Spatio-temporal variation in a seed bank of a semi-arid region in northeastern Brazil

This study aimed to evaluate variations in the seed bank within a 3-year temporal series in order to answer the following questions: 1) Does the seed bank's species richness and seed density differ among climatic seasons and between years? 2) Are there differences in the richness and density of seed banks between the litter and mineral soil? 3) Can the seed bank's species richness and seed density be explained by characteristics such as the previous year's precipitation and soil depth (litter or mineral soil)? The samples were collected from litter and mineral soil (0–5 cm), in 210 sub-plots, during the dry and rainy seasons of each year (August 2005 through February 2008). Overall, 79 species were recorded. On average, 1 168, 304 and 302 seeds.m⁻² were recorded in the seed bank during years I, II and III, respectively. This study showed that the Caatinga's seed bank is rich in herbaceous species, yet species' density and richness are low in the litter. Furthermore, about 43% of the variation in species richness and density was explained by soil depth (litter and mineral soil) and previous years' rainfall.     

Impact of herbivores on nitrogen cycling: contrasting effects of small and large species

Herbivores are reported to slow down as well as enhance nutrient cycling in grasslands. These conflicting results may be explained by differences in herbivore type. In this study we focus on herbivore body size as a factor that causes differences in herbivore effects on N cycling. We used an exclosure set-up in a floodplain grassland grazed by cattle, rabbits and common voles, where we subsequently excluded cattle and rabbits. Exclusion of cattle lead to an increase in vole numbers and a 1.5-fold increase in net annual N mineralization at similar herbivore densities (corrected to metabolic weight). Timing and height of the mineralization peak in spring was the same in all treatments, but mineralization in the vole-grazed treatment showed a peak in autumn, when mineralization had already declined under cattle grazing. This mineralization peak in autumn coincides with a peak in vole density and high levels of N input through vole faeces at a fine-scale distribution, whereas under cattle grazing only a few patches receive all N and most experience net nutrient removal. The other parameters that we measured, which include potential N mineralization rates measured under standardized laboratory conditions and soil parameters, plant biomass and plant nutrient content measured in the field, were the same for all three grazing treatments and could therefore not cause the observed difference. When cows were excluded, more litter accumulated in the vegetation. The formation of this litter layer may have added to the higher mineralization rates under vole grazing, through enhanced nutrient return through litter or through modification of microclimate. We conclude that different-sized herbivores have different effects on N cycling within the same habitat. Exclusion of large herbivores resulted in increased N annual mineralization under small herbivore grazing.     

The impact of sarcoptic mange Sarcoptes scabiei on the British fox Vulpes vulpes population

• 1 Disease epizootics can significantly influence host population dynamics and the structure and functioning of ecological communities. Sarcoptic mange Sarcoptes scabiei has dramatically reduced red fox populations Vulpes vulpes in several countries, including Britain, although impacts on demographic processes are poorly understood. We review the literature on the impact of mange on red fox populations, assess its current distribution in Britain through a questionnaire survey and present new data on resultant demographic changes in foxes in Bristol, UK.
• 2 A mange epizootic in Sweden spread across the entire country in < 10 years resulting in a decline in fox density of up to 95%; density remained lowered for 15–20 years. In Spain, mange has been enzootic for > 75 years and is widely distributed; mange presence was negatively correlated with habitat quality.
• 3 Localized outbreaks have occurred sporadically in Britain during the last 100 years. The most recent large‐scale outbreak arose in the 1990s, although mange has been present in south London and surrounding environs since the 1940s. The questionnaire survey indicated that mange was broadly distributed across Britain, but areas of perceived high prevalence (> 50% affected) were mainly in central and southern England. Habitat type did not significantly affect the presence/absence of mange or perceived prevalence rates. Subjective assessments suggested that populations take 15–20 years to recover.
• 4 Mange appeared in Bristol's foxes in 1994. During the epizootic phase (1994–95), mange spread through the city at a rate of 0.6–0.9 km/month, with a rise in infection in domestic dogs Canis familiaris c. 1–2 months later. Juvenile and adult fox mortality increased and the proportion of females that reproduced declined but litter size was unaffected. Population density declined by > 95%.
• 5 In the enzootic phase (1996–present), mange was the most significant mortality factor. Juvenile mortality was significantly higher than in the pre‐mange period, and the number of juveniles classified as dispersers declined. Mange infection reduced the reproductive potential of males and females: females with advanced mange did not breed; severely infected males failed to undergo spermatogenesis. In 2004, Bristol fox population density was only 15% of that in 1994.

     

A test of Brown's principle in the intertidal limpet Collisella scabra (Gould, 1846)

Aim Brown's principle predicts that a species will peak in density near its range centre, and decline gradually towards the margins of its geographical distribution. The decline is assumed to reflect a decrease in individual performance near range margins. I test this abundance–performance hypothesis by comparing patterns in density and size across the northern half of the geographical distribution of the marine patellogastropod Collisella scabra (Gould, 1846). Location Collisella scabra is a high intertidal patellogastropod species distributed along the Pacific coast of North America from Cape Mendocino (CA, USA) to southern Baja California (Mexico). I surveyed 11 research sites spanning c. 36–44° N. Methods In each of the 11 research sites I surveyed four distinct microhabitats, and compared spatial patterns in density and in the size of solitary limpets. Results Both density and size were highly variable across the species range. Density peaked near the northern range margin, and showed greater variance at small spatial scales (< 10 km) than at large scales (> 100 km). In contrast, large size occurred uniformly across the survey area, and size was strongly associated with microhabitat. Main conclusion Collisella scabra does not show spatial patterns of density or performance that are consistent with Brown's principle. The underlying assumptions of Brown's principle may conflict with specific characteristics of C. scabra's life history and/or patterns of environmental variation across its range. Because such conflicts may be common in a large number of marine and terrestrial species, the generality of Brown's principle is questioned.     

Impact of management treatments on waterfowl use of dense nesting cover in the Canadian parklands

Periodic treatment of established stands of dense nesting cover (DNC) is a recommended practice to maintain cover quality, but little information exists on the magnitude and duration of treatment effects on nesting waterfowl. During 1998–2001, we examined the effect of management treatments on vegetative characteristics and waterfowl nest success and density in fields of DNC seeded to introduced and native grass and forb mixes in the parklands of Saskatchewan and Manitoba. We measured vegetation height–density and litter depth within fields and located and monitored 1,927 duck nests within 33–42 fields/yr ranging in size from 6 ha to 62 ha. We considered a series of models examining the influence of grass type and management treatment (GTMT) and years post-management (YPM) on vegetative characteristics, nest success, and nest density while including covariates potentially affecting these response variables. Visual obstruction and litter depth were lowest in native-burned fields and greatest in introduced-hayed fields. Visual obstruction was low the year following management, peaked 2–3 YPM, and remained at intermediate levels through ≥6 YPM. Litter depth remained low for the first 3 YPM and increased thereafter. Nest success and nest density varied little among GTMT. Nest success was high (14.3%) the year following a management treatment, low (6.5%) at 2 YPM, and moderate thereafter. Nest success decreased with percent cropland in the surrounding landscape. Nest density was 0.7 nests/ha the first year following management, increased to approximately 1.3 nests/ha in years 2–3, and declined back to approximately 0.7 nests/ha for ≥6 YPM. Nest density decreased with field size and increased with the area of small wetlands, percent cropland, and percent wetland within surrounding landscapes. Nest density tracked vegetation density as expected and our results indicate a possible trade-off between nest density and nest success. Given ancillary data on small mammal and insect prey in our study fields, and evidence from other studies, we speculate that DNC fields may act as prey reservoirs during years of peak vegetative density with a consequent reduction in nest survival. Therefore, management to increase waterfowl production based on our results needs to consider the interaction of treatment effects, competing habitats, and surrounding landscape composition. © 2011 The Wildlife Society.     

Small stonefly predators affect microbenthic and meiobenthic communities in stream leaf packs

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Litter size variation in hypothalamic gene expression determines adult metabolic phenotype in Brandt's voles (Lasiopodomys brandtii)

Background

Early postnatal environments may have long-term and potentially irreversible consequences on hypothalamic neurons involved in energy homeostasis. Litter size is an important life history trait and negatively correlated with milk intake in small mammals, and thus has been regarded as a naturally varying feature of the early developmental environment. Here we investigated the long-term effects of litter size on metabolic phenotype and hypothalamic neuropeptide mRNA expression involved in the regulation of energy homeostasis, using the offspring reared from large (10–12) and small (3–4) litter sizes, of Brandt''s voles (Lasiopodomys brandtii), a rodent species from Inner Mongolia grassland in China.

Methodology/Principal Findings

Hypothalamic leptin signaling and neuropeptides were measured by Real-Time PCR. We showed that offspring reared from small litters were heavier at weaning and also in adulthood than offspring from large litters, accompanied by increased food intake during development. There were no significant differences in serum leptin levels or leptin receptor (OB-Rb) mRNA in the hypothalamus at weaning or in adulthood, however, hypothalamic suppressor of cytokine signaling 3 (SOCS3) mRNA in adulthood increased in small litters compared to that in large litters. As a result, the agouti-related peptide (AgRP) mRNA increased in the offspring from small litters.

Conclusions/Significance

These findings support our hypothesis that natural litter size has a permanent effect on offspring metabolic phenotype and hypothalamic neuropeptide expression, and suggest central leptin resistance and the resultant increase in AgRP expression may be a fundamental mechanism underlying hyperphagia and the increased risk of overweight in pups of small litters. Thus, we conclude that litter size may be an important and central determinant of metabolic fitness in adulthood.     

Growth and survival of larvae of Thaumetopoea pinivora inside and outside a local outbreak area

1 Outbreaks of herbivorous insects tend to be spatially restricted, possibly because of demographic differences between inside and outside the outbreak area. In some cases, the margin of the outbreak area is distinct, allowing comparisons of adjacent areas that may identify factors leading to such differences in abundance. The northern pine processionary moth Thaumetopoea pinivora presently occurs at outbreak densities within a well‐defined area of approximately 3000 ha on the island of Gotland, south Sweden. We investigated whether cohorts of young larvae (first and second instar) had higher growth rate and survival inside or outside the outbreak area. 2 Group‐feeding appears to promote outbreaks in certain insect groups. Because T. pinivora larvae are highly social, we also compared larval performance between groups of different sizes inside and outside of the outbreak area: ‘small’ (33 eggs/group) and ‘normal’ (100 eggs/group). 3 Averaged over group size, whole colony mortality through the first two instars was two‐fold higher in the non‐outbreak area compared with the outbreak area. Mortality of individual larvae in the surviving colonies, however, did not differ between the two areas. There were only small differences in food quality (toughness, nitrogen content) between the areas, with no detectable effects on larval performance. 4 Larval relative growth rate did not differ between reduced and normal‐sized groups, which is surprising given that growth rate is known to increase with group size in other group‐feeding lepidopterans. 5 Reduced group size negatively affected larval survival, particularly in the outbreak area; by contrast, normal‐sized groups survived equally well in the two areas. Wood ants (Formica spp.) were more common outside the outbreak area, and appeared to be the main cause of colony mortality at low larval density. A different result was observed with regard to per‐capita mortality, which was higher in the outbreak area. We speculate that this could have been due to solitary predators being locally specialized on T. pinivora in the high‐density area.     

Relations among birth condition, maternal condition, and postnatal growth in captive common marmoset monkeys (Callithrix jacchus)

The present study characterizes the relations among maternal condition, litter size, birth condition, and growth in body weight for a population of common marmosets. The subjects of the study were marmosets born into a single colony between 1994 and 2001. Three sets of analyses were conducted to answer the following questions: 1) Is there a relationship between litter size, maternal condition, and birth condition? In the study population, maternal body weight, maternal age, litter size, and birth condition were related in a complex fashion. Birth weight and prenatal long‐bone growth, as reflected in knee–heel length, were both related to maternal age, with older mothers supporting higher prenatal growth. Age and maternal condition appeared to interact as determinants of long‐bone growth, as the combination of older and larger mothers resulted in significantly longer knee–heel lengths in their offspring. 2) Is there a relationship between birth condition or maternal condition and subsequent growth or final adult size? The early growth rate in this population was similar to early growth rates reported for three different marmoset colonies, suggesting that early growth may be relatively inflexible in this species. However, within this population, the variation that did occur in early growth rate was related to birth weight and maternal weight. Later growth and adult weight were related to birth weight and litter size: small twin infants displayed slower later growth rates and were smaller as adults than twins that began life at a higher birth weight, while the birth weight of triplets was not related to adult size. In these marmosets, small infants that were the result of increased litter size differed from small infants whose small birth size resulted from other factors. This reinforces the proposal that the causes of low birth weight will be relevant to the development of the marmoset as a model of prenatal environmental effects. Am. J. Primatol. 62:83–94, 2004. © 2004 Wiley‐Liss, Inc.     

Changes in the kinetics of follicular growth in response to selection for large litter size in mice

The present study examined a randomly selected control line (C) and a large litter size-selected line of mice (S1) to determine changes in the kinetics of follicular growth that have occurred in response to selection for large litter size. For each follicle type (FT), the number of healthy and atretic follicles, length of components of granulosa cell cycle, follicular growth rate, and follicular flux were determined microscopically from serially sectioned ovaries of Lines C and S1 mice. Selection for litter size significantly increased the number of small and medium, and some large follicle-size classes. While selection for litter size did not change the overall incidence of atresia at proestrus, it did decrease the incidence of atresia in the large Type 7 follicles by 19%. Selection for litter size also increased ovarian weight at proestrus. Selection for litter size increased the rate of growth through FT 3a, 5a, and 5b, and reduced the time required for follicles to grow from primordial to Graafian follicles from 39.1 days in Line C to 33.4 days in Line S1. Selection for large litter size also increased the flux of follicles through follicle Types 3a to 5b by 72%, and through follicle Types 6 and 7 by 21%. Genetic variation was found in many aspects of the kinetics of follicular growth.     

HOW MAMMALS PRODUCE LARGE-BRAINED OFFSPRING

Two explanations for species differences in neonatal brain size in eutherian mammals relate the size of the brain at birth to maternal metabolic rate. Martin (1981, 1983) argued that maternal basal metabolic rate puts an upper bound on the mother's ability to supply energy to the fetus, thereby limiting neonatal brain size. Hofman (1983) proposed that gestation length in mammals is constrained by maternal metabolic rate, implying an indirect constraint on neonatal brain size. Since individuals of precocial species have much larger neonatal brain sizes and are gestated longer for a given maternal body size than individuals of altricial species, Martin's and Hofman's ideas also require that mothers of precocial offspring have higher metabolic rates for their body sizes than mothers of altricial offspring. Data on 116 mammal species from 13 orders show that neither neonatal brain size nor gestation length is correlated with maternal metabolic rate when maternal body-size effects are removed. For a given maternal size, there is no difference in metabolic rates between precocial and altricial species, despite a two-fold difference between them in average neonatal brain size. However, neonatal brain size is strongly correlated with gestation length and litter size, independently of maternal size and metabolic rate. Analyses conducted within orders replicated the findings for gestation length and suggested that neonatal brain size may be at best only weakly related to metabolic rate. Differences in neonatal brain size appear to have evolved primarily with species differences in gestation length and litter size but not with differences in metabolic rate; large-brained offspring are typically produced from litters of one that have been gestated for a long time relative to maternal size. We conclude that species differences in relative neonatal brain size reflect different life-history tactics rather than constraints imposed by metabolic rate.     

A note on the relationship between outbreaks of the oriental tussock moth,Euproctis flava and weather conditions

A relationship between outbreaks of the oriental tussock moth, Euproctis flava, in Akita Prefecture and weather conditions have been analyzed in the present paper. The results obtained are summarized as follows: Outbreaks were, in most cases, observed at intervals of 7–9 years, and the population fluctuation was considered to be regular, when small rises of population density were taken into consideration. The condition of low temperature and decreased percentage of sunny hours in spring and summer has a great influence upon the occurrence of outbreaks, and a big outbreak seems to come when a peak of the intrinsic population fluctuation occurs just simultaneously with such a weather condition.     

Ramet size equalisation in a clonal plant,Phragmites australis

The influence of shading from older generations of dead culms (standing litter) on density, growth rate and development of size structure at the ramet level was investigated in a pure stand of Phragmites australis by experimental neutral shading of plots after removal of standing litter. Initial differences in height distribution between autumn and spring cohorts disappeared in the course of shoot growth. The Gini coefficients of shoot heights and estimated shoot weights indicated that the size structure of the shoots became more equal with increasing mean size in both shaded and unshaded plots. Relative growth rate for height (RHGR) and weight of individual shoots was negatively related to shoot size during the early and presumably storage-dependent growth period, suggesting a strong support for growth of smaller shoots. No etiolation was indicated by mean or maximum height in shaded and unshaded plots, or by the relationship between shoot height and weight. Mean shoot density was significantly lower in shaded than in unshaded plots in one of two shade treatment years. A regression model indicated a small but significant effect of shoot density on the approximately linear relationship between RHGR and the logarithm of height. The growth rate of small shoots was slightly larger at low than at high shoot density. Therefore, it is suggested that the shade from standing litter in P. australis stands can decrease shoot natality in the spring cohort, and thereby increase the support to fewer small shoots.     

Modelling the use of NPV for the biological control of Asian gypsy moth Lymantria dispar invading New Zealand

1 The accidental introduction of the Asian strain of gypsy moth (AGM) Lymantria dispar (L.) to New Zealand poses a major threat to New Zealand's forestry industry. To aid eradication and control decisions in the event of its establishment, a model was developed for the effect of nuclear polyhedrosis virus (NPV) as biological control for AGM in New Zealand. 2 The model simulates within‐ and between‐season gypsy moth population dynamics, including temperature‐dependent development, density dependence through predation and resource limitation, and interactions with NPV. 3 Following its introduction to New Zealand, AGM is predicted to increase more quickly and have more severe outbreaks than the European strain. In the absence of predators, the model predicts initial outbreaks then damped oscillations to an equilibrium. 4 In the model, a single application of NPV (2500 GPIB ha^?1) at the time of maximum larval density gave up to 80% suppression of peak larval densities in the following year. The same level of suppression was achieved in the absence of predators. 5 In the long term, the model predicted that spraying when an outbreak was just beginning gave best results (a 50–70% reduction of the following two outbreaks). Simulation of threshold spraying resulted in NPV application, on average, every 7 years and suppression of outbreak densities by 40–70%. Following a single application, NPV was maintained in the population as a classical biological control agent, giving approximately a 20% reduction in outbreak densities. 6 Eradication of AGM using NPV was possible if larval densities were very low.     

Primate life histories

An animal's life history can be summarized by key variables that account for its life course from conception to death. Biological parameters that are of interest relate to reproductive effort and developmental rates (e.g., gestation length, neonatal weight, prenatal and postnatal growth rates, weaning age, and weaning weight) and the rate of reproduction (e.g., age at first and last reproduction, interbirth interval, the number of offspring per litter, birth rate, and the intrinsic rate of natural increase [r_max]). The rather obvious fact that such variables differ from species to species and from individual to individual has been the subject of much interest since the late 1960s, following the observation that species seem to be arranged in a spectrum that ranges from small animals that breed rapidly and develop early, have many young per litter, and have short lives, to large animals that breed slowly and develop late, have few young per litter, and have long lives. © 1998 Wiley-Liss, Inc.     

Effects of agricultural development on processing of tussock leaf litter in high country New Zealand streams

1. We compare the rates and mechanisms of processing of tussock (Chionochloa spp.) leaf litter in six New Zealand streams draining grassland catchments that contrast in the extent to which they have been developed for pasture. 2. Rates of processing, measured as rate of weight loss of leaf packs and rate of leaf softening, were at the slow end of the spectrum for vascular plant processing. Processing was faster at developed sites, mediated mainly through the influence of oxidized nitrogen concentration on microbial activity. 3. Few invertebrate shredders colonized leaf packs and it is unlikely that invertebrates had an appreciable effect on leaf processing in our study streams, which do not effectively retain leaf litter. Very small headwater tributaries appear to retain leaf litter and possess a more abundant shredder community. 4. Measures of leaf processing in our six streams were significantly correlated with Petersen's (1992) RCE score of stream condition. We discuss the potential for using rate of leaf litter processing as a method of bioassessment. 5. Even the most degraded stream in our study is classed as ‘good’ using the RCE inventory system. Human impact in the Taieri River is relatively small compared with the degradation observed in some parts of the world.     

DIET AND DISPERSION OF THE GREAT ARGUS ARGUSIANUS ARGUS

The food and dispersion of Great Argus were studied in two West Malaysian forested areas from 1975 to 1978. Invertebrates on the litter surface lived at low density and were typically large and solitary; those beneath the litter were abundant but typically tiny and colonial. Great Argus feed mainly on fallen fruit but rely for protein on the large, solitary litter-surface invertebrates and were never seen scratching for food. Adult males maintain small, discrete home ranges in which they forage alone. Much of their time is spent resting, which reduces energy expenditure. Females also forage alone and may be similarly inactive. Great Argus are very large and their requisite large invertebrate prey cannot be obtained beneath the litter; scratching has been eliminated as a foraging technique. A strategy of litter-surface searching necessitates solitariness, to avoid intraspecific feeding competition. The large size of Great Argus is related to a low rate of energy turnover and energy conservation by prolonged resting. Both sexes are probably territorial during the breeding season and larger territories then may limit population density.     

Mammalian reproductive strategies: A generalized relation of litter size to body size

Summary A generalized relationship of litter size to mammalian body size was predicted by a graph model. The model was used to generate hypotheses explaining specific features of variation in gestation time, relative litter weight, birth weight, and reproductive capacity. The predictions were tested by means of data from the literature.Mammals were assumed to maximize neonatal survival of offspring to the limits allowed by litter weight per female body weight. Gestation time correlated negatively with the foetal growth rate of relative litter weight. Gestation time did not correlate with the foetal growth rate of individual offspring.Relative litter weight correlated negatively with adult body weight. This relationship was explained by the higher assimilation rate per unit weight relative to metabolic rate in small mammals.Birth weight correlated positively with body weight. However, small mammals produce larger offspring than predicted by the linear relationship of birth weight to body weight in large mammals. There is obviously a minimum birth weight which cannot be decreased without special arrangements for parental care.The prediction of the relationship of litter size to body size was derived from the relations of relative litter weight and birth weight to body weight. In small mammals (less than 1 kg) litter the correlation was negative. When litter size was compared with body length, the correlation was positive in small mammals (less than 30 cm) and negative in large mammals. In both sets of data there was a negative overall correlation between litter size and body size.Reproductive capacity, defined as the number of offspring per season, correlated negatively with life-span.     

东方田鼠种群密度制约的迟滞效应

本研究中用成年东方田鼠不同时间不同密度笼养后（笼内雌雄各半）,低密度配对饲养,观察各阶段东方田鼠繁殖指标的差异。试鼠144只,处理时分为LL（低密度长时间）（2只/笼,共20笼,90d）、HL（高密度长时间）（8只/笼,共5笼,90d）、HM（高密度中等时间）（8只/笼,共4笼,20d）与HS（高密度短时间）（8只/笼,共4笼,10d）4组。处理后在低密度条件下观察繁殖情况（胎仔数、怀孕率、分娩频率以及产仔间隔等）,直至180 d。实验数据按时间划分为3个部分统计：0~90 d为第一阶段(不同密度处理期),90~109 d为过渡阶段（处理后的过渡期）,109 d以后为第二阶段（正常低密度配对繁殖期）。结果表明：经过长时间高密度处理后,平均产仔数显著减少,组间平均分娩频率无显著差异。怀孕率由HL组到HM组,再到LL和HS组显著增加。在第一阶段,除了LL组之外,其余各组都未发现繁殖。过渡阶段的怀孕率组间差异显著,LL组以及HM组显著高于HL组。第二阶段的怀孕率以及胎仔数的组间差异显著,都为LL、HS两组较高,HL、HM两组较低。平均分娩频率、平均胎仔数以及产仔间隔各组间无显著差异。结论：不同持续时间的密度效应有较大差别。对于东方田鼠,20d的高密度处理相比于10d更能对其繁殖起到抑制作用。恢复低密度后,存在种群的迟滞性密度制约。     

Life history traits in a cyclic ecosystem: a field experiment on the arctic fox

The reproduction of many species depends strongly on variation in food availability. The main prey of the arctic fox in Fennoscandia are cyclic small rodents, and its number of litters and litter size vary depending on the phase of the rodent cycle. In this experiment, we studied if the arctic fox adjusts its reproduction as a direct response to food abundance, in accordance with the food limitation hypothesis, or if there are additional phase-dependent trade-offs that influence its reproduction. We analysed the weaning success, i.e. proportion of arctic fox pairs established during mating that wean a litter in summer, of 422 pairs of which 361 were supplementary winter fed, as well as the weaned litter size of 203 litters of which 115 were supplementary winter fed. Females without supplementary winter food over-produced cubs in relation to food abundance in the small rodent increase phase, i.e. the litter size was equal to that in the peak phase when food was more abundant. The litter size for unfed females was 6.38 in the increase phase, 7.11 in the peak phase and 3.84 in the decrease phase. The litter size for supplementary winter-fed litters was 7.95 in the increase phase, 10.61 in the peak phase and 7.86 in the decrease phase. Thus, feeding had a positive effect on litter size, but it did not diminish the strong impact of the small rodent phase, supporting phase-dependent trade-offs in addition to food determining arctic fox reproduction.