Long-term temporal changes of plant phenology in the Western Mediterranean

Plants are altering their life cycles in response to current climatic change around the globe. More than 200 000 records for six phenological events (leaf unfolding, flowering, fruit ripening, fruit harvesting, leaf falling and growing season) of 29 perennial species for the period 1943–2003 recorded throughout Spain provide the longest temporal and the broadest spatial assessment of plant phenology changes in the Mediterranean region. The overwhelming majority of the 118 studied phenophases shifted their dates in recent decades. Such changes differed among phenological events. Leaf unfolding, flowering and fruiting are markedly advancing (?0.48, ?0.59 and ?0.32 days yr^?1, respectively), but only since the mid‐1970s. Anemophilous have advanced more days their flowering than entomophilous. However, some species have delayed and others have advanced their leaf falling dates and as a result only a weak shift was observed in this event for the whole of the studied species (+0.12 days yr^?1). The growing season lengthened by 18 days, which implies an increase of 8% in the life of annual leaves. Such an increase was achieved mainly through the advance of leaf unfolding dates in the spring, one of the most productive times of year for vegetation in the Mediterranean. Shifts in the plant calendar were accompanied as well by long‐term changes in the range of onset dates in 39% of studied phenophases. Leaf unfolding, flowering and growing season tended to reduce spatial variability, reflecting a faster and more synchronized onset (or duration) of phenophases across the study area. Changes in spatial variability may aggravate calendar mismatching with other trophic levels resulting from changes in dates. Because temporal responses differed markedly among species, calendar guilds of plants have changed, which suggests alterations of interspecific relationships in plant communities from Mediterranean ecosystems.     

气候波动对西安39种木本植物展叶始期及其积温需求的影响

过去几十年来暖春等异常气候事件发生的频次和强度显著增加, 使植物春季物候期发生了明显变化。但异常气候事件对植物春季物候积温需求的影响仍不清楚, 限制了对未来物候变化预测精度的提升。该研究利用西安植物园1963-2018年39种木本植物的展叶始期和相应气象数据, 首先根据3-4月平均气温划分了偏冷年、正常年和偏暖年, 对比了冷暖年相对于正常年的展叶始期变化。其次, 利用3种积温算法计算了各植物逐年的展叶始期积温需求, 比较了积温需求在冷暖年和正常年的差异。最后, 评估了传统积温模型在模拟偏冷或偏暖年展叶始期时的误差。结果表明, 所有植物的展叶始期在偏暖年比正常年平均早8.6天, 而在偏冷年平均晚8.2天。在偏暖年, 大多数物种展叶始期的积温需求(以5 ℃为阈值, 平均257.5度日)显著高于正常年(平均195.1度日); 在偏冷年的积温需求(平均168.0度日)低于正常年, 但在统计上差异不显著。就不同类群而言, 古老类群相对于年轻类群在偏冷年的推迟天数更多, 积温需求变化较小, 但在偏暖年无显著差异。不同生活型间物候与积温需求变化也无显著差异。造成偏暖年积温需求增加的可能原因是偏暖年冬季气温较高, 导致植物受到的冷激程度减轻, 从而抑制了后续的展叶。在正常年, 积温模型模拟木本植物展叶始期的平均误差仅为0.4-1.9天。在偏暖年和偏冷年, 模拟值分别比观测值平均早4.1天和晚3.0天。因此在预测未来物候变化时, 需要考虑气候波动条件下的积温需求变化。     

Effects of leaf emergence on leaf lifespan are independent of life form and successional status

Abstract The longevity of a leaf is related to the benefit that the plant is able to derive from it. This benefit varies among seasons and as more leaves emerge, such that leaf lifespan can be limited by canopy position rather than physiological age. Using interval‐censored failure time analysis, we investigate leaf lifespan for 34 Mediterranean species in a previously published dataset involving species with different life forms and functional strategies. Failure time regression models were used to determine leaf lifespan, and to investigate how these effects varied among species. Median lifespan estimated for each species with two methods differed by less than 10% on average, but varied from 0.02–19.5% depending on the shape of the underlying failure time distribution. Within shoots, later‐emerging leaves had shorter lifespans for species with longer periods of leaf emergence, and the reverse was true for species with short emergence. Having accounted for the within‐shoot effect, leaves emerging in spring had shorter lifespans, particularly herbaceous species, whereas the reverse was true woody species. These effects were consistent among life forms and successional stages, and consistent with theories of within‐shoot translocation of resources following self‐shading.     

Climatic change is advancing the phenology of moth species in Ireland

Recent increases in global temperatures have contributed to advancing phenology of plants and animals. These increases in temperature have been shown to affect the phenological phases (phenophases) of plants and birds in Ireland, but less is known about the effect on the phenophases of Irish insects. Records of the flight periods of 59 species of Irish moths over the past 35 years (1974–2009) were obtained from a public monitoring group. Observations were analysed across the country using generalized additive models (GAMs) weighted by total yearly population numbers for each species. The results of the statistical analyses showed that 45 of the 59 species studied have a significantly earlier first sighting date now than when observations began. With this earlier emergence, 44 of the 59 species also have a significantly longer flight season over the same 35‐year period. The extent of these changes varies across the country and by species life history. In particular, species emerging in spring are advancing at a much faster rate than species emerging during the summer. Many of these changes in first sighting are negatively correlated with rising temperatures in Ireland, particularly in late spring and early summer (May and June). The variation in phenological advancement in the moth species of Ireland is extremely complex and may be influenced more by species life history than by the phenology of interacting species, such as host plants.     

Evolutionary size changes in plants of the south‐west Pacific

Aim To investigate evolutionary changes in the size of leaves, stems and seeds of plants inhabiting isolated islands surrounding New Zealand. Location Antipodes, Auckland, Campbell, Chatham, Kermadec, Three Kings and Poor Knights Islands. Methods First, we compared the size of leaves and stems produced by 14 pairs of plant taxa between offshore islands and the New Zealand mainland, which were grown in a common garden to control for environmental effects. Similar comparisons of seed sizes were made between eight additional pairs of taxa. Second, we used herbarium specimens from 13 species pairs to investigate scaling relationships between leaf and stem sizes in an attempt to pinpoint which trait might be under selection. Third, we used herbarium specimens from 20 species to test whether changes in leaf size vary among islands located at different latitudes. Lastly, we compiled published records of plant heights to test whether insular species in the genus Hebe differed in size from their respective subgenera on the mainland. Results Although some evidence of dwarfism was observed, most insular taxa were larger than their mainland relatives. Leaf sizes scaled positively with stem diameters, with island taxa consistently producing larger leaves for any given stem size than mainland species. Leaf sizes also increased similarly among islands located at different latitudes. Size changes in insular Hebe species were unrelated to the average size of the respective subgenera on the mainland. Main conclusions Consistent evidence of gigantism was observed, suggesting that plants do not obey the island rule. Because our analyses were restricted to woody plants, results are also inconsistent with the ‘weeds‐to‐trees’ hypothesis. Disproportionate increases in leaf size relative to other plant traits suggest that selection may favour the evolution of larger leaves on islands, perhaps due to release from predation or increased intra‐specific competition.     

Phenology and climate change: a long-term study in a Mediterranean locality

It is well documented that plant and animal phenology is changing in response to recent climate warming in the Palaearctic. However, few long-term data sets are currently available in the Mediterranean basin. The present study reports long-term temporal trends of several phenophases of 45 plants, 4 insects and 6 migratory insectivorous birds. Dynamic factor analyses performed with plant phenophases showed that most of those events occurring at spring and summer had common trends toward the advancement, especially since mid-1970s. However, during these last decades, insect phenology showed a steeper advance than plant phenology, suggesting an increase of decoupling of some plant–insect interactions, such as those between pollinators and flowers or herbivorous insects and their plant resources. All trans-Saharan bird species showed highly significant temporal trends in all studied phenophases (some of them covering most of the last century). In two species, the duration of stay is increasing due to both earlier arrivals and later departures. On the other hand, two wintering species showed a significant advancement in their arrival dates, while an opposite pattern were found for departures of each one. Only one of these species increased significantly its wintering stay. Bird departures were not related to local climate in any species. Our results demonstrate a key role of local temperatures behind interannual variability of most plant and insects phenophases, with especial emphasis in those occurring in spring and summer. Therefore, the common signal towards the advancement recorded since mid-1970s resulted from the recent rise in temperatures.     

Phenology in central Europe – differences and trends of spring phenophases in urban and rural areas

In order to examine the impacts of both large-scale and small-scale climate changes (urban climate effect) on the development of plants, long-term observations of four spring phenophases from ten central European regions (Hamburg, Berlin, Cologne, Frankfurt, Munich, Prague, Vienna, Zurich, Basle and Chur) were analysed. The objective of this study was to identify and compare the differences in the starting dates of the pre-spring phenophases, the beginning of flowering of the snowdrop (Galanthus nivalis) and forsythia (Forsythia sp.), and of the full-spring phenophases, the beginning of flowering of the sweet cherry (Prunus avium) and apple (Malus domestica), in urban and rural areas. The results indicate that, despite regional differences, in nearly all cases the species studied flower earlier in urbanised areas than in the corresponding rural areas. The forcing in urban areas was about 4 days for the pre-spring phenophases and about 2 days for the full-spring phenophases. The analysis of trends for the period from 1951 to 1995 showed tendencies towards an earlier flowering in all regions, but only 22% were significant at the 5% level. The trends for the period from 1980 to 1995 were much stronger for all regions and phases: the pre-spring phenophases on average became earlier by 13.9 days/decade in the urban areas and 15.3 days/decade in the rural areas, while the full-spring phenophases were 6.7 days earlier/decade in the urban areas and 9.1 days/decade earlier in the rural areas. Thus rural areas showed a higher trend towards an earlier flowering than did urban areas for the period from 1980 to 1995. However, these trends, especially for the pre-spring phenophases, turned out to be extremely variable. Received: 21 October 1999 / Revised: 5 April 2000 / Accepted: 25 April 2000     

Different temperature perception in high‐elevation plants: new insight into phenological development and implications for climate change in the alpine tundra

In alpine habitats, predicted warmer and longer growing seasons will influence plant phenology, with important implications for species adaptation and vegetation dynamics. However, little is known on the temperature sensitivity of different phenophases and on the characteristics allowing phenological variation among and within species. By integrating interannual micro‐climatic variability with experimental warming, we explored how the phenology of three alpine species is influenced by temperature and what mechanisms underlie intra‐ and inter‐specific phenological differences. The present study demonstrated that alpine plants have different temperature responses during their reproductive cycle, do not have constant thermal thresholds and heat‐use efficiencies to achieve the seed dispersal stage and can change their temperature sensitivity to flower along snowmelt gradients. In addition, the length of the reproductive cycle, which proved to be species‐specific under experimental warming, does not seem to be the only life‐history trait under selective pressure due to the short‐length of the snow‐free period. In a warming climate scenario, the phenology of sexual reproduction will be considerably altered, and alpine plants may be subjected to changes in population dynamics driven by altered perception of environmental cues appropriate for coordinating the timing of key life‐history events.     

Is N‐resorption efficiency related to secondary compounds and leaf longevity in coexisting plant species of the arid Patagonian Monte,Argentina?

Leaf longevity and nutrient resorption efficiency are important strategies to conserve plant nutrients. Theory suggests a negative relationship between them and also proposes that high concentration of phenolics in long‐lived leaves may reduce nutrient resorption. In order to provide new evidence on these relationships, we explored whether N‐resorption efficiency is related to leaf longevity, secondary compounds and other leaf traits in coexisting plant species of different life forms in the arid Patagonian Monte, Argentina. We assessed N‐resorption efficiency, green leaf traits (leaf mass per area (LMA), leaf longevity and lignin, total soluble phenolics and N concentrations) and N concentration in senescent leaves of 12 species of different life forms (evergreen shrubs, deciduous shrubs and perennial grasses) with contrasting leaf traits. We found that leaf longevity was positively correlated to LMA and lignin, and negatively correlated to N concentration in green leaves. N concentrations both in green and senescent leaves were positively related. N‐resorption efficiency was not associated with the concentration of secondary compounds (total soluble phenolics and lignin) but it was negatively related to LMA and leaf longevity and positively related to N concentration in green leaves. Furthermore, leaf traits overlapped among life forms highlighting that life forms are not a good indicator of the functional properties (at least in relation to nutrient conservation) of species. In conclusion, our findings indicated that differences in N‐resorption efficiency among coexisting species were more related to N concentration in green leaves, leaf lifespan and LMA than to the presence of secondary compounds at least those assessed in our study (soluble phenolics and lignin). Accordingly, N‐resorption efficiency seems to be modulated, at least in part, by the productivity–persistence trade‐off.     

Impact of climate change on plant phenology in Mediterranean ecosystems

Plant phenology is strongly controlled by climate and has consequently become one of the most reliable bioindicators of ongoing climate change. We used a dataset of more than 200 000 records for six phenological events of 29 perennial plant species monitored from 1943 to 2003 for a comprehensive assessment of plant phenological responses to climate change in the Mediterranean region. Temperature, precipitation and North Atlantic Oscillation (NAO) were studied together during a complete annual cycle before phenological events to determine their relative importance and potential seasonal carry‐over effects. Warm and dry springs under a positive phase of NAO advance flowering, leaf unfolding and fruiting dates and lengthen the growing season. Spatial variability of dates (range among sites) was also reduced during warm and dry years, especially for spring events. Climate during previous weeks to phenophases occurrence had the greatest impact on plants, although all events were also affected by climate conditions several months before. Immediate along with delayed climate effects suggest dual triggers in plant phenology. Climatic models accounted for more than 80% of variability in flowering and leaf unfolding dates, and in length of the growing season, but for lower proportions in fruiting and leaf falling. Most part of year‐to‐year changes in dates was accounted for temperature, while precipitation and NAO accounted for <10% of dates' variability. In the case of flowering, insect‐pollinated species were better modelled by climate than wind‐pollinated species. Differences in temporal responses of plant phenology to recent climate change are due to differences in the sensitivity to climate among events and species. Spring events are changing more than autumn events as they are more sensitive to climate and are also undergoing the greatest alterations of climate relative to other seasons. In conclusion, climate change has shifted plant phenology in the Mediterranean region.     

Phenological response to climate change in China: a meta‐analysis

The change in the phenology of plants or animals reflects the response of living systems to climate change. Numerous studies have reported a consistent earlier spring phenophases in many parts of middle and high latitudes reflecting increasing temperatures with the exception of China. A systematic analysis of Chinese phenological response could complement the assessment of climate change impact for the whole Northern Hemisphere. Here, we analyze 1263 phenological time series (1960–2011, with 20+ years data) of 112 species extracted from 48 studies across 145 sites in China. Taxonomic groups include trees, shrubs, herbs, birds, amphibians and insects. Results demonstrate that 90.8% of the spring/summer phenophases time series show earlier trends and 69.0% of the autumn phenophases records show later trends. For spring/summer phenophases, the mean advance across all the taxonomic groups was 2.75 days decade^?1 ranging between 2.11 and 6.11 days decade^?1 for insects and amphibians, respectively. Herbs and amphibians show significantly stronger advancement than trees, shrubs and insect. The response of phenophases of different taxonomic groups in autumn is more complex: trees, shrubs, herbs and insects show a delay between 1.93 and 4.84 days decade^?1, while other groups reveal an advancement ranging from 1.10 to 2.11 days decade^?1. For woody plants (including trees and shrubs), the stronger shifts toward earlier spring/summer were detected from the data series starting from more recent decades (1980s–2000s). The geographic factors (latitude, longitude and altitude) could only explain 9% and 3% of the overall variance in spring/summer and autumn phenological trends, respectively. The rate of change in spring/summer phenophase of woody plants (1960s–2000s) generally matches measured local warming across 49 sites in China (R = ?0.33, P < 0.05).     

Broad-scale reciprocity in an avian seed dispersal mutualism

Aim Coevolved relationships between individual species of birds and plants rarely occur in seed dispersal mutualisms. This study evaluates whether reciprocal relationships may occur between assemblages of bird and plant species. Location Vancouver Island, British Columbia, Canada (48°50′‐N, 125°22′‐W). Methods The distribution and fruiting phenologies of seven shrub species were compared to seasonal changes in habitat selection and seed dispersal by six fruit‐eating bird species. Results Shrub species inhabiting forest understorey habitat had earlier fruiting phenologies than shrub species inhabiting forest edge habitat along lake and bog margins. Birds showed a parallel pattern in habitat selection, being more abundant in the forest understorey early in the fruiting season, and more abundant in the forest edge later in the season. Rates of seed deposition covaried with avian habitat selection, in such a way that birds directed seed dispersal into habitats preferred by shrubs. Conclusions These results depict a broad‐scale pattern in the abundance of birds and fruits indicative of reciprocal interactions. Seasonal changes in seed dispersal to each habitat appear to reinforce the relationship between shrub habitat affinities and fruiting phenologies. Phenological differences between habitats may also reinforce seasonal changes in avian habitat selection. Therefore, although reciprocal interactions between pairs of bird and plant species are rare, broad‐scale reciprocal relationships may occur between assemblages of bird and plant species.     

城市化对上海市城乡梯度上植物爆芽与展叶物候的影响

城市化影响植物物候,春季物候,如爆芽和展叶是植物响应城市化的客观生物指示。然而在城乡梯度上植物物候如何变化,以及物候与城市化程度是否存在定量关系,尚不明确。利用上海市2条城乡梯度上的6种木本植物的春季物候观测数据,以及距市中心距离作为城市化程度的代理指标,研究植物的爆芽和展叶物候期在城乡梯度上的变化及其与城市化程度的关系。研究结果表明：6种城市木本植物的爆芽和展叶物候期呈现随着距市中心距离的增加而延迟的趋势;城市化程度代理指标与爆芽和展叶物候期之间存在线性关系,城市化代理指标距市中心距离可以解释物候期延迟的31.5%—96.7%;物候期延迟时间因物种和距离城市距离而异,爆芽期延迟时间为3—15 d,展叶始期延长时间为4—13 d,展叶盛期延长时间为3—9 d。上述结果对理解城市化对植物物候的影响机制,评估城市化对城市生态系统的影响具有积极意义。     

Implications of climate change for North American wood warblers (Parulidae)

Since 1912, Neotropical–Nearctic migrant birds may now have up to 20 fewer days to travel between Southern Illinois, where spring is arriving later, and Northern Minnesota, where spring is arriving earlier, to exploit optimal habitat conditions (expanding leaves and caterpillar activity) for refueling and breeding. As case studies of the effect of climate change on bird migration, I analyzed two long‐term data sets of arrival times for eight species of northern breeding migratory wood warblers (Parulidae) gathered over a 100 year period in east‐central Illinois (IL, USA) and a 40 year period in western Minnesota (MN, USA). Six (IL) and seven (MN) of the wood warbler species showed no significant tendency to migrate earlier in response to earlier springs in their breeding range. These results suggest that climate change may force many species of long‐distance migratory songbirds to become uncoupled in the spring from their food resources that are driven by temperature.     

Possible impacts of climate change on natural vegetation in Saxony (Germany)

Recent climate changes have had distinct impacts on plant development in many parts of the world. Higher air temperatures, mainly since the end of the 1980s, have led to advanced timing of phenological phases and consequently to an extension of the general growing season. For this reason it is interesting to know how plants will respond to future climate change. In this study simple phenological models have been developed to estimate the impact of climate change on the natural vegetation in Saxony. The estimations are based on a regional climate scenario for the state of Saxony. The results indicate that changes in the timing of phenophases could continue in the future. Due to distinct temperature changes in winter and in summer, mainly the spring and summer phases will be advanced. Spring phenophases, such as leafing or flowering, show the strongest trends. Depending on the species, the average timing of these phenophases could be advanced by 3–27 days by 2050. Phenophases in autumn show relatively small changes. Thus, the annual growth period of individual trees will be further extended, mainly because of the shift of spring phases. Frequent droughts in summer and in autumn can compensate for the earlier leafing of trees, because in this case leaf colouring and leaf fall would start some weeks earlier. In such cases, the growing period would not be really extended, but shifted to the beginning of the year.     

Incidence of Fungal Necrotrophic and Biotrophic Pathogens in Pioneer and Shade‐tolerant Tropical Rain Forest Trees

This study assessed the hypothesis that plant life history traits determine the incidence of fungal biotrophic and necrotrophic pathogens in pioneer vs. shade‐tolerant tropical plant species. Considering that pioneer species mainly invest in induced defenses, we expected a negative relationship between the incidence of biotrophic and necrotrophic pathogens; in contrast, as shade‐tolerant species invest heavily in constitutive defenses, we expected to find no correlation between the incidence of biotrophic and necrotrophic pathogens. These ideas were evaluated by assessing standing levels of fungal damage in a set of pioneer and shade‐tolerant species from the Lacandona tropical rain forest (Mexico). The results showed that among pioneer plant species, leaves with biotrophic lesions were between 34 and 44 percent more abundant than those with necrotic lesions. In contrast, among shade‐tolerant species, the proportions of leaves with necrotic lesions were 17–23 percent higher than those of leaves with injuries caused by biotrophic pathogens. Our study suggests that tropical tree species might present different defense strategies depending on the life‐style of the pathogens that attack them, and the life history strategy of the attacked host plant species. Thus, the host constitutive and induced defenses, as well as the mechanisms used by different types of pathogens to circumvent those defenses maybe responsible for the patterns of attack observed in perennial tropical plants. Abstract in Spanish is available at http://www.blackwell‐synergy.com/loi/btp .     

Field evidence for earlier leaf-out dates in alpine grassland on the eastern Tibetan Plateau from 1990 to 2006

Worldwide, many plant species are experiencing an earlier onset of spring phenophases due to climate warming. Rapid recent temperature increases on the Tibetan Plateau (TP) have triggered changes in the spring phenology of the local vegetation. However, remote sensing studies of the land surface phenology have reached conflicting interpretations about green-up patterns observed on the TP since the mid-1990s. We investigated this issue using field phenological observations from 1990 to 2006, for 11 dominant plants on the TP at the levels of species, families (Gramineae—grasses and Cyperaceae—sedges) and vegetation communities (alpine meadow and alpine steppe). We found a significant trend of earlier leaf-out dates for one species (Koeleria cristata). The leaf-out dates of both Gramineae and Cyperaceae had advanced (the latter significantly, starting an average of 9 days later per year than the former), but the correlation between them was significant. The leaf-out dates of both vegetation communities also advanced, but the pattern was only significant in the alpine meadow. This study provides the first field evidence of advancement in spring leaf phenology on the TP and suggests that the phenology of the alpine steppe can differ from that of the alpine meadow. These findings will be useful for understanding ecosystem responses to climate change and for grassland management on the TP.     

Interacting effects of elevated CO2, nutrient availability and plant species on a generalist invertebrate herbivore

By affecting plant growth and phytochemistry elevated CO₂ may have indirect effects on the performance of herbivores. These effects show considerable variability across studies and may depend on nutrient availability, the carbon/nutrient‐balance in plant tissues and the secondary metabolism of plants. We studied the responses to elevated CO₂ and different nutrient availability of 12 herbaceous plant species differing in their investment into secondary compounds. Caterpillars of the generalist herbivore Spodoptera littoralis were reared on the leaves produced and their consumption and growth rates analysed. Elevated CO₂ resulted in a similar increase of biomass in all plant species, whereas the positive effect of fertilization varied among plant species. Specific leaf weight was influenced by elevated CO₂, but the effect depended on nutrient level and identity of plant species. Elevated CO₂ increased the C/N ratio of the leaves of most species. Caterpillars consumed more leaf material when plants were grown under elevated CO₂ and low nutrients. This indicates compensatory feeding due to lower tissue quality. However, the effects of elevated CO₂, nutrient availability and plant species identity on leaf consumption interacted. Both the effects of CO₂ and nutrient availability on the relative growth rate of the herbivore depended on the plant species. The feeding rate of S. littoralis on plant species that do not produce nitrogen‐containing secondary compounds (NCSC) was higher under low nutrient availability. In contrast, in plants producing NCSC nutrient availability had no effect on the feeding rate. This suggests that compensatory feeding in response to low nutrient contents may not be possible if plants produce NCSC. We conclude that elevated CO₂ causes species‐specific changes in the quality of plant tissues and consequently in changes in the preferences of herbivores for plant species. This could result in changes in plant community composition.     

A matter of timing: changes in the first date of arrival and last date of departure of Australian migratory birds

Although there is substantial evidence that Northern Hemisphere species have responded to climatic change over the last few decades, there is little documented evidence that Southern Hemisphere species have responded in the same way. Here, we report that Australian migratory birds have undergone changes in the first arrival date (FAD) and last date of departure (LDD) of a similar magnitude as species from the Northern Hemisphere. We compiled data on arrival and departure of migratory birds in south‐east Australia since 1960 from the published literature, Bird Observer Reports, and personal observations from bird watchers. Data on the FAD for 24 species and the LDD for 12 species were analyzed. Sixteen species were short‐ to middle‐distance species arriving at their breeding grounds, seven were long‐distance migrants arriving at their nonbreeding grounds, and one was a middle‐distance migrant also arriving at its nonbreeding ground. For 12 species, we gathered data from more than one location, enabling us to assess the consistency of intraspecific trends at different locations. Regressions of climate variables against year show that across south‐east Australia average annual maximum and minimum temperatures have increased by 0.17°C and 0.13°C decade^?1 since 1960, respectively. Over this period there has been an average advance in arrival of 3.5 days decade^?1; 16 of the 45 time‐series (representing 12 of the 24 species studied) showed a significant trend toward earlier arrival, while only one time‐series showed a significant delay. Conversely, there has been an average delay in departure of 5.1 days decade^?1; four of the 21 departure time‐series (four species) showed a significant trend toward later departure, while one species showed a significant trend toward earlier departure. However, differences emerge between the arrival and departure of short‐ to middle‐distance species visiting south‐east Australia to breed compared with long‐distance species that spend their nonbreeding period here. On average, short‐ to middle‐distance migrants have arrived at their breeding grounds 3.1 days decade^?1 earlier and delayed departure by 8.1 days decade^?1, thus extending the time spent in their breeding grounds by ～11 days decade^?1. The average advance in arrival at the nonbreeding grounds of long‐distance migrants is 6.8 days decade^?1. These species, however, have also advanced departure by an average of 6.9 days decade^?1. Hence, the length of stay has not changed but rather, the timing of events has advanced. The patterns of change in FAD and LDD of Australian migratory birds are of a similar magnitude to changes undergone by Northern Hemisphere species, and add further evidence that the modest warming experienced over the past few decades has already had significant biological impacts on a global scale.     

Long-term breeding phenology shift in royal penguins

The Earth's climate is undergoing rapid warming, unprecedented in recent times, which is driving shifts in the distribution and phenology of many plants and animals. Quantifying changes in breeding phenology is important for understanding how populations respond to these changes. While data on shifts in phenology are common for Northern Hemisphere species (especially birds), there is a dearth of evidence from the Southern Hemisphere, and even fewer data available from the marine environment. Surface air temperatures at Macquarie Island have increased by 0.62°C during the 30-year study period (0.21°C decade(-1)) and royal penguins (Eudyptes schlegeli) commenced egg laying on average three days earlier in the 1990s than during the 1960s. This contrasts with other studies of Southern Ocean seabirds; five of nine species are now breeding on average 2.1 days later than during the 1950s. Despite the different direction of these trends, they can be explained by a single underlying mechanism: resource availability. There was a negative relationship between the Southern Annular Mode (SAM) and median laying date of royal penguins, such that low-productivity (low SAM) years delayed laying date. This accords with the observations of other seabird species from the Antarctic, where later laying dates were associated with lower sea ice and lower spring productivity. The unifying factor underpinning phenological trends in eastern Antarctica is therefore resource availability; as food becomes scarcer, birds breed later. These changes are not uniform across the region, however, with resource increases in the subantarctic and decreases in eastern Antarctica.