Impact of Puumala virus infection on maturation and survival in bank voles: a capture-mark-recapture analysis

Many zoonotic diseases are caused by rodent-borne viruses. Major fluctuations in the transmission of these viruses have been related to large changes in reservoir host population numbers due to external factors. However, the impact of the pathogen itself on the demography of its reservoir host is often overlooked. We investigated the impact of Puumala virus (PUUV) on survival and reproductive maturation probability of its reservoir host, the bank vole (Myodes glareolus). Three years (2004-06) of data from nine independent sites in southern Belgium were collected and analyzed with a capture-mark-recapture (CMR) method that includes statistical correction for the variation in capture probability of voles. A multistate model based on four states of reproductive activity and PUUV immunoglobulin G (IgG) antibody status was used to estimate survival and probability of transition from one reproductive or infection state to another. Although survival estimates for reproductively active voles were similar between infected and noninfected individuals, PUUV infection in reproductively inactive voles decreased mean monthly survival by 14%. PUUV infection was associated with a threefold increase in the probability of reproductive maturation in bank voles. Moreover, the probability of PUUV IgG seroconversion was three times higher for reproductively active voles compared to reproductively inactive voles. Our model indicates that PUUV infection may alter bank vole population dynamics by affecting both survival and maturation in its host. Additional studies, using CMR methodology with shorter time intervals between trapping sessions and possibly a longer duration, are needed to confirm these findings.     

Effects of competition and season on survival and maturation of young bank vole females

In territorial microtines intra-specific density dependent processes can limit the maturation of individuals during the summer of their birth. This may have demographic consequences by affecting the number and the age distribution of breeding individuals in the population. Little is known about this process on a community level, though populations of many northern microtine species fluctuate in synchrony and are known to interfere socially with each other. We experimentally studied the influence of the field vole Microtus agrestis on maturation, breeding, space use and survival of weanling bank voles, Clethrionomys glareolus. Two additive competition experiments on bank vole populations were conducted in large outdoor enclosures, half of them additionally housing a field vole population. In a mid-summer experiment low population density and absence of older breeding females minimised intra-specific competition. Survival was not affected by the presence of field voles. Season had a significant effect on both the probability of maturation and breeding of the weanlings. Competition with field voles significantly delayed breeding, and coupled with seasonal effects decreased the probability of breeding. In a late-summer experiment breeding and survival of bank vole weanlings were studied for three weeks as part of a high density breeding bank vole population. Weanlings did not mature at all nor were their space use and survival affected by the presence of field voles. Our results show that competition with other species can also have an impact on breeding of immatures. In an extreme seasonal environment, even a short delay of breeding may decrease survival chances of offspring. Seasonal and competition effects together may thus limit the contribution of year born females to reproductive output of the population. Other studies have shown that adult breeding bank voles suffer lower survival in the presence of field voles, but this study showed no survival effects on the weanlings. Thus it might be beneficial for weanlings to stay immature especially in the end of the breeding season and postpone reproduction to the next breeding season if densities of competing species are high.     

Moose recruitment in relation to bilberry production and bank vole numbers along a summer temperature gradient in Norway

The plant stress hypothesis states that plant stress factors other than herbivory improve herbivore performance due to changes in the content of nutritive or defensive compounds in the plants. In Norway, the bilberry (Vaccinium myrtillus) is important forage for the bank vole (Myodes glareolus) in winter and for the moose (Alces alces) in summer and autumn. The observed peaks in bank vole numbers after years with high production of bilberries are suggested to be caused by increased winter survival of bank voles due to improved forage quality. High production of bilberries should also lead to higher recruitment rates in moose in the following year. We predict, however, that there is an increasing tendency for a 1-year delay of moose indices relative to vole indices with decreasing summer temperatures, because low temperatures prolong the period needed by plants to recover in the vole peak year, and thus positively affect moose reproduction also in the succeeding year. In eight out of nine counties in south-eastern Norway, there was a positive relationship between the number of calves observed per female moose during hunting and a bilberry seed production index or an autumn bank vole population index. When dividing the study area into regions, there was a negative relationship between a moose-vole time-lag index and the mean summer temperature of the region. These patterns suggest that annual fluctuations in the production of bilberries affect forage quality, but that the effect on moose reproduction also depends on summer temperatures.     

Predator population dynamics under a cyclic prey regime: numerical responses,demographic parameters and growth rates

The consequences of cyclic fluctuations in abundance of prey species on predator continue to improve our understanding of the mechanisms behind population regulation. Among predators, vole‐eating raptors usually respond to changes in prey abundance with no apparent time‐lag and therefore contradict predictions from the predator–prey theory. In such systems, the interplay between demographic traits and population growth rate in relation to prey abundance remains poorly studied, yet it is crucial to characterize the link between ecological processes and population changes. Using a mechanistic approach, we assessed the demographic rates associated to the direct and indirect numerical responses of a specialist raptor (Montagu's harrier) to its cyclic prey (common vole), using long term data from two adjacent study sites in France. First‐year survival rates were weakly affected by vole abundance, probably due to the fact that Montagu's harriers are trans‐Saharan migrants and thus escape the vole collapse occurring in autumn–winter. Recruitment of yearling as well as breeding propensity of experienced adult females were strongly affected by vole abundance and at least partially shaped the trajectory of the breeding population. We argued that the strong density dependent signal detected in predator time series was mostly the phenomenological consequence of the positive direct numerical response of harriers to vole abundance. Accounting for this, we proposed a method to assess density dependence in predator relying on a cyclic prey. Finally, the variation in Montagu's harrier population growth rates was best explained by overwinter growth rates of the prey population and to a lesser extent by previous residual predator density.     

Woodland recovery after suppression of deer: cascade effects for small mammals, wood mice (Apodemus sylvaticus) and bank voles (Myodes glareolus)

Over the past century, increases in both density and distribution of deer species in the Northern Hemisphere have resulted in major changes in ground flora and undergrowth vegetation of woodland habitats, and consequentially the animal communities that inhabit them. In this study, we tested whether recovery in the vegetative habitat of a woodland due to effective deer management (from a peak of 0.4-1.5 to <0.17 deer per ha) had translated to the small mammal community as an example of a higher order cascade effect. We compared deer-free exclosures with neighboring open woodland using capture-mark-recapture (CMR) methods to see if the significant difference in bank vole (Myodes glareolus) and wood mouse (Apodemus sylvaticus) numbers between these environments from 2001-2003 persisted in 2010. Using the multi-state Robust Design method in program MARK we found survival and abundance of both voles and mice to be equivalent between the open woodland and the experimental exclosures with no differences in various metrics of population structure (age structure, sex composition, reproductive activity) and individual fitness (weight), although the vole population showed variation both locally and temporally. This suggests that the vegetative habitat--having passed some threshold of complexity due to lowered deer density--has allowed recovery of the small mammal community, although patch dynamics associated with vegetation complexity still remain. We conclude that the response of small mammal communities to environmental disturbance such as intense browsing pressure can be rapidly reversed once the disturbing agent has been removed and the vegetative habitat is allowed to increase in density and complexity, although we encourage caution, as a source/sink dynamic may emerge between old growth patches and the recently disturbed habitat under harsh conditions.     

Evidence for different bottom-up mechanisms in wood mouse ( Apodemus sylvaticus ) and bank vole ( Myodes glareolus ) population fluctuations in Southern Norway

Animals that feed on forest tree seeds, such as Apodemus mice, increase in number after a mast year. At high latitudes, there is a similar delayed response by Myodes voles to high seed crops of bilberry (Vaccinium myrtillus), but here the mechanism is hypothesised to be increased forage quality, caused by a trade-off between reproduction and defence in the plants. Both Apodemus mice and Myodes voles eat berries, but only the latter feed on bilberry plants. Hence, only Myodes voles are predicted to respond to bilberry peak years. A second prediction is that the effect should last longer than any possible direct impacts of bilberries, because the plants would not be able to rebuild their defence until the succeeding summer. During a 21-year snap-trapping study of small rodents in Southern Norway, the spring population of bank vole (Myodes glareolus) was positively related to a bilberry seed index of the previous year, indicating increased winter survival, whereas the wood mouse (Apodemus sylvaticus) was not affected. Also the succeeding autumn population index of the bank vole was positively related to the bilberry index of the previous year, even when controlling for spring population levels. The wood mouse population responded to mast years of sessile oak (Quercus petraea), whereas seeds of Norway spruce (Picea abies) seemed to have some impact on both species. It is concluded that these rodents are mainly limited from below, but by different mechanisms for the granivorous and the herbivorous species.     

Spatial synchrony in vole population fluctuations – a field experiment

Three mechanisms have been proposed to induce spatial synchrony in fluctuations of small mammal populations: climate‐related environmental effects, predation and dispersal. We conducted a field experiment in western Finland to evaluate the relative roles of these mechanisms in inducing spatial synchrony among cyclic populations of field voles Microtus agrestis. The study was conducted during the increase and peak phases of a vole population cycle on four agricultural field sites situated 1.5–7.0 km apart. Each field contained two 0.5‐ha fenced enclosures and one 1‐ha unfenced control area. One enclosure per field allowed access by small mustelid predators and the other by avian predators; all enclosures prevented the dispersal of voles. The unfenced control areas allowed access by all predators as well as dispersal by voles. Enclosed vole populations were in a treatment‐wise asynchronous phase before the predator access treatments were applied. The growth rates of all enclosed populations were tightly synchronized during the course of the experiment. Conversely, synchrony both among the unfenced populations and between the fenced and unfenced populations was practically non‐existent. During winter, in the increase phase of the cycle, vole populations in all treatments declined to low densities due to a seasonal effect of winter food depletion. During summer, in the peak year of the vole cycle, all populations fluctuated in synchrony. At this time, both small mustelids and birds of prey appeared to be abundant enough to induce synchrony. Dispersal was identified as a potential contributor to synchronization, but the magnitude of its effects could not be reliably discerned. Our results indicate that no single mechanism can account for the observed patterns of spatial synchrony among cyclic northern vole populations. Rather, spatial synchronization is induced by different mechanisms, namely seasonality and predation, acting successively during different seasons and phases of the vole cycle.     

Strong “bottom‐up” influences on small mammal populations: State‐space model analyses from long‐term studies

“Bottom‐up” influences, that is, masting, plus population density and climate, commonly influence woodland rodent demography. However, “top‐down” influences (predation) also intervene. Here, we assess the impacts of masting, climate, and density on rodent populations placed in the context of what is known about “top‐down” influences. To explain between‐year variations in bank vole Myodes glareolus and wood mouse Apodemus sylvaticus population demography, we applied a state‐space model to 33 years of catch‐mark‐release live‐trapping, winter temperature, and precise mast‐collection data. Experimental mast additions aided interpretation. Rodent numbers in European ash Fraxinus excelsior woodland were estimated (May/June, November/December). December–March mean minimum daily temperature represented winter severity. Total marked adult mice/voles (and juveniles in May/June) provided density indices validated against a model‐generated population estimate; this allowed estimation of the structure of a time‐series model and the demographic impacts of the climatic/biological variables. During two winters of insignificant fruit‐fall, 6.79 g/m² sterilized ash seed (as fruit) was distributed over an equivalent woodland similarly live‐trapped. September–March fruit‐fall strongly increased bank vole spring reproductive rate and winter and summer population growth rates; colder winters weakly reduced winter population growth. September–March fruit‐fall and warmer winters marginally increased wood mouse spring reproductive rate and September–December fruit‐fall weakly elevated summer population growth. Density dependence significantly reduced both species' population growth. Fruit‐fall impacts on demography still appeared after a year. Experimental ash fruit addition confirmed its positive influence on bank vole winter population growth with probable moderation by colder temperatures. The models show the strong impact of masting as a “bottom‐up” influence on rodent demography, emphasizing independent masting and weather influences; delayed effects of masting; and the importance of density dependence and its interaction with masting. We conclude that these rodents show strong “bottom‐up” and density‐dependent influences on demography moderated by winter temperature. “Top‐down” influences appear weak and need further investigation.     

Characterising demographic contributions to observed population change in a declining migrant bird

Populations of Afro‐Palearctic migrant birds have shown severe declines in recent decades. To identify the causes of these declines, accurate measures of both demographic rates (seasonal productivity, apparent survival, immigration) and environmental parameters will allow conservation and research actions to be targeted effectively. We used detailed observations of marked breeding birds from a ‘stronghold’ population of whinchats Saxicola rubetra in England (stable against the declining European trend) to reveal both on‐site and external mechanisms that contribute to population change. From field data, a population model was developed based on demographic rates from 2011 to 2014. Observed population trends were compared to the predicted population trends to assess model‐accuracy and the influence of outside factors, such as immigration. The sensitivity of the projected population growth rate to relative change in each demographic rate was also explored. Against expectations of high productivity, we identified low seasonal breeding success due to nocturnal predation and low apparent first‐year survival, which led to a projected population growth rate (λ) of 0.818, indicating a declining trend. However, this trend was not reflected in the census counts, suggesting that high immigration was probably responsible for buffering against this decline. Elasticity analysis indicated λ was most sensitive to changes in adult survival but with covariance between demographic rates accounted for, most temporal variation in λ was due to variation in productivity. Our study demonstrates that high quality breeding habitat can buffer against population decline but high immigration and low productivity will expose even such stronghold populations to potential decline or abandonment if either factor is unsustainable. First‐year survival also appeared low, however this result is potentially confounded by high natal dispersal. First‐year survival and/or dispersal remains a significant knowledge gap that potentially undermines local solutions aimed at counteracting low productivity.     

The tragedy of the commons: unsustainable population structure of Iberian red deer in hunting estates

Hunting can influence population structure with consequences in ecological and evolutionary processes. Populations of Iberian red deer (Cervus elaphus hispanicus) in Spain occur under two different management regimes: fenced and unfenced (open) estates. We compared census data, hunting bags and data from hunted individuals between both types of estates. Harvest on stags was moderate in fenced estates but strong in open ones, probably due to the competition between neighbouring landowners over the same deer populations. On the contrary, female culling was low in open estates compared to fenced ones. As a result, populations in open estates have mostly young males and strongly female-biased sex ratios. Female-biased population structure in open estates did not result in higher number of males being harvested per year compared with fenced estates, probably due to negative effects on development, survival and reproduction, and harvested males were younger, and hence, with smaller antlers. There is published evidence for undesirable effects of biased population sex ratio and age structure in these red deer populations. Our results indicate that this type of management may be unsustainable and recommend that harvest on males in open estates should be reduced and that on females increased, in order to maintain a more balanced population structure that may allow sustainable population dynamics and the operation of natural evolutionary processes.     

Recruitment rates exhibit high elasticity and high temporal variation in populations of a short-lived perennial herb

Empirical studies for different life histories have shown an inverse relationship between elasticity (i.e. the proportional contribution to population growth rate) and temporal variation in vital rates. It is accepted that this relationship indicates the effect of selective pressures in reducing variation in those life‐history traits with a major impact on fitness. In this paper, we sought to determine whether changes in environmental conditions affect the relationship between elasticity of vital rates and their temporal variation, and whether vital rates with simultaneously large elasticity and temporal variation might represent a characteristic life‐history strategy. We used demographic data on 13 populations of the short‐lived Hypericum cumulicola over 5–6 years, in three time‐since‐fire classes. For each population of each time‐since‐fire, we computed the mean matrix over years and its respective elasticity matrix, and the coefficients of variation in matrix entries over study years as an estimate of temporal variability. We found that mean elasticity negatively significantly correlated with temporal variation in vital rates in populations (overall eight out of 13) included in each time‐since‐fire. However, seedling recruitment exhibited both high elasticity and high temporal variation in almost all study populations. These results indicated that (1) the general relationship between elasticity and temporal variation in vital rates was not modified by environmental changes due to time‐since‐fire, and (2) high elasticity and high temporal variation in seedling recruitment in H. cumulicola is a particular trait of the species' life history. After seed survival in the soil seed bank, seedling recruitment represents the most important life‐history trait influencing H. cumulicola population growth rate (and fitness). The high temporal variability in seedling recruitment suggests that this trait is determined by environmental cues, leading to an increase in population size and subsequent replenishment of the seed bank in favorable years.     

Temporal patterns of genetic variation across a 9-year-old aerial seed bank of the shrub Banksia hookeriana (Proteaceae)

The pattern of accumulation of genetic variation over time in seed banks is poorly understood. We examined the genetic structure of the aerial seed bank of Banksia hookeriana within a single 15-year-old population in fire-prone southwestern Australia, and compared genetic variation between adults and each year of a 9-year-old seed bank using amplified fragment length polymorphism (AFLP). B. hookeriana is well suited to the study of seed bank dynamics due to the canopy storage of its seeds, and because each annual crop can be identified. A total of 304 seeds from nine crop years and five maternal plants were genotyped, along with 113 plants from the adult population. Genetic variation, as assessed by the proportion of polymorphic markers (P(p)) and Shannon's index (I), increased slightly within the seed bank over time, while gene diversity (H(j)), did not change. P(p), I, and H(j) all indicated that genetic variation within the seed bank quickly approached the maximal level detected. Analysis of molecular variance revealed that less than 4% of variation could be accounted for by variation among seeds produced in different years, whereas there was greater differentiation among maternal plants (12.7%), and among individual seeds produced by different maternal plants (83.4%). With increasing population age, offspring generated each year were slightly more outbred, as indicated by an increase in the mean number of nonmaternal markers per offspring. There were no significant differences for H(j) or I between adults and the seed bank. Viability of seeds decreased with age, such that the viability of 9-year-old seeds was half that of 2-year-old seeds. These results suggest that variable fire frequencies have only limited potential to influence the amount of genetic variation stored within the seed bank of B. hookeriana.     

Ecological longevity of Polaskia chende (Cactaceae) seeds in the soil seed bank,seedling emergence and survival

• Soil seed banks are essential elements of plant population dynamics, enabling species to maintain genetic variability, withstand periods of adversity and persist over time, including for cactus species. However knowledge of the soil seed bank in cacti is scanty. In this study, over a 5‐year period we studied the seed bank dynamics, seedling emergence and nurse plant facilitation of Polaskia chende, an endemic columnar cactus of central Mexico.
• P. chende seeds were collected for a wild population in Puebla, Mexico. Freshly collected seeds were sown at 25 °C and 12‐h photoperiod under white light, far‐red light and darkness. The collected seeds were divided in two lots, the first was stored in the laboratory and the second was use to bury seeds in open areas and beneath a shrub canopy. Seeds were exhumed periodically over 5 years. At the same time seeds were sown in open areas and beneath shrub canopies; seedling emergence and survival were recorded over different periods of time for 5 years.
• The species forms long‐term persistent soil seed banks. The timing of seedling emergence via germination in the field was regulated by interaction between light, temperature and soil moisture. Seeds entered secondary dormancy at specific times according to the expression of environmental factors, demonstrating irregular dormancy cycling.
• Seedling survival of P. chende was improved under Acacia constricta nurse plants. Finally, plant facilitation affected the soil seed bank dynamics as it promoted the formation of a soil seed bank, but not its persistence.

     

The role of the seed bank,seed rain and the timing of disturbance in gap regeneration

Abstract. Question: Does the degree and timing of disturbance contribute significantly to the pattern and process of regeneration in plant communities as a consequence of the availability and number of species of propagules present? Location: Acid grassland at 230 m a.s.l., eastern Scotland, UK. Methods : Plots were surface disturbed or had their soil profile inverted at monthly intervals at 12 dates during a year. Seed bank and seed rain were assessed at each treatment time. The effect of disturbance intensity and timing on the regenerating vegetation was assessed. Results: Removing the seed bank significantly slowed regeneration, as it contributed 43 % of developing cover after one year where it was present. At an individual seed level, seed in the seed rain had a much higher likelihood of contributing to the regenerating vegetation than a seed in the seed bank. Some species showed a reliance on the seed bank for regeneration, and hence there was a significant difference in the vegetation that developed between plots with the seed bank intact and those where it was removed. Winter disturbed plots (little seed rain) had slower rates of re‐vegetation than summer disturbed plots. Timing had little effect on species composition, though a significantly higher cover of perennial forb species developed on the winter disturbed plots. Conclusion: Removing the contribution of the seed bank had a greater effect on the composition of regenerating vegetation than the effect of seasonal variation on the seed rain.     

Density-dependent variation in body mass of voles

We studied inter-annual, spatial and sexual variation in the body mass of bank volesMyodes glareolus Schreber, 1780 and grey-sided volesMyodes rufocanus Sundevall, 1846 using live trappings from two grids on the southand north-facing slopes of a mountain valley in Southern Norway. Variation in spring density of the four populations was consistent with cyclic dynamics (n=7,s-values >0.5). Individuals caught on the south-facing slope were larger than those caught on the north-facing slope. Reproductively mature bank vole males were smaller than females, whereas reproductively mature grey-sided vole males were larger than females. Body mass was related to density in both species. In bank voles, we found a direct positive density dependence caused by a higher rate of survival at higher densities resulting from individual allocation of resources from reproduction to survival and growth. In grey-sided voles, we found a negative delayed density dependence resulting from grazing on preferred plants that determined the resource available for individual vole growth the following year.     

田鼠种群波动的原因和时间

本文总结了橙腹田鼠（Microtus ochrogaster）和草原田鼠（M．permsylvanicus）25年的种群统计学研究结果和结论。探讨了田鼠种群波动周期性、诱发种群波动以及导致波动期间峰值变异的因素。并对种群存活值和繁殖活动的作用进行了分析和评价。根据两种田鼠种群波动周期性、波动峰值出现的时间以及特定年份峰值的高度等特征,证明两物种波动均具有不稳定性。两种田鼠存活值的变化是由特定年份是否发生波动以及波动峰值出现的时间决定。增加初始阶段的种群密度及时间长度是造成两种动物种群波动峰值不同的主要原因。橙腹田鼠种群停止增长的原因是存活值降低,而草原田鼠则是繁殖活动减少。据推测,与种群波动初始密度相关的种群死亡率的差异是由捕食者的净效应（Net effect）决定的,调控两种群密度的因素均为非密度的其它生态学因子。由于特定年份田鼠种群捕食压力的不确定性,导致了橙腹田鼠和草原田鼠种群波动的不稳定性。     

Spatiotemporal dynamics of Puumala hantavirus in suburban reservoir rodent populations

The transmission of pathogens to susceptible hosts is dependent on the vector population dynamics. In Europe, bank voles (Myodes glareolus) carry Puumala hantavirus, which causes nephropathia epidemica (NE) in humans. Fluctuations in bank vole populations and epidemics in humans are correlated but the main factors influencing this relationship remain unclear. In Belgium, more NE cases are reported in spring than in autumn. There is also a higher incidence of human infections during years of large vole populations. This study aimed to better understand the link between virus prevalence in the vector, vole demography, habitat quality, and human infections. Three rodent populations in different habitats bordering Brussels city, Belgium, were studied for two years. The seroprevalence in voles was influenced first by season (higher in spring), then by vole density, vole weight (a proxy for age), and capture site but not by year or sex. Moreover, voles with large maximal distance between two captures had a high probability for Puumala seropositivity. Additionally, the local vole density showed similar temporal variations as the number of NE cases in Belgium. These results showed that, while season was the main factor influencing vole seroprevalence, it was not sufficient to explain human risks. Indeed, vole density and weight, as well as the local habitat, were essential to understanding the interactions in these host‐pathogen dynamics. This can, in turn, be of importance for assessing the human risks.     

Synchrony in short-term fluctuations of moose calf body mass and bank vole population density supports the mast depression hypothesis

Inter-annual variations in body mass of moose, Alces alces , in Norway and Sweden have been considered as most likely due to direct or indirect effects of weather, but so far predictions of autumn body mass of moose calves on the basis of weather data have given a poor fit to data. A striking, but hitherto unnoticed, feature of several time series on body mass of moose calves from south-eastern Norway is an apparently regular 3–4-year fluctuation pattern. This short-term fluctuation could be due to regular variations in forage quality, e.g. caused by a cyclic seed production of some important food plants, as envisaged by the "mast depression" hypothesis. One plant species important as food for moose calves in autumn is bilberry, Vaccinium myrtillus , which usually produces high seed crops (masts) at intervals of 3–4 years. Populations of the bank vole, Clethrionomys glareolus, which feeds on bilberry shoots in winter, are known to peak in bilberry post-mast years. In two study areas in Norway, there was a positive correlation between the autumn body mass of moose calves and the autumn population index of bank vole in the succeeding year. In the northern area there was an additional positive effect of summer precipitation, whereas in the southern area there was an additional negative effect of summer temperature. In both areas, however, the effect of weather was less pronounced than that of the bank vole index. These results support the mast depression hypothesis.     

Temperature thresholds of physically dormant seeds and plant functional response to fire: variation among species and relative impact of climate change

Variation in dormancy thresholds among species is rarely studied but may provide a basis to better understand the mechanisms controlling population persistence. Incorporating dormancy‐breaking temperature thresholds into existing trait frameworks could improve predictions regarding seed bank persistence, and subsequently species resilience in response to fire, climate change and anthropogenic management. A key ecological strategy for many species from fire‐prone ecosystems is the possession of a long‐lived seed bank, ensuring recovery after fire. Physical dormancy is dominant in these ecosystems and maintaining this dormancy is directly linked to seed bank persistence. We identified a suite of seed‐related factors relevant to maintaining populations in fire‐prone regions for 14 co‐occurring physically dormant species. We measured variation in initial levels of dormancy and then applied experimental heating treatments, based on current seasonal temperatures and those occurring during fires, to seeds of all study species. Additionally, higher seasonal temperature treatments were applied to assess response of seeds to temperatures projected under future climate scenarios. Levels of germination response and mortality were determined to assess how tightly germination response was bound to either fire or seasonal cues. Six species were found to have dormancy cues bound to temperatures that only occur during fires (80°C and above) and were grouped as having obligate pyrogenic dormancy release. The remaining species, classified as having facultative pyrogenic dormancy, had lower temperature dormancy thresholds and committed at least 30% of seeds to germinate after summer‐temperature treatments. Evidence from this study supports including dormancy‐breaking temperature thresholds as an attribute for identifying functional types. High temperature thresholds for breaking dormancy, found in our obligate pyrogenic group, appear to be a fire‐adapted trait, while we predict that species in the facultative group are most at risk to increased seed bank decay resulting from elevated soil temperatures under projected climate change.     

Size traits and site conditions determine changes in seed bank structure caused by grazing exclusion in semiarid annual plant communities

1. Contrasting patterns of change in the seed bank of natural grasslands are frequently found in response to grazing by domestic herbivores. Here, we studied the hypotheses that a) patterns of change in seed bank density and composition in response to grazing depend on spatial variation in resource availability and productivity, and b) that variation among species in patterns of seed bank response to grazing is linked to differences in species size traits (i.e. size of plant, dispersal unit and seed). 2. Effects of sheep grazing exclusion on the seed bank were followed during five years in a semiarid Mediterranean annual plant community in Israel. Seed bank density and composition were measured in autumn, before the rainy season, inside and outside fenced exclosures in four neighboring topographic sites differing in vegetation characteristics, soil resources and primary productivity: Wadi (dry stream terraces, high productive site), Hilltop, South‐ and North‐facing slopes (less productive sites). 3. Topographic sites differed in seed density (range ca 2500–18000 seed m^?2) and in seed bank response to grazing exclusion. Fencing increased seed density by 78, 51 and 18% in the Wadi, South‐ and North‐facing slopes, respectively, but had no effect in the Hilltop. At the species level, grazing exclusion interacted with site conditions in determining species seed bank density, with larger or opposite changes in the high productive Wadi compared to the other less productive sites. 4. Changes in seed bank structure after grazing exclusion were strongly related to species size traits. Grazing exclusion favored species with large size traits in all sites, while seed density of tiny species decreased strongly in the high productive Wadi. Species with medium and small size traits showed lesser or no responses. 5. The size of plants, dispersal units and seeds were strongly correlated to each other, thus confounding the evaluation of the relative importance of each trait in the response of species to grazing and site conditions. We propose that the relative importance of plant size vs seed size in the response to grazing changes with productivity level.