstudied. Here, we synthesize literature on phenology, plasticity, and adaptation to suggest ways in which phenological responses to climate may vary across species’ ranges and review the empirical evidence for and against these hypotheses. We then outline how phenological plasticity could facilitate or hinder persistence and potential consequences of phenological plasticity in range expansions, including phenological cues, shifts in correlated traits, altered species interactions, and effects on gene flow. Finally, we suggest future avenues for research, such as characterizing reaction norms for phenology across a species’ range and in beyond-the-range transplant experiments. Given the prevalence and magnitude of phenological shifts, future work should carefully dissect its costs and benefits for population persistence, and incorporate phenological plasticity into models predicting species’ persistence and geographic range shifts under climate change.

Winter temperatures predominate in spring phenological responses to warming

Research on woody plant species highlights three major cues that shape spring phenological events: chilling, forcing and photoperiod. Increasing research on the phenological impacts of climate change has led to debate over whether chilling and/or photoperiod cues have slowed phenological responses to warming in recent years. Here we use a global meta-analysis of all published experiments to test the relative effects of these cues. Almost all species show strong responses to all three cues, with chilling being the strongest and photoperiod the weakest. Forecasts from our findings for Central Europe suggest that spring phenology will continue to advance, as stalling effects of chilling generally appear above 4 °C warming in this region. Our results unify both sides of the debate over phenological cues: while all species may respond to all cues strongly in experimental conditions, in current environmental conditions the dominant signal of climate change is from increased forcing. Spring phenology is influenced by chilling, forcing and photoperiod cues; the phenological response to warming from anthropogenic climate change may be slowed by chilling or photoperiod. Plant species respond to all cues in experiments but under environmental conditions, forcing predominates.

What drives phenological synchrony? Warm springs advance and desynchronize flowering in oaks

Annually variable and synchronous seed production, or masting, is often correlated with environmental factors and in oaks involves differential pollination success that depends on phenological synchrony in flowering. The synchronization of phenology of flowering was thought to be driven by temperature during flowering (microclimatic hypothesis). We tested an alternative, whereby phenological synchronization is driven by the timing of the onset of flowering (photoperiod-sensitivity hypothesis). This hypothesis assumes that flowering synchrony is driven by interaction between daylength and temperature, and individual variation in sensitivity to daylength as a phenological cue. We used long-term (23–26 years) records of airborne pollen in Quercus robur, Q. petraea, Q. ilex, and Q. humilis. Late pollen seasons were short, as predicted by photoperiod-sensitivity hypothesis. The onset of pollen seasons was delayed as preseason temperatures cooled over the last three decades at our Mediterranean sites, which was paralleled by shortening in pollen seasons, providing additional support for the photoperiod-sensitivity hypothesis. Global warming under the microclimatic hypothesis is predicted to lead to less frequent reproductive failures and thus decreased variability and synchrony of mast seeding. In contrast, warming under the photoperiod-sensitivity hypothesis should advance the onset of and desynchronize flowering, a pattern supported by our data. This pattern suggests that global warming will lead to more frequent vetoes and more stochastic and variable patterns of oak reproduction.

Advanced phenology of intraguild predators shifts herbivore host plant preference and performance

1. The abundance of insect herbivores is mediated by interactions with higher and lower trophic levels. This research asks (i) how phenological change across trophic levels affects host plant quality and selection for aphids, and (ii) what higher trophic level mechanisms drive aphid abundance.2. Ligusticum porteri is a perennial host for the sap‐feeding aphid Aphis asclepiadis and intraguild mirid predators (chiefly Lygus hesperus) in Colorado. We used long‐term observational data to discover that aphids and mirids respond differently to phenological cues. These unique responses can impact aphid abundance through changes to host plant selection and quality.3. We used behavioural choice assays to assess how advanced mirid phenology influences aphid host plant selection. More alates landed and reproduced on mirid‐free control plants relative to host plants with prior mirid feeding. However, this preference did not correlate with aphid performance when we compared aphid relative growth rates between treatments. This suggests that advanced mirid phenology would impact aphid populations more through host plant choice, rather than reductions in host quality. The addition of mirids to experimental aphid colonies also demonstrated reduced aphid colony growth via predation.4. We measured plant cues involved in host selection and found differences in volatile composition between plants with prior mirid feeding compared to control plants, providing the potential for aphids to detect enemy‐free space using volatile cues.

Tactical departures and strategic arrivals: Divergent effects of climate and weather on caribou spring migrations

AbstractThe Arctic has been warming rapidly, affecting ecological processes across the region. Caribou and reindeer (Rangifer tarandus) is a keystone Arctic species undergoing declines in many parts of its range, but definitive links between climate and populations remain elusive. The conspicuous and dramatic mass migration of many caribou populations, during which nearly all pregnant females move from wintering ranges to calving grounds shortly before giving birth, may be an important link between climate and caribou populations. The drivers of migration, however, are similarly mysterious. It is unknown, for example, whether caribou respond to immediate phenological cues, anticipate conditions on calving grounds, or are driven by lagged effects related to physical condition. To investigate the drivers of migration, we analyzed movement data from over 1000 individual caribou from seven major herds, spanning 3000 km across Alaska, Yukon, Northwest Territories (NWT), and Nunavut in Canada, from 1995 to 2017. We developed a hierarchical model to estimate migration departure and arrival times, and analyzed these variables against global climate indices and local weather conditions, exploring immediate and lagged effects, as well as snowmelt timing and vegetation indices. We discovered a continent‐wide synchrony in spring migration departure times, driven mainly by large‐scale, ocean‐driven climate indices (Pacific Decadal Oscillation, Arctic Oscillation, and North Atlantic Oscillation). However, we also found that the speed of migration was highly plastic with later migration departure times followed by shorter migration durations. This plasticity made arrival timing independent of departure timing and its respective drivers. Rather, arrival timing depended strongly on weather conditions from the previous summer: cooler and windier summers generally led to earlier arrival at calving grounds the following year. We suggest that maternal body condition, mainly influenced by conditions that limit insect harassment, is a major factor for earlier spring migration arrival timing, and therefore earlier calving and higher survival rates. We place these results in the context of mechanistic links between climate and caribou population dynamics. Long‐term and large‐scale observations of migratory animals can provide insights into the mechanisms by which long‐distance, collective migrants may adapt to dynamic and unpredictable environments.

The influence of spectral composition on spring and autumn phenology in trees.

Several recent reviews highlight the molecular mechanisms that underpin phenological responses to temperature and photoperiod; however, these have mostly overlooked the influence of solar radiation and its spectral composition on these processes. For instance, solar radiation in the blue and ultraviolet (UV) regions of the spectrum, as well as the red/far-red (R:FR) ratio, can influence spring and autumn phenology. Solar radiation reaching the Earth changes diurnally and seasonally; however, rising global temperatures, latitudinal range shifts and light pollution are likely to produce novel combinations of phenological cues for tree species. Here, we review the literature on phenological responses to spectral composition. Our objective was to explore the natural variation in spectral composition using radiative transfer models and to reveal any species-specific or ecotype-specific responses relating to latitudinal origin. These responses are likely to be most pronounced at high latitudes where spectral composition varies most throughout the year. For instance, trees from high latitudes tend to be more sensitive to changes in R:FR than those from low latitudes. The effects of blue light and UV radiation on phenology have not been studied as much as those of R:FR, but the limited results available suggest both could be candidate cues affecting autumn leaf colouration and senescence. Failure of more-southern species and ecotypes to adapt and use spectral cues during northwards range shifts could result in mistimed phenology, potentially resulting in frost damage, reduced fitness and limited range expansion. Future areas for research should look to establish how consistently different functional types of tree respond to spectral cues and identify photoreceptor-mediated mechanisms that allow plants to combine information from multiple light cues to coordinate the timing of phenological events. It should then be feasible to consider the synchronous or sequential action of light cues within a hierarchy of environmental factors regulating phenology.

Breeding phenology, provisioning behaviour, and unusual patterns of life history variation across an anthropogenic heterogeneous landscape.

Understanding how resource use and life history variation influence a population's response to modified, fragmented landscapes is a major challenge for ecologists. We investigated the phenology, life history decisions and provisioning behaviour of a generalist passerine-the great tit-across a heavily managed woodland landscape. Contrary to most previous studies on this species, reproductive investment and success were lower in deciduous than in coniferous woodland fragments. This could not be explained by differences in provisioning behaviour; instead population density was considerably higher in deciduous woodlands, suggesting birds did not follow an ideal free distribution. Clutch size declined with lay date amongst populations breeding in coniferous woodland fragments, but these populations also displayed pronounced seasonal declines in the proportion of fledglings produced per egg and fledgling mass. In contrast, and against patterns observed in other similar study systems, clutch size did not change with lay date in mixed-species deciduous woodland fragments. Furthermore, the proportion of young fledged and fledgling condition remained stable throughout the season, even though the quality of food provisioned to nestlings increased over the season. Local recruitment was negligible, suggesting that plasticity rather than natural selection played a key role in driving the patterns observed. The unusual patterns we report are likely explained by the fragmented nature of the landscape, and unreliable phenological cues in a mixed-species tree community coupled with low food availability. They contrast with those reported from most other populations situated in continuous woodland, demonstrating that caution is needed when generalising across different contexts and ecosystems.

Identifying phenological functional types in savanna trees

Savannas are notable for the varied patterns of leaf display in the tree layer. Characterising this variation, and assessing the range of phenological strategies apparent in savanna ecosystems will be helpful for understanding savanna tree life-histories and their response to global change. We recorded leaf phenological metrics on a monthly basis for 28 woody species for one year in a semi-arid savanna at Nylsvley Nature Reserve, South Africa. Using these metrics four plant functional types were described: (1) evergreen species, which retained full canopy throughout the year; (2) semi-evergreen species, which lost a significant amount of canopy during the dry season, but not less than 60%; (3) obligate deciduous species in which the canopy was leafless for about 3 months during the dry season; and (4) rain-stimulated species, in which canopy flushing was driven by rainfall patterns. Of the 28 species, 18% were evergreen, 18% obligate deciduous, 32% semi-evergreen and 32% were rain-stimulated species. These results confirm that phenological strategies are variable in savannas, and highlight that there are numerous cues controlling the timing of leaf out and leaf drop. By defining distinct phenological strategies we can predict which plant functional types will have phenological cues that will be vulnerable to shifting climates.

Phenological cues intrinsic in indigenous knowledge systems for forecasting seasonal climate in the Delta State of Nigeria.

Shifts in the timing of phenological events in plants and animals are cited as one of the most robust bioindicators of climate change. Much effort has thus been placed on the collection of phenological datasets, the quantification of the rates of phenological shifts and the association of these shifts with recorded meteorological data. These outputs are of value both in tracking the severity of climate change and in facilitating more robust management approaches in forestry and agriculture to changing climatic conditions. However, such an approach requires meteorological and phenological records spanning multiple decades. For communities in the Delta State of Nigeria, small-scale farming communities do not have access to meteorological records, and the dissemination of government issued daily to seasonal forecasts has only taken place in recent years. Their ability to survive inter-annual to inter-decadal climatic variability and longer-term climatic change has thus relied on well-entrenched indigenous knowledge systems (IKS). An analysis of the environmental cues that are used to infer the timing and amount of rainfall by farmers from three communities in the Delta State reveals a reliance on phenological events, including the croaking of frogs, the appearance of red millipedes and the emergence of fresh rubber tree and cassava leaves. These represent the first recorded awareness of phenology within the Delta State of Nigeria, and a potentially valuable source of phenological data. However, the reliance of these indicators is of concern given the rapid phenological shifts occurring in response to climate change.

Phylogenetic conservatism and trait correlates of spring phenological responses to climate change in northeast China.

Climate change has resulted in major changes in plant phenology across the globe that includes leaf‐out date and flowering time. The ability of species to respond to climate change, in part, depends on their response to climate as a phenological cue in general. Species that are not phenologically responsive may suffer in the face of continued climate change. Comparative studies of phenology have found phylogeny to be a reliable predictor of mean leaf‐out date and flowering time at both the local and global scales. This is less true for flowering time response (i.e., the correlation between phenological timing and climate factors), while no study to date has explored whether the response of leaf‐out date to climate factors exhibits phylogenetic signal. We used a 52‐year observational phenological dataset for 52 woody species from the Forest Botanical Garden of Heilongjiang Province, China, to test phylogenetic signal in leaf‐out date and flowering time, as well as, the response of these two phenological traits to both temperature and winter precipitation. Leaf‐out date and flowering time were significantly responsive to temperature for most species, advancing, on average, 3.11 and 2.87 day/°C, respectively. Both leaf‐out and flowering, and their responses to temperature exhibited significant phylogenetic signals. The response of leaf‐out date to precipitation exhibited no phylogenetic signal, while flowering time response to precipitation did. Native species tended to have a weaker flowering response to temperature than non‐native species. Earlier leaf‐out species tended to have a greater response to winter precipitation. This study is the first to assess phylogenetic signal of leaf‐out response to climate change, which suggests, that climate change has the potential to shape the plant communities, not only through flowering sensitivity, but also through leaf‐out sensitivity.

Experimentally increased prebreeding male social behaviour has no effect on female breeding phenology and performance

In the context of breeding phenology, social behaviour (especially song) has long been considered an important ‘supplemental cue’ that females integrate with other environmental information to determine timing of egg laying. However, to our knowledge, no studies have experimentally manipulated song in the wild in the context of female breeding phenology and performance. We studied natural variation in, and response to experimental manipulation of, prebreeding song and social behaviour in European starlings, Sturnus vulgaris, to determine whether male behaviour acts as a phenological cue determining female timing of egg laying and subsequent, postlaying breeding performance. In our highly synchronous system, natural variation in prebreeding male song quality and singing effort was surprisingly high, and singing increased slightly closer to egg-laying date. There was a strong prebreeding response of both males and females to the playback treatment, but no effect on female breeding phenology (egg-laying date) or performance (nestbox occupancy, egg size, clutch size, total chicks hatched, total chicks fledged or double-brooding behaviour). Our 3-year study finds no evidence that male prebreeding song is a cue determining female breeding phenology or performance in starlings.

Passerines may be sufficiently plastic to track temperature-mediated shifts in optimum lay date.

Projecting the fates of populations under climate change is one of global change biology's foremost challenges. Here, we seek to identify the contributions that temperature-mediated local adaptation and plasticity make to spatial variation in nesting phenology, a phenotypic trait showing strong responses to warming. We apply a mixed modeling framework to a Britain-wide spatiotemporal dataset comprising >100000 records of first egg dates from four single-brooded passerine bird species. The average temperature during a specific time period (sliding window) strongly predicts spatiotemporal variation in lay date. All four species exhibit phenological plasticity, advancing lay date by 2-5days°C(-1) . The initiation of this sliding window is delayed further north, which may be a response to a photoperiod threshold. Using clinal trends in phenology and temperature, we are able to estimate the temperature sensitivity of selection on lay date (B), but our estimates are highly sensitive to the temporal position of the sliding window. If the sliding window is of fixed duration with a start date determined by photoperiod, we find B is tracked by phenotypic plasticity. If, instead, we allow the start and duration of the sliding window to change with latitude, we find plasticity does not track B, although in this case, at odds with theoretical expectations, our estimates of B differ across latitude vs. longitude. We argue that a model combining photoperiod and mean temperature is most consistent with current understanding of phenological cues in passerines, the results from which suggest that each species could respond to projected increases in spring temperatures through plasticity alone. However, our estimates of B require further validation.

Phenological dynamics of the invasive plant Acacia longifolia in Portugal

SummaryPhenological shifts may play an important role in the success of invasive species. In the coastal regions of continental Portugal, Acacia longifolia is one of the most widespread invasive plants, but there are significant gaps in our understanding and interpretation of its phenology. This study is the first to investigate the phenological variation of A. longifolia and its association with climate in populations representing a temperature and precipitation gradient in mainland Portugal. The results highlighted significant variations in the vegetative (leaf initiation) and reproductive phenophases (bud formation, flowering and fruiting) among populations. Overall, leafing was observed throughout the year, bud formation occurred between summer and spring, anthesis between late winter and the beginning of spring, and fruiting started in late winter. Most of the reproductive phenophases varied across elevation or latitudinal gradients, with populations that were subjected to more stressful climatic conditions advancing their flowering and fruiting phenophases. The occurrence of each phenophase was strongly associated with the climatic conditions in the previous 5–12 months, suggesting that plants receive their phenological cues well in advance of their phenological response. Among climatic factors, temperature and irradiance were the most significant predictors of the phenological cycle of A. longifolia, but precipitation also exerted a greater influence on budding and flowering of the species. The phenological response of A. longifolia in a variety of sites represents an important step towards understanding how its phenology may react to the projected climate change in the Mediterranean region.