Comparison of cold tolerance in eggs of two cicadas, Cryptotympana facialis and Graptopsaltria nigrofuscata, in relation to climate warming

The population of the cicada Cryptotympana facialis began to increase in Osaka, Japan, during the late 20th century. Climate warming is considered a major cause, although the relationship between temperature and the cicada population increase remains unclear. By examining cold tolerance in overwintering eggs of C. facialis in relation to another cicada, Graptopsaltria nigrofuscata , whose population has recently decreased in Osaka, we tested the hypothesis that warming has caused the population increase of C. facialis by decreasing egg mortality due to winter temperatures. A short-term (24 h) cold exposure experiment demonstrated that the half-lethal temperatures (LT50) of C. facialis and G. nigrofuscata were −23.3°C and −28.9°C, respectively, although these extreme low temperatures never occurred in Osaka during the 20th century. Prolonged exposure to −5°C for up to 30 days had no harmful effects on the hatching rate in either species. Overwintering mortality was also assessed under naturally fluctuating conditions by transferring eggs to cooler elevated sites that mimicked the environment prior to the current warming. Eggs of C. facialis that overwintered at the cooler site exhibited similar hatching rates to those maintained at the original site. The results of these experiments consistently indicated that overwintering eggs of C. facialis possess adequate tolerance to the low temperatures of the 20th century. Therefore, we rejected our initial hypothesis that recent increases in the C. facialis population have been caused by warming-related reductions in overwintering egg mortality.     

Two sides of a coin: host‐plant synchrony fitness trade‐offs in the population dynamics of the western spruce budworm

Conifer‐feeding budworms emerge from overwintering sites as small larvae in early spring, several days before budburst, and mine old needles. These early‐emerging larvae suffer considerable mortality during this foraging period as they disperse in search of available, current‐year buds. Once buds flush, surviving budworms construct feeding shelters and must complete maturation before fresh host foliage senesces and lignifies later in the summer. Late‐developing larvae suffer greater mortality and survivors have lower fecundity when feeding on older foliage. Thus, there is a seasonal trade‐off in fitness associated with host synchrony: early‐emerging budworms have a greater risk of mortality during spring dispersal but gain better access to the most nutritious foliage, while, on the other hand, late‐emerging larvae incur a lower risk during the initial foraging period but must contend with rapidly diminishing resource quality at the end of the feeding period. We investigate the balance that results from these early‐season and late‐season synchrony fitness trade‐offs using the concept of the phenological window. Parameters associated with the variation in the phenological window are used to estimate generational fitness as a function of host‐plant synchrony. Because defoliation modifies these relationships, it is also included in the analysis. We show that fitness trade‐offs characterizing the phenological window result in a robust synchrony relationship between budworm and host plant over a wide geographic range in southern British Columbia, Canada.     

Warming counteracts defoliation‐induced mismatch by increasing herbivore‐plant phenological synchrony

Climate change is altering phenology; however, the magnitude of this change varies among taxa. Compared with phenological mismatch between plants and herbivores, synchronization due to climate has been less explored, despite its potential implications for trophic interactions. The earlier budburst induced by defoliation is a phenological strategy for plants against herbivores. Here, we tested whether warming can counteract defoliation‐induced mismatch by increasing herbivore‐plant phenological synchrony. We compared the larval phenology of spruce budworm and budburst in balsam fir, black spruce, and white spruce saplings subjected to defoliation in a controlled environment at temperatures of 12, 17, and 22°C. Budburst in defoliated saplings occurred 6–24 days earlier than in the controls, thus mismatching needle development from larval feeding. This mismatch decreased to only 3–7 days, however, when temperatures warmed by 5 and 10°C, leading to a resynchronization of the host with spruce budworm larvae. The increasing synchrony under warming counteracts the defoliation‐induced mismatch, disrupting trophic interactions and energy flow between forest ecosystem and insect populations. Our results suggest that the predicted warming may improve food quality and provide better growth conditions for larval development, thus promoting longer or more intense insect outbreaks in the future.     

A perspective on match/mismatch of phenology in community contexts

Climate change has significant impacts on phenology of various organisms in a species‐specific manner. Facing this problem, the match/mismatch hypothesis that phenological (a)synchrony with resource availability strongly influences recruitment success of a consumer population has recently received much attention. In this article, we discuss extending the conventional pairwise concept and demonstrate a community module‐based approach as an initial step for exploring community consequences of species‐specific phenological shifts caused by climate change. Our multispecies match/mismatch perspective leads to the prediction that phenological (a)synchrony among interacting species critically affects not only population recruitment of species but also key dynamical features of ecological communities such as trophic cascades, competitive hierarchies, and species coexistence. Explicit identification and consideration of species relationships is therefore desirable for a better understanding of seasonal community dynamics and thus community consequences of climate change‐induced phenological shifts.     

Food web de‐synchronization in England's largest lake: an assessment based on multiple phenological metrics

Phenological changes have been observed globally for marine, freshwater and terrestrial species, and are an important element of the global biological ‘fingerprint’ of climate change. Differences in rates of change could desynchronize seasonal species interactions within a food web, threatening ecosystem functioning. Quantification of this risk is hampered by the rarity of long‐term data for multiple interacting species from the same ecosystem and by the diversity of possible phenological metrics, which vary in their ecological relevance to food web interactions. We compare phenological change for phytoplankton (chlorophyll a), zooplankton (Daphnia) and fish (perch, Perca fluviatilis) in two basins of Windermere over 40 years and determine whether change has differed among trophic levels, while explicitly accounting for among‐metric differences in rates of change. Though rates of change differed markedly among the nine metrics used, seasonal events shifted earlier for all metrics and trophic levels: zooplankton advanced most, and fish least, rapidly. Evidence of altered synchrony was found in both lake basins, when combining information from all phenological metrics. However, comparisons based on single metrics did not consistently detect this signal. A multimetric approach showed that across trophic levels, earlier phenological events have been associated with increasing water temperature. However, for phytoplankton and zooplankton, phenological change was also associated with changes in resource availability. Lower silicate, and higher phosphorus, concentrations were associated with earlier phytoplankton growth, and earlier phytoplankton growth was associated with earlier zooplankton growth. The developing trophic mismatch detected between the dominant fish species in Windermere and important zooplankton food resources may ultimately affect fish survival and portend significant impacts upon ecosystem functioning. We advocate that future studies on phenological synchrony combine data from multiple phenological metrics, to increase confidence in assessments of change and likely ecological consequences.     

Disentangling climate change effects on species interactions: effects of temperature, phenological shifts, and body size

Climate-mediated shifts in species’ phenologies are expected to alter species interactions, but predicting the consequences of this is difficult because phenological shifts may be driven by different climate factors that may or may not be correlated. Temperature could be an important factor determining effects of phenological shifts by altering species’ growth rates and thereby the relative size ratios of interacting species. We tested this hypothesis by independently manipulating temperature and the relative hatching phenologies of two competing amphibian species. Relative shifts in hatching time generally altered the strength of competition, but the presence and magnitude of this effect was temperature dependent and joint effects of temperature and hatching phenology were non-additive. Species that hatched relatively early or late performed significantly better or worse, respectively, but only at higher temperatures and not at lower temperatures. As a consequence, climate-mediated shifts in hatching phenology or temperature resulted in stronger or weaker effects than expected when both factors acted in concert. Furthermore, consequences of phenological shifts were asymmetric; arriving relatively early had disproportional stronger (or weaker) effects than arriving relatively late, and this varied with species identity. However, consistent with recent theory, these seemingly idiosyncratic effects of phenological shifts could be explained by species-specific differences in growth rates across temperatures and concordant shifts in relative body size of interacting species. Our results emphasize the need to account for environmental conditions when predicting the effects of phenological shifts, and suggest that shifts in size-structured interactions can mediate the impact of climate change on natural communities.     

Trophic mismatch and its effects on the growth of young in an Arctic herbivore

In highly seasonal environments, timing of breeding of organisms is typically set to coincide with the period of highest resource availability. However, breeding phenology may not change at a rate sufficient to keep up with rapid changes in the environment in the wake of climate change. The lack of synchrony between the phenology of consumers and that of their resources can lead to a phenomenon called trophic mismatch, which may have important consequences on the reproductive success of herbivores. We analyzed long‐term data (1991–2010) on climate, plant phenology and the reproduction of a long‐distance Arctic migrant, the greater snow goose (Chen caerulescens atlantica), in order to examine the effects of mismatched reproduction on the growth of young. We found that geese are only partially able to adjust their breeding phenology to compensate for annual changes in the timing of high‐quality food plants, leading to mismatches of up to 20 days between the two. The peak of nitrogen concentration in plants, an index of their nutritive quality for goslings, occurred earlier in warm springs with an early snow melt. Likewise, mismatch between hatch dates of young and date of peak nitrogen was more important in years with early snow melt. Gosling body mass and structural size at fledging was reduced when trophic mismatch was high, particularly when the difference between date of peak nitrogen concentration and hatching was >9 days. Our results support the hypothesis that trophic mismatch can negatively affect the fitness of Arctic herbivores and that this is likely to be exacerbated by rising global temperatures.     

中国东北城乡植被物候时空变化及其对地表温度的响应

以中国东北地区的沈阳、长春、哈尔滨3个大城市及其周边的乡村为研究单元,在像元尺度上采用小波变换法对长时间序列中分辨率成像光谱仪(Moderate-resolution Imaging Spectroradiometer, MODIS)增强植被指数(Enhanced Vegetation Index, EVI)数据滤除噪声数据后重建平滑的EVI曲线,基于EVI曲线,采用动态阈值法提取出研究区2009—2016年植被关键物候期参数指标,即植被生长季开始时间(Start of Growing Season, SOS)和结束时间(End of Growing Season, EOS),分析各研究单元植被物候时空变化特征及其对地表温度的响应特征。结果表明:各研究单元SOS和EOS值的空间分布图存在明显的城乡差异。每一个像元所属的实际位置距离城区中心越近,其SOS值越小,EOS值越大,表明植被生长季开始日期早结束日期晚,整个植被生长期时间变长。各研究单元植被物候参数指标的年际变化趋势具有一定的相似性,即SOS随时间均呈现出提前趋势,且城区和乡村的SOS年际变化趋势保持一致,变化速率各不相同。研究区2012年的SOS值是研究时段内的最大值,从植被物候期反映来看,该年是一个最冷年,这与当年受寒潮影响,出现暴雪,低温等极端天气的气候现象相吻合。各研究单元年均地表温度(Land Surface Temperature,LST)与对应的植被关键物候期参数均有显著的相关性,SOS与LST呈显著负相关,EOS与LST呈高度正相关。即植被物候同期的平均温度越高,植被生长季的起始时间越早,结束时间越晚。     

Fitness Consequences of Timing of Migration and Breeding in Cormorants

In most bird species timing of breeding affects reproductive success whereby early breeding is favoured. In migratory species migration time, especially arrival at the breeding grounds, and breeding time are expected to be correlated. Consequently, migration time should also have fitness consequences. However, in contrast to breeding time, evidence for fitness consequences of migration time is much more limited. Climate change has been shown to negatively affect the synchrony between trophic levels thereby leading to directional selection on timing but again direct evidence in avian migration time is scarce. We here analysed fitness consequences of migration and breeding time in great cormorants and tested whether climate change has led to increased selection on timing using a long-term data set from a breeding colony on the island of Vorsø (Denmark). Reproductive success, measured as number of fledglings, correlated with breeding time and arrival time at the colony and declined during the season. This seasonal decline became steeper during the study period for both migration and breeding time and was positively correlated to winter/spring climate, i.e. selection was stronger after warmer winters/springs. However, the increasing selection pressure on timing seems to be unrelated to climate change as the climatic variables that were related to selection strength did not increase during the study period. There is indirect evidence that phenology or abundances of preferred prey species have changed which could have altered selection on timing of migration and breeding.     

Interactive effects of environmental stress and inbreeding on reproductive traits in a wild bird population

1. Conservation biologists are concerned about the interactive effects of environmental stress and inbreeding because such interactions could affect the dynamics and extinction risk of small and isolated populations, but few studies have tested for these interactions in nature. 2. We used data from the long-term population study of song sparrows Melospiza melodia on Mandarte Island to examine the joint effects of inbreeding and environmental stress on four fitness traits that are known to be affected by the inbreeding level of adult birds: hatching success, laying date, male mating success and fledgling survival. 3. We found that inbreeding depression interacted with environmental stress to reduce hatching success in the nests of inbred females during periods of rain. 4. For laying date, we found equivocal support for an interaction between parental inbreeding and environmental stress. In this case, however, inbred females experienced less inbreeding depression in more stressful, cooler years. 5. For two other traits, we found no evidence that the strength of inbreeding depression varied with environmental stress. First, mated males fathered fewer nests per season if inbred or if the ratio of males to females in the population was high, but inbreeding depression did not depend on sex ratio. Second, fledglings survived poorly during rainy periods and if their father was inbred, but the effects of paternal inbreeding and rain did not interact. 6. Thus, even for a single species, interactions between the inbreeding level and environmental stress may not occur in all traits affected by inbreeding depression, and interactions that do occur will not always act synergistically to further decrease fitness.     

Consequences of a changing environment on the breeding phenology and reproductive success components in a long-distance migratory bird

Migratory birds have a narrow time window to breed, especially in the Arctic, where early nesting typically yields the highest reproductive success. We assessed temporal changes (1991–2015) in reproductive success components in relation to timing of breeding in greater snow geese (Chen caerulescens atlantica). This species breeds in the Canadian Arctic, a region that has experienced a strong warming trend. We tested the effect of laying or hatching date, year and their interaction on six reproductive components: Total clutch laid, nesting success, egg survival, hatching success, prefledging, and postfledging survival. Over 25 years, mean laying date changed little, even though it advanced 1.8 days in early breeders and was delayed 3 days in late breeders. Likewise, the number of eggs in nests initiated early in the season decreased by 0.6 egg, whereas in late nests it increased by 0.3 egg. Success of nests initiated early and late in the season was lower than nests initiated near the population mean, and consistently increased over time. The proportion of eggs surviving to partial predation and postfledging survival decreased with laying date but the pattern did not change over time. In contrast, prefledging survival was not affected by laying date initially but declined in nests initiated late in the season toward the end of the study. Overall, nests initiated close to the population mean showed little temporal change for most components of reproductive success and seem to be less affected by environmental change than nests initiated early and late in the season.     

Are polyphagous geometrid moths with flightless females adapted to budburst phenology of local host species?

The majority of studies demonstrating local adaptation of insect herbivores involve sessile species, particularly those with a parthenogentic phase to their life history or endophagous "parasites" of plants. Current arguments suggest the strength of selection determines whether local adaptation can or cannot take place. Therefore local adaptation should not be limited to species with such traits. We studied the ability of three polyphagous geometrid moths with flightless adult females ( Erannis defoliaria , Operophtera brumata and O. fagata ) to synchronise their egg hatching with the budburst of a local host species in north east Scotland. A strong selection for hatching time is expected among generalist moths given the large variation in budburst phenology and an inability to hatch in synchrony with budburst decreases moth fitness substantially. In two successive seasons, we trapped emerging females from patches of five host species and recorded the temperature sum needed for 50% egg hatch of each brood laid by the trapped females. The hatching times of broods were compared against the average budburst time of the maternal host species in the study area. In addition, the trapping dates of each female were recorded. Only O. brumata showed synchrony with egg hatch and budburst which suggests local phenological adaptation to different host species . This could be maintained by selection and partial reproductive isolation between populations dwelling on different host species. No phenological adaptation was found in the other common geometrids of the study area.     

Simulated climate warming alters phenological synchrony between an outbreak insect herbivore and host trees

As the world’s climate warms, the phenologies of interacting organisms in seasonally cold environments may advance at differing rates, leading to alterations in phenological synchrony that can have important ecological consequences. For temperate and boreal species, the timing of early spring development plays a key role in plant–herbivore interactions and can influence insect performance, outbreak dynamics, and plant damage. We used a field-based, meso-scale free-air forest warming experiment (B4WarmED) to examine the effects of elevated temperature on the phenology and performance of forest tent caterpillar (Malacosoma disstria) in relation to the phenology of two host trees, aspen (Populus tremuloides) and birch (Betula papyrifera). Results of our 2-year study demonstrated that spring phenology advanced for both insects and trees, with experimentally manipulated increases in temperature of 1.7 and 3.4 °C. However, tree phenology advanced more than insect phenology, resulting in altered phenological synchrony. Specifically, we observed a decrease in the time interval between herbivore egg hatch and budbreak of aspen in both years and birch in one year. Moreover, warming decreased larval development time from egg hatch to pupation, but did not affect pupal mass. Larvae developed more quickly on aspen than birch, but pupal mass was not affected by host species. Our study reveals that warming-induced phenological shifts can alter the timing of ecological interactions across trophic levels. These findings illustrate one mechanism by which climate warming could mediate insect herbivore outbreaks, and also highlights the importance of climate change effects on trophic interactions.     

中国东部温带植被生长季节的空间外推估计

利用地面植物物候和遥感归一化差值植被指数（NDVI）数据,以及一种物候-遥感外推方法,实现植被生长季节从少数站点到较多站点的空间外推。结果表明：（1）在1982～1993年期间,中国东部温带地区植被生长季节多年平均起讫日期的空间格局与春季和秋季平均气温的空间格局相关显著;（2）在不同纬度带和整个研究区域,植被生长季节结束日期呈显著推迟的趋势,而开始日期则呈不显著提前的趋势,这与欧洲和北美地区植被生长季节开始日期显著提前而结束日期不显著推迟的变化趋势完全不同;（3）北部纬度带的植被生长季节平均每年延长1．4～3．6d,全区的植被生长季节平均每年延长1．4d,与同期北半球和欧亚大陆植被生长季节延长的趋势数值相近;（4）植被生长季节结束日期的显著推迟与晚春至夏季的区域性降温有关,而植被生长季节开始日期的不显著提前则与晚冬至春季气温趋势的不稳定变化有关;（5）在年际变化方面,植被生长季节开始和结束日期分别与2～4月份平均气温和5～6月份平均气温呈负相关关系。     

Desiccation tolerance in fully developed embryos of two cicadas, Cryptotympana facialis and Graptopsaltria nigrofuscata

In two Japanese cicadas, Cryptotympana facialis and Graptopsaltria nigrofuscata , with different habitat distributions, fully developed embryos hatch in response to high humidity due to rainfall. Despite the advantage of hatching on rainy days, this trait burdens embryos with an extra period of desiccation until the unpredictable advent of rain. We compared the ability of the fully developed embryos of these cicadas to endure periods of low humidity. Eggs were exposed to a combination of different humidities (43% and 75% relative humidity, RH) and durations (0–15 days), and then transferred to an environment with 100% RH to stimulate hatching. In both species, total hatching rates decreased as duration increased, although there was no significant effect of humidity. In C. facialis , a considerable proportion of the eggs hatched during the desiccation period, and the hatching rate was higher at 75% RH than at 43% RH. After transfer to 100% RH, most hatching occurred within a day regardless of the desiccation level. In G. nigrofuscata , no nymphs hatched during the desiccation periods. However, more eggs required more than a day after transfer to 100% RH to hatch after desiccation at 43% RH than at 75% RH. Consequently, the overall proportion of timely hatching of eggs (eggs hatching within a day of moisture supply) was higher after desiccation at 43% RH in C. facialis , but it was higher after desiccation at 75% RH in G. nigrofuscata . These different physiological responses of the two species may reflect adaptation to habitat dryness.     

Breeding success and chronology of Wood Storks Mycteria americana in northern and central Florida, U.S.A.

The breeding success and chronology of Wood Storks Mycteria americana were studied at eight colonies in northern and central Florida during 1981–1985. Mean ± s.d. clutch size for all colony-years was 3.07 ± 0.56 (n = 2694 nests), with three-egg clutches (72%) most frequent. Mean clutch size among all colonies and years ranged from 2.73 ± 0.55 to 3.41 ± 0.61. Many colonies exhibited significant negative trends in clutch size with, hatching date because of a proportional decrease in four-egg clutches later in the season. Mean colony clutch size was not correlated with nest numbers, nesting density or mean hatching date within most years. Mean ± s.d. number of fledglings for all colonies and years was 1.29 ± 1.16 fledglings per nest (n = 2812 nests). Mean annual fledging rates in colonies ranged from 0 (colony failed) to 2.66 fledglings per nest. Most breeding failure occurred prior to egg hatching, and the second highest mortality occurred between hatching and 2 weeks of age. Four-egg clutches fledged more storks than three-egg clutches, which in turn were more successful than two-egg clutches. However, all clutch sizes showed similar fledgling per egg rates. The seasonal decline in productivity was associated proportionally with smaller clutch sizes later in the breeding season. An increase in mean hatching date was correlated with an increase in latitude. There was greater within-year breeding synchrony among colonies than interyear breeding synchrony within each colony. Breeding synchrony was not correlated with mean hatching date, latitude, longitude, nest numbers or nesting density.     

Climate change and the optimal flowering time of annual plants in seasonal environments

Long‐term phenology monitoring has documented numerous examples of changing flowering dates during the last century. A pivotal question is whether these phenological responses are adaptive or not under directionally changing climatic conditions. We use a classic dynamic growth model for annual plants, based on optimal control theory, to find the fitness‐maximizing flowering time, defined as the switching time from vegetative to reproductive growth. In a typical scenario of global warming, with advanced growing season and increased productivity, optimal flowering time advances less than the start of the growing season. Interestingly, increased temporal spread in production over the season may either advance or delay the optimal flowering time depending on overall productivity or season length. We identify situations where large phenological changes are necessary for flowering time to remain optimal. Such changes also indicate changed selection pressures. In other situations, the model predicts advanced phenology on a calendar scale, but no selection for early flowering in relation to the start of the season. We also show that the optimum is more sensitive to increased productivity when productivity is low than when productivity is high. All our results are derived using a general, graphical method to calculate the optimal flowering time applicable for a large range of shapes of the seasonal production curve. The model can thus explain apparent maladaptation in phenological responses in a multitude of scenarios of climate change. We conclude that taking energy allocation trade‐offs and appropriate time scales into account is critical when interpreting phenological patterns.     

Phenological asynchrony between herbivorous insects and their hosts: signal of climate change or pre-existing adaptive strategy?

Climate change alters phenological relations between interacting species. We might expect the historical baseline, or starting-point, for such effects to be precise synchrony between the season at which a consumer most requires food and the time when its resources are most available. We synthesize evidence that synchrony was not the historical condition in two insect–plant interactions involving Edith''s checkerspot butterfly (Euphydryas editha), the winter moth (Operophtera brumata) and their host plants. Initial observations of phenological mismatch in both systems were made prior to the onset of anthropogenically driven climate change. Neither species can detect the phenology of its host plants with precision. In both species, evolution of life history has involved compromise between maximizing fecundity and minimizing mortality, with the outcome being superficially maladaptive strategies in which many, or even most, individuals die of starvation through poor synchrony with their host plants. Where phenological asynchrony or mismatch with resources forms the starting point for effects of anthropogenic global warming, consumers are particularly vulnerable to impacts that exacerbate the mismatch. This vulnerability likely contributed to extinction of a well-studied metapopulation of Edith''s checkerspot, and to the skewed geographical pattern of population extinctions underlying a northward and upward range shift in this species.     

Do long-term changes in sea surface temperature at the breeding areas affect the breeding dates and reproduction performance of Mediterranean loggerhead turtles? Implications for climate change

Global climate change is likely to have an important influence on the phenology, behaviour and population dynamics of many species. We investigate climatic related changes in the breeding phenology of Mediterranean loggerhead marine turtles Caretta caretta over a 19 year period and the potential relationship between these changes and reproductive success and performance. We found that the studied population has experienced fluctuating sea surface temperatures (SST) with an increasing trend during the last century. With increasing spring SST there is a trend towards earlier nesting. However, there is no significant relationship between SST and nesting season, defined as the duration between the first recorded emergence and the last nest laid. Our analyses indicate that marine turtles display phenological changes, and thus maintain favorable thermal conditions at the nesting sites. Furthermore, increasing spring SST was correlated with decreasing clutch size and increasing hatching success that resulted in an apparent lack of correlation between SST and hatchling production. This apparent independence might be misleading since it only holds for a limited range of SST values. Thus, if we estimate the effect of climate change on loggerhead population growth as neutral, based on the apparent independence between SST and total number of hatchlings, we will be underestimating the population extinction risk.