Hierarchical path analysis of deer responses to direct and indirect effects of climate in northern forest

A problem in climate studies has been on how to treat causal chains of explanations and both direct and indirect effects. Mammals in strongly seasonal environments of the boreal forest typically lose condition during winter and gain mass (and reproduce) during the summer season when biomass and plant quality peak. Mass decay of large herbivores during winter is due to direct effects of winter weather, such as increased costs of movement, thermoregulation and reduced access to food when snow is deep. Deer condition during summer is thought to be affected mainly indirectly by weather through plants. High spring temperature speeds up plant development, and deep snow can delay phenology in early summer. Current statistical modelling does not take into account these mechanistic pathways. We used hierarchical Bayes modelling to more mechanistically link global climate, local weather and plant phenology to autumn body mass of red deer in Norway. Red deer were much more affected indirectly through trophic interactions. No solid evidence of direct effects of snow depth was found on autumn body mass. We discuss the implications of our results relative to our ability to predict effects of global change on large mammalian herbivores in the boreal forest.     

Survival and Body Condition of Raccoons at the Edge of the Range

ABSTRACT We investigated the influence of intrinsic and extrinsic variables on overwinter survival of raccoons (Procyon lotor; n = 114) at the northern edge of their distribution. A Cox proportional hazard model identified winter severity as the variable with the greatest influence on raccoon survival (β = 1.08). Autumn body condition estimates (20.5 ± 0.46% total body fat) were relatively stable across years even though we observed large differences in autumn food indices. Variations in autumn body condition did not explain heterogeneity observed in overwinter survival nor the spring condition in which raccoons emerged. Relatively constant autumn body condition suggests reliable availability of anthropogenic food resources may negate variations observed in natural food items on which raccoons rely during hyperphagia. Conversely, spring body condition did vary among years and was highly correlated with winter severity. Accordingly, we also observed a strong inverse relationship with overwinter survival and winter severity. Our findings indicate winter climatic constraints are important factors governing the northern limit of raccoon distribution and changes in winter severity could have important implications in further range expansion of this species.     

Inter-specific synchrony of two contrasting ungulates: wild boar (Sus scrofa) and roe deer (Capreolus capreolus)

Very few studies on ungulates address issues of inter-specific synchrony in population responses to environmental variation such as climate. Depending on whether annual variation in performance of ungulate populations is driven by direct or indirect (trophic) interactions, very different predictions regarding the pattern of inter-specific synchrony can be derived. We compared annual autumn body mass variation in roe deer (Capreolus capreolus) and wild boar (Sus scrofa) from Poland over the period 1982–2002, and related this to variation in winter and summer climate and plant phenological development [the Normalized Difference Vegetation Index (NDVI), derived from satellites]. Roe deer fawns (∼1.3 kg increase from year 1982 to 2002) and yearlings both increased markedly in mass over years. There was also an increase for wild boar mass over years (∼4.2 kg increase for piglets from 1982 to 2002). Despite our failure to link annual body mass to spring or winter conditions or the NDVI, the body mass of roe deer and wild boar fluctuated in synchrony. As this was a field roe deer population, and since wild boar is an omnivore, we suggest this may be linked to annual variation and trends in crop structure (mainly rye). We urge future studies to take advantage of studying multiple species in order to gain further insight into processes of how climate affect ungulate populations. Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible for authorized users.     

Maternal winter body mass and not spring phenology determine annual calf production in an Arctic herbivore

Warming of the Arctic has resulted in earlier snowmelt and green‐up of plants in spring, potentially disrupting the synchrony between plant phenology and breeding phenology in herbivores. A negative relationship between offspring survival in West Greenland caribou and the timing of vegetation emergence was the first finding of such a mismatch in Arctic mammals. However, other studies indicate that the energy for foetal growth and early lactation is predominantly drawn from stored energy reserves typical of ‘capital’ breeders, suggesting that conditions well before spring influence calf production more than the timing of spring onset. Here we use 20 years of observations of marked Svalbard reindeer to evaluate determinants of annual recruitment, as measured by the presence of a calf at foot in mid‐summer. Spring temperatures and the enhanced vegetation index were used as proxies for spring onset, while data on body mass and pregnancy rates in late winter allowed us to determine maternal condition and the reproductive status before spring. Pregnancy rate, offspring survival and annual recruitment were all strongly correlated with average late winter adult female body mass (r = 0.87; r = 0.83; r = 0.92, respectively). Contrary to the findings in West Greenland, neither early calf survival nor annual recruitment were correlated with the two measures of annual variation in spring phenology (r = – 0.07, p = 0.8 and r = – 0.15, p = 0.6, respectively). We also revisit the Greenland data and reveal that the pattern of covariance between early and late measures of fecundity, as well as between early measures of fecundity and offspring survival, correspond with the results from Svalbard. Our results emphasize that conditions affecting maternal body mass during winter explain close to all the variation in recruitment, questioning the importance of the role of a mismatch between plant phenology and calving date.     

Predicting animal performance through climatic and plant phenology variables: The case of an omnivore hibernating species in Finland

Annual variation in the environment is expected to influence individual performance, e.g. measured as body condition, such as body mass or fat deposition, through its direct or indirect effects on food abundance and availability. Such environmental variation is traditionally measured by climatic observation, but recently, measures of environmental phenology obtained from satellite images have been successfully used. We examined the performance of climatic and plant phenology variables in explaining body condition of an invasive omnivore species: the raccoon dog Nyctereutes procyonoides. We collected data on fat deposition of juveniles in southern Finland from the end of June to the beginning of November. A four-parametric logistic model was fitted separately for each province to the data by non-linear regression procedure and the residuals were compared to the expected average as measure of individual performance. These values were then analysed with respect to the environmental variables. Climatic variables describing spring conditions performed better than plant phenology variables in explaining the variation in fat deposition. Harsh spring conditions negatively affected the amount of fat deposed during the growing season. Plant phenology variables, effective in explaining life history traits in herbivores, might not reflect variation in food abundance and quality for omnivore species. We propose that in Europe raccoon dogs will benefit from climate warming, because of a longer growing season, but increased spring precipitation in the form of snow at higher latitudes might compensate for the effect of greater primary productivity and outline the border of their expansion towards harsher environments.     

Are responses of herbivores to environmental variability spatially consistent in alpine ecosystems?

Animal responses to global climate variation might be spatially inconsistent. This may arise from spatial variation in factors limiting populations' growth or from differences in the links between global climate patterns and ecologically relevant local climate variation. For example, the North Atlantic Oscillation (NAO) has a spatially consistent relation to temperature, but inconsistent spatial relation to snow depth in Scandinavia. Furthermore, there are multiple mechanistic ways by which climate may limit animal populations, involving both direct effects through thermoregulation and indirect pathways through trophic interactions. It is conceptually appealing to directly model the predicted mechanistic links. This includes the use of climate variables mimicking such interactions, for example, to use growing degree days (GDD) as a proxy for plant growth rather than average monthly temperature. Using a unique database of autumn body mass of 83331 domestic lambs from the period 1992–2007 in four alpine ranges in Norway, we demonstrate the utility of hierarchical, mechanistic path models fitted using a Bayesian approach to analyse explicitly predicted relationships among environmental variables and between lamb body mass and the environmental variables. We found large spatial variation in strength of responses of autumn lamb body mass to the NAO, to a proxy for plant growth in spring (the Normalized Difference Vegetation Index, NDVI) and effects even differed in direction to local summer climate. Average local temperature outperformed GDD as a predictor of the NDVI, whereas the NAO index in two areas outperformed local weather variables as a predictor of lamb body mass, despite the weaker mechanistic link. Our study highlights that spatial variation in strength of herbivore responses may arise from several processes. Furthermore, mechanistically more appealing measures do not always increase predictive power due to scale of measurement and since global measures may provide more relevant “weather packages” for larger scales.     

Over-winter survival in subadult European rabbits: weather effects,density dependence,and the impact of individual characteristics

The survival probability of an individual may be limited by density-dependent mechanisms and by environmental stochasticity, but can also be modified by individual characteristics. In our study, we investigated over-winter survival of subadults of an enclosed European rabbit Oryctolagus cuniculus population in a temperate zone habitat over the period 1992–2002. We: (1) selected for appropriate models to explain individual variation in over-winter survival and the animals autumn body mass, the latter was used as a measure of the individual pre-winter body condition; and (2) aimed to compare the sensitivity of the target variables on the realised variation of the factors considered. Model selection based on information theory revealed that individual over-winter survival was best explained by the combination of autumn body mass, winter temperature, population density and sex, where the probability of survival was higher in females than in males. According to this model, the probability of survival reacted most sensitively to variation in the autumn body mass and in winter temperature. Individual autumn body mass was best explained by the combination of the date of birth, population density, and weather conditions by means of the percentage of rainy days during the first 2 months after the animals had emerged above ground, where the autumn body mass was negatively related to the percentage of rainy days. The chosen model suggested that the autumn body mass reacted most sensitively to variation in the date of birth. Combining these models, we found that weather conditions during two different periods of time as well as population density, sex and the date of birth operated together to determine the probability of over-winter survival. In particular, the study points out the high impact of environmental stochasticity on over-winter survival: (1) by direct effects of winter temperature conditions, and (2) by the indirect action of weather conditions to which the animals were exposed during the early period of juvenile development.     

Large herbivore migration plasticity along environmental gradients in Europe: life‐history traits modulate forage effects

The most common framework under which ungulate migration is studied predicts that it is driven by spatio–temporal variation in plant phenology, yet other hypotheses may explain differences within and between species. To disentangle more complex patterns than those based on single species/ single populations, we quantified migration variability using two sympatric ungulate species differing in their foraging strategy, mating system and physiological constraints due to body size. We related observed variation to a set of hypotheses. We used GPS‐collar data from 537 individuals in 10 roe Capreolus capreolus and 12 red deer Cervus elaphus populations spanning environmental gradients across Europe to assess variation in migration propensity, distance and timing. Using time‐to‐event models, we explored how the probability of migration varied in relation to sex, landscape (e.g. topography, forest cover) and temporally‐varying environmental factors (e.g. plant green‐up, snow cover). Migration propensity varied across study areas. Red deer were, on average, three times more migratory than roe deer (56% versus 18%). This relationship was mainly driven by red deer males which were twice as migratory as females (82% versus 38%). The probability of roe deer migration was similar between sexes. Roe deer (both sexes) migrated earliest in spring. While territorial male roe deer migrated last in autumn, male and female red deer migrated around the same time in autumn, likely due to their polygynous mating system. Plant productivity determined the onset of spring migration in both species, but if plant productivity on winter ranges was sufficiently high, roe deer were less likely to leave. In autumn, migration coincided with reduced plant productivity for both species. This relationship was stronger for red deer. Our results confirm that ungulate migration is influenced by plant phenology, but in a novel way, that these effects appear to be modulated by species‐specific traits, especially mating strategies.     

东北地区植被物候时序变化

植被与气候的关系非常密切,植被物候可作为气候变化的指示器。东北地区位于我国最北部,是气候变化的敏感区域,研究该区植被物候对气候变化的响应对阐明陆地生态体统碳循环具有重要意义。利用GIMMS AVHRR遥感数据集得到了东北地区阔叶林、针叶林、草原和草甸4种植被25a(1982—2006年)的物候时序变化,得出4种植被春季物候都表现出先提前后推迟的现象,秋季物候的变化则比较复杂,阔叶林和针叶林整体上呈现出秋季物候推迟的趋势,草原和草甸则表现为提前-推迟-提前的趋势。应用偏最小二乘(Partial Least Squares)回归分析了该区域植被物候与气候因子之间的关系,结果表明:春季温度与阔叶林、针叶林和草甸春季物候负相关,前一年冬季温度与草原春季物候正相关,降水与植被春季物候的关系有点复杂;4种植被秋季物候与夏季温度均呈正相关,除草原外,其余3种植被秋季物候均与夏季降水负相关。植被春季物候可能主要受温度影响,而秋季物候很可能主要受降水控制。     

Testing the match–mismatch hypothesis in bighorn sheep in the context of climate change

In species with long gestation, females commit to reproduction several months before parturition. If cues driving conception date are uncoupled from spring conditions, parturition could be mistimed. Mismatch may increase with global change if the rate of temporal changes in autumn cues differs from the rate of change in spring conditions. Using 17 years of data on climate and vegetation phenology, we show that autumn temperature and precipitation, but not vegetation phenology, explain parturition date in bighorn sheep. Although autumn cues drive the timing of conception, they do not predict conditions at parturition in spring. We calculated the mismatch between individual parturition date and spring green-up, assessed whether mismatch increased over time and investigated the consequences of mismatch on lamb neonatal survival, weaning mass and overwinter survival. Mismatch fluctuated over time but showed no temporal trend. Temporal changes in green-up date did not lead to major fitness consequence of mismatch. Detailed data on individually marked animals revealed no effect of mismatch on neonatal or overwinter survival, but lamb weaning mass was negatively affected by mismatch. Capital breeders might be less sensitive to mismatch than income breeders because they are less dependent on daily food acquisition. Herbivores in seasonal environments may access sufficient forage to sustain lactation before or after the spring ‘peak’ green-up, and partly mitigate the consequences of a mismatch. Thus, the effect of phenological mismatch on fitness may be affected by species life history, highlighting the complexity in quantifying trophic mismatches in the context of climate change.     

Experimental evidence of cost of lactation in a low risk environment for a long-lived mammal

In a previous experiment we have documented that organisms adopt a risk-sensitive reproductive allocation when summer reproductive investment competes with survival in the coming winter ( Bårdsen et al. 2008 ). This tradeoff is present through autumn female body mass, which acts as an insurance against unpredictable winter environmental conditions. We tested this hypothesis experimentally on female reindeer experiencing stable and benign winter feeding conditions. Additional supplementary feeding and removal of newborns represented two sets of experimental manipulations. Females in the supplementary feeding group increased more in winter body mass relative to control females. This manipulation, however, did not have any effect on summer body mass development for neither females nor offspring, but we found a positive effect of feeding on offspring birth mass for smaller females. In contrast, offspring removal did have a positive effect on summer body mass development as females in this group were larger in the autumn relative to control females. In essence, we documented two immediate effects as: (1) supplementary feeding did have a positive effect on spring body mass for smaller females; and (2) offspring removal did increase the female summer somatic growth as this had a positive effect on female autumn body mass. Additionally, we tested for lagged effects, but we could not document any biologically significant effects of neither manipulation in the coming spring. The fact that we only found rather weak effects of both manipulations was as expected for risk sensitive individuals experiencing benign environmental conditions over many years.     

Density- and Size-Dependent Winter Mortality and Growth of Late Chaoborus flavicans Larvae

Winter processes such as overwinter survival and growth of individuals can have wide-ranging consequences for population dynamics and communities within and across seasons. In freshwater organisms winter processes have been mainly studied in fish despite that invertebrates also have substantial impacts on lake and pond food webs. One of the major invertebrate consumers in lake and ponds is the planktonic larvae of the dipteran insect Chaoborus spec. However, while much is known about Chaoborus feeding ecology, behaviour and structuring role in food webs, its winter ecology and how it affects its populations are poorly understood. Here size- and density-dependent winter mortality and body growth of late Chaoborus flavicans larvae were quantified over naturally occurring size and density ranges in autumn and under natural winter conditions using two field enclosure experiments. Winter mortality increased with autumn density but decreased with autumn body size while winter growth rates decreased with autumn density and body sizes. There was also a density- and size-independent background mortality component. The proportion of pupae found in spring decreased strongly and exponentially with autumn density. These results may explain the commonly observed univoltine life cycle and multi-annual density fluctuations in northern Chaoborus populations. They further demonstrate the relevance of winter processes and conditions for freshwater invertebrates and ecosystems.     

Lower body mass and higher metabolic rate enhance winter survival in root voles,Microtus oeconomus

Although the biological significance of individual variation in physiological traits is widely recognized, studies of their association with fitness in wild populations are surprisingly scarce. We investigated the effect of individual phenotypic variation in body mass, resting (RMR) and peak metabolic rates (PMR) on mortality of the root vole Microtus oeconomus. Body mass and metabolic rates varied significantly among consecutive years and were also age dependent, as individuals born in late summer and autumn were characterized by significantly lower body mass and metabolic rates than animals born earlier. At the beginning of winter voles born in spring and early summer exhibited reduced body mass and metabolic rates, whereas animals born later maintained lower body mass and RMR, which may be interpreted as phenotypic plasticity enhancing the probability of survival. Body mass had no significant effect on vole survival during summer. In contrast, smaller individuals were characterized by lower mortality during early winter, whereas higher body mass was positively associated with survival later in the season. High body‐mass‐corrected RMR positively affected survival in both summer and winter. The effect of PMR was apparent only during winter, though its direction (and correlation with RMR) varied among years. Deep snow cover negatively affected the survival of voles in both early and late winter. Ambient temperature was positively associated with winter survival, except for late winter, when rising temperature caused flooding of vole habitat. We conclude that the lack of consistency in the directionality and strength of the effects of body mass and metabolic rates on winter survival does not undermine their importance, but rather demonstrates the ability of individuals to adjust metabolic rate to changing environmental conditions. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 297–309.     

Survival of Adult Female Bighorn Sheep Following a Pneumonia Epizootic

Beginning in the early 1900s, poly-factorial, poly-microbial pneumonia was identified as a disease affecting bighorn sheep (Ovis canadensis) and it continues to threaten bighorn populations, posing an ongoing management challenge. In May and June 2013, a pneumonia outbreak linked to the pathogen Mycoplasma ovipneumoniae led to an all-age die-off of desert bighorn sheep (O. c. nelsoni) at Old Dad Peak in the Kelso Mountains of the Mojave Desert in California, USA. Subsequently, we observed clinical signs of respiratory disease among bighorn sheep in multiple neighboring ranges. Our objective was to investigate post-outbreak survival of adult female bighorn across 9 populations from 2014 to 2017 in the Mojave Desert and evaluate the relationship between M. ovipneumoniae infection and survival, while testing effects of range factors that could potentially influence differences in adult female survival (i.e., forage quality, winter precipitation, population abundance). We fitted adult females with radio-collars following the outbreak and collected serum and nasal swab samples for competitive enzyme-linked immunosorbent assay (cELISA) and polymerase chain reaction (PCR) testing to determine exposure and infection status at time of capture. We tracked survival of 115 adult females with radio-collars and used the known-fate model in Program MARK to evaluate effects and estimate survival from November 2013 to March 2017. Annual survival was negatively correlated with positive infection status at capture but varied across populations with respect to differences in range conditions. Summer and autumn forage quality, as represented by mean normalized difference vegetation index (NDVI) values for summer and autumn, was positively correlated with overwinter survival, whereas winter precipitation (a proxy for winter severity) was negatively correlated with overwinter survival. Population abundance was negatively correlated with annual survival, suggesting a potential density-dependent effect. Model-averaged annual survival estimates ranged from 0.700 ± 0.07 (SE) to 0.945 ± 0.026 for infected individuals and 0.896 ± 0.03 to 0.983 ± 0.011 for uninfected individuals. We conclude that summer and autumn forage quality, indexed by NDVI, may partially offset the negative effect associated with M. ovipneumoniae infection on host survival. Our survival modeling results suggest that chronic infection may have afflicted adult females that were PCR-positive (i.e., infected with M. ovipneumoniae) at time of capture. We propose programmatic re-testing of infected individuals to assess pathogen persistence at the individual level and evaluate whether selective culling might potentially help to reduce prevalence and transmission within populations. © 2020 The Wildlife Society.     

Using a proxy of plant productivity (NDVI) to find key periods for animal performance: the case of roe deer

Animals in seasonal environments are affected by climate in very different ways depending on season and part of the climatic effects operates indirectly through the plants. Vegetation conditions in spring and summer are regarded as decisive for the reproductive success and the offspring's condition of large herbivores, but objective ways to determine key periods during the growing season have not been done often due to limitations in plant data. Using satellite data (NDVI), we determined how plant productivity from birth to fall influences the following winter body mass of roe deer fawns. We do this in two populations, the first inhabiting the low productive Chizé reserve in south western France with an oceanic climate and the second from Trois Fontaines, a highly productive forest with continental climate in east France. The effect of plant productivity was similar for male and female fawn mass, as expected from the weak intensity of sexual selection in roe deer life history traits. We found contrasting results between sites, with a strong effect of plant productivity in spring (April-May) in the Chizé population, but no effect in the Trois Fontaines population. The relatively low variability in winter fawn body mass could account for the absence of NDVI effects at Trois Fontaines. However, such results might also point to a limitation in the use of the NDVI, since the relationship between the canopy and the plant productivity at the ground level might be weak in the highly productive forest of Trois Fontaines.     

Temperature alone does not explain phenological variation of diverse temperate plants under experimental warming

Anthropogenic climate change has altered temperate forest phenology, but how these trends will play out in the future is controversial. We measured the effect of experimental warming of 0.6–5.0 °C on the phenology of a diverse suite of 11 plant species in the deciduous forest understory (Duke Forest, North Carolina, USA) in a relatively warm year (2011) and a colder year (2013). Our primary goal was to dissect how temperature affects timing of spring budburst, flowering, and autumn leaf coloring for functional groups with different growth habits, phenological niches, and xylem anatomy. Warming advanced budburst of six deciduous woody species by 5–15 days and delayed leaf coloring by 18–21 days, resulting in an extension of the growing season by as much as 20–29 days. Spring temperature accumulation was strongly correlated with budburst date, but temperature alone cannot explain the diverse budburst responses observed among plant functional types. Ring‐porous trees showed a consistent temperature response pattern across years, suggesting these species are sensitive to photoperiod. Conversely, diffuse‐porous species responded differently between years, suggesting winter chilling may be more important in regulating budburst. Budburst of the ring‐porous Quercus alba responded nonlinearly to warming, suggesting evolutionary constraints may limit changes in phenology, and therefore productivity, in the future. Warming caused a divergence in flowering times among species in the forest community, resulting in a longer flowering season by 10‐16 days. Temperature was a good predictor of flowering for only four of the seven species studied here. Observations of interannual temperature variability overpredicted flowering responses in spring‐blooming species, relative to our warming experiment, and did not consistently predict even the direction of flowering shifts. Experiments that push temperatures beyond historic variation are indispensable for improving predictions of future changes in phenology.     

Host plant-mediated variation in overwintering site quality: implications for the size and composition of populations of Acanthoscelides alboscutellatus (Coleoptera: Bruchidae)

This study provides an example of how variation in the quality of overwintering sites provided by the host plant of an insect seed predator can influence both the probability of overwintering survival and the size and composition of postwintering populations. Thus, the concept of host plant quality is extended to include variation in the suitability of the overwintering site of temperate region insects that overwinter within, or in habitats created by, their host plant. Adult Acanthoscelides alboscutellatus (Horn) (Coleoptera: Bruchidae) overwinter inside the fruit of Ludwigia alternifolia (L.) (Onagraceae). In early winter, however, fruits begin to dehisce, i.e., one or more of the fruit's four sides and/or top are shed. Variation in the onset and extent of dehiscence creates a range of overwintering habitats that vary in exposure to ambient conditions. In this study the frequency of possible overwintering sites in natural populations of L. alternifolia was determined by monitoring the phenology of fruit dehiscence from October through May in two populations for four years and for a third population for three years. Winter survivorship of adult A. alboscutellatus was assessed experimentally in eight environments representative of the conditions created by variation in dehiscence. These environments were produced by crossing four levels of exposure (degree of dehiscence) with two locations of the overwintering site, i.e., above or on the ground surface. The onset, phenology, and overall frequency of fruit dehiscence varied markedly among populations and years. Exposure, location, and their interaction had strong effects on survival and accounted for 80% of the observed variation in winter survival. Survivorship was higher on than above the ground, and in both locations decreased with increasing exposure. Thus, variation in fruit dehiscence among L. alternifolia populations will influence the size of postwintering A. alboscutellatus populations by dictating the quality of overwintering sites. Adult beetles that over-winter inside indehiscent fruit experience selection for small body size, associated with high mortality, when they attempt to exit the fruit at eclosion. As a consequence, the frequency of fruit dehiscence at eclosion coupled with the relative survival rates of adults within indehiscent fruit will determine the body size composition of postwintering populations and hence the response to selection for small body size in this species.     

Flowering phenology and reproduction of a forest understorey plant species in response to the local environment

Temperate forest understorey plants are subjected to a strong seasonality in their optimal growing conditions. In winter and early spring, low temperatures are suboptimal for plant growth while light becomes limited later in spring season. We can thus expect that differences in plant phenology in relation to spatiotemporal environmental variation will lead to differences in reproductive output, and hence selection. We specifically studied whether early flowering, a paradoxical pattern that is observed in many plant species, is an adaptive strategy, and whether selection for early flowering was confounded with selection for flower duration or was attributable to environmental variables. We used Geum urbanum as a study species to investigate the effect of relevant environmental factors on the species’ flowering phenology and the consequences for plant reproductive output. We monitored the phenology of four to six plants in each of ten locations in a temperate deciduous forest (Belgium). We first quantified variation in flowering time within individuals and related this temporal variation to individual flower reproductive output. Then, we studied inter-individual variation here-in and linked this to reproduction at the plant level, hence studying the selection differential. We found that flowering within individual plants of Geum urbanum was spread over a long period from June to October. Reproductive output of individual flowers, measured as total seed mass per flower, declined during the season. We found no indication for selection for early flowering but rather for longer flower duration. Larger plants had an earlier flowering onset and a higher seed mass, which suggests that these factors covary and are condition dependent. None of the studied environmental variables could explain plant size, although soil pH and to a lesser extent light availability had a positive direct effect on seed mass per plant. Finally, we suggest that the high intra-individual variation in flowering time, which might be a risk spreading strategy of the plant in the presence of seed predation, limits the potential for selection on flowering phenology.

     

西安4种落叶乔木物候期对气候变化的响应

以西安1979—2018年的气候资料和植物物候观测资料为基础,采用分段回归和趋势倾向率等方法,分析了毛白杨(Populus tomentosa)、杜梨(Pyrus betulifolia)、七叶树(Aesculus chinensis)和灯台树(Bothrocaryum controversum)4种落叶乔木展叶盛期和叶全变色期的生长趋势,使用偏相关分析探讨了气温、降水和日照时数与物候期的关系并通过偏最小二乘回归(Partial Least Squares, PLS)判断气候变量对物候期的综合影响。结果表明：(1)1979—2018年,4种乔木的生长季长度延长,整体表现为春季物候期提前,秋季物候期推迟;(2)展叶盛期物候指标与叶全变色期物候指标,转折均发生在1982年;转折后,物候特征变化显著,春季物候的提前速率和秋季物候的推迟速率加快,展叶盛期平均提前3.8d/10a,叶全变色期平均推迟4.7d/10a;(3)展叶盛期与春季气温表现为极显著负相关,叶变色期与秋季气温表现为显著正相关;降水对植物物候的影响不显著;春季物候与日照时数呈现极显著负相关关系,秋季物候期与日照时数呈不显著正相关...     

Observational evidence of risk-sensitive reproductive allocation in a long-lived mammal

Organisms should adopt a risk-sensitive reproductive allocation when summer reproductive allocation competes with survival in the coming winter. This trade off is shown through autumn female body mass, which acts as an insurance against unpredictable winter environmental conditions. We tested this hypothesis on female reindeer in a population that has experienced a time period of dramatic increase in abundance. Environmental conditions during winter were fairly stable (with the exception of 1 year). We conclude that increased population abundance (perhaps in interaction with winter environmental conditions) could have represented a worsening of winter environmental conditions as both autumn offspring and spring female body mass decreased during the course of the study. Moreover, we found that the cost of reproduction was related to environmental conditions as: (1) autumn body mass was larger for barren than for lactating females, and this difference was temporally highly variable; (2) lactating females produced smaller offspring than barren ones in the following year; and (3) reproductive output (offspring size) decreased over time. We also found evidence of quality effects as lactating females had a higher reproductive success in the following year. In sum, a worsening of winter conditions lead to: (1) decreased reproductive output; (2) lowered autumn body mass for lactating females; and (3) increased body mass for barren females. Since females reduce their reproductive allocation as winter conditions becomes more severe, we conclude that reindeer have adopted a risk-sensitive reproductive allocation.