Life history patterns in female moose (Alces alces): the relationship between age,body size,fecundity and environmental conditions

I examined the relationship between age, body size and fecundity in 833 female moose (Alces alces) from 14 populations in Sweden sampled during 1989–1992. Data on population density, food availability and climatic conditions were also collected for each population. Age and body mass were both significantly positively related to fecundity, measured as ovulation rate, among female moose. The relationship between the probability of ovulation and body mass was dependent on age with (1) a higher body mass needed in younger females for attaining a given fecundity, and (2) body mass having a stronger effect on fecundity in yearling (1.5 year) than in older (2.5 year) females. Thus, a 40 kg increase in yearling body mass resulted in a 42% increase in the probability of ovulation as compared to a 6% increase in older females. The lower reproductive effort per unit body mass, and the relatively stronger association between fecundity and body mass in young female moose compared to older ones, is likely to primarily represent a mechanism that trades off early maturation against further growth, indicating a higher cost of reproduction in young animals. In addition to age and body mass, population identity explained a significant amount of the individual variation in fecundity, showing that the relationship between body mass and fecundity was variable among populations. This variation was in turn related to the environment, in terms of climatic conditions forcing female moose living in relatively harsh/more seasonal climatic conditions to attain a 22% higher body mass to achive the same probability of multiple ovulation (twinning) as females living in climatically milder/less seasonal environments. The results suggests that the lower fecundity per unit body mass in female moose living in climatically harsh/more seasonal environments may be an adaptive response to lower rates of juvenile survival, compared to females experiencing relatively milder/less seasonal climatic conditions.     

Effects of range position,inter-annual variation and density on demographic transition rates of <Emphasis Type="Italic">Hornungia petraea</Emphasis> populations

The central-marginal model assumes unfavourable and more variable environmental conditions at the periphery of a species’ distribution range to negatively affect demographic transition rates, finally resulting in reduced population sizes and densities. Previous studies on density-dependence as a crucial factor regulating plant population growth have mainly focussed on fecundity and survival. Our objective is to analyse density-dependence in combination with the effect of inter-annual variation and range position on all life stages of an annual plant species, Hornungia petraea, including germination and seed incorporation into the seed bank. As previous studies on H. petraea had revealed a pattern opposite to existing theory with lower population densities at the distribution centre in Italy than at the periphery in Germany, we hypothesised that (1) demographic transition rates are lower, (2) the inter-annual variation in demographic transition rates is higher and (3) the intensity of density-dependence is weaker in Italy than in Germany. To analyse demographic transition rates, we used an autoregressive covariance strategy for repeated measures including density and inter-annual variation. All the three hypotheses were confirmed, but the impact of range position, density-dependence and inter-annual variation differed among the transition steps. All transition rates except fecundity were higher in the German populations than in the Italian populations. Germination rate and incorporation rate into the seed bank were strongly density-dependent. Central populations showed a larger inter-annual variation in fecundity and winter survival rate. Winter survival rate was the only transition step with a stronger density-dependence in peripheral populations. In most cases, these differences between distribution centre and periphery would not have emerged without taking density-dependence and inter-annual variation into account. We conclude that including range position, inter-annual variation and density-dependence in one single statistical model is an important tool for the interpretation of demographic patterns regarding the central-marginal model. Electronic Supplementary Material Supplementary material is available for this article at     

DEVELOPMENTAL STABILITY IN LEAVES OF CLARKIA TEMBLORIENSIS (ONAGRACEAE) AS RELATED TO POPULATION OUTCROSSING RATES AND HETEROZYGOSITY

Four natural populations of Clarkia tembloriensis, whose levels of heterozygosity and rates of outcrossing were previously found to be correlated, are examined for developmental instability in their leaves. From the northern end of the species range, we compare a predominantly selfing population (t? = 0.26) with a more outcrossed population (t? = 0.84), which is genetically similar. From the southern end of the range, we compare a highly selfing population (t? = 0.03) with a more outcrossed population (t? = 0.58). We measured developmental stability in the populations using two measures of within-plant variation in leaf length as well as calculations of fluctuating asymmetry (FA) for several leaf traits. Growth-chamber experiments show that selfing populations are significantly more variable in leaf length than more outcrossed populations. Developmental instability can contribute to this difference in population-level variance. Plants from more homozygous populations tend to have greater within-plant variance over developmentally comparable nodes than plants from more heterozygous populations, but the difference is not significant. At the upper nodes of the plant, mature leaf length declines steadily with plant age, allowing for a regression of leaf length on node. On average, the plants from more homozygous populations showed higher variance about the regression (MSE) and lower R² values, suggesting that the decline in leaf length with plant age is less stable in plants from selfing populations than in plants from outcrossing populations. Fluctuating asymmetry (FA) was calculated for four traits within single leaves at up to five nodes per plant. At the early nodes of the plant where leaf arrangement is opposite, FA was also calculated for the same traits between opposite leaves at a node. Fluctuating asymmetry is significantly greater in the southern selfing population than in the neighboring outcrossed population. Northern populations do not differ in FA. Fluctuating asymmetry can vary significantly between nodes. The FA values of different leaf traits were not correlated. We show that developmental stability can be measured in plants using FA and within-plant variance. Our data suggest that large differences in breeding system are associated with differences in stability, with more inbred populations being the least stable.     

Morphological traits and common illness in newborn infants

The present study focussed on neonates morbidity in connection with the morphological traits in the individual relative to the population average and in connection with the fluctuating asymmetry (FA) in 8 bilateral traits. The sample comprised two categories of newborn infants: 1) 1088 full term children, in which body weight (W), body height (H) and head circumference (HC) were measured on the day of birth; and 2) 216 infants of various gestation age (from 26 to 41 weeks) in which 6 measurements of upper and lower arm breadth, plus ear length and breadth on both sides of the body, were made during the first 72 hours of life. Relevant vital statistics were also obtained from parents. The following major categories of morbidity were observed in our samples: jaundice, respiratory diseases, cardiovascular diseases, urinary tract infections (only in the second sample), haematological diseases and ‘varia’. The results show that W, H, HC and W/H² index are significantly correlated with a predisposition to various categories of morbidity. For W and W/H² index, this correlation remains significant even after adjustment for diverse anthropometric and demographic variables. In the case of urinary tract infection, the contribution of HC also remains significant even after the adjustment procedure. Gestational age and cardiovascular illnesses yielded significant correlations with mean FA. There was an evident decrease in mean FA upon increase of gestational age. The mean FA of infants suffering from cardiovascular morbidity displays the same trend, although it is consistently higher compared to the total group of infants in the respective gestational age.     

Macro-alga population shows low but significant heterogeneity in developmental instability with no detectable association with individual fitness

We studied among-individual variation in developmental instability (DI) and fitness-related parameters in 80 individual plants of Fucus vesiculosus (Phaeophyta). To minimize differential environmental effects, plants were sampled from one environmentally homogenous population. DI was measured as fluctuating asymmetry (FA) of four bilaterally symmetric traits (branch length, receptacle length and width, and bladder width) for an average total of 30 structures per individual. FA levels varied significantly among individual plants, consistent with a coefficient of variation of 0.12 for organism-wide DI or 0.13–0.21 for trait-specific DI. These values are lower than estimates for other organisms, suggesting that the genetic heterogeneity in DI was low. The data provide some evidence for organism-wide DI, but simulations show that organism-wide and trait-specific variation cannot be conclusively separated. Growth rate of branch tips was determined experimentally, demonstrating significant variation among individuals. FA was not significantly correlated with growth rate or with morphological variables associated with fecundity, age, size, and health. At the same time, the signs of all the correlation coefficients were consistent with the expectation of a negative relationship between DI and fitness. The simulations indicated that the correlation between FA and the underlying DI was comparatively strong (high hypothetical repeatability), implying that the lack of significant associations between FA and fitness variables reflected a weak relationship between DI and these fitness parameters. This weak relationship may be related to the low amount of DI variation in the study population.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 92 , 277–286.     

Testing the Relationship between Human Occupancy in the Landscape and Tadpole Developmental Stress

Amphibian population declines are widespread; the main causal factors are human related and include habitat fragmentation due to agriculture, mining, fires, and urban development. Brazil is the richest country in species of amphibians, and the Brazilian regions with the greatest amphibian diversity are experiencing relatively high rates of habitat destruction, but there are presently relatively few reports of amphibian declines. It is thus important to develop research methods that will detect deterioration in population health before severe declines occur. We tested the use of measurements of fluctuating asymmetry (FA) taken on amphibian larvae to detect anthropogenic stress. We hypothesized that greater human occupancy in the landscape might result in more stressful conditions for amphibians. We conducted this study at the Espinhaço mountain range in southeastern Brazil, using as a model an endemic species (Bokermannohyla saxicola, Hylidae). We chose two tadpole denticle rows and eye-nostril distance as traits for FA measurement. We measured percent cover of human-altered habitats in the landscape around tadpole sampling points and measured FA levels in sampled tadpoles. We found FA levels to differ among localities but found no relationship between human modification of the landscape and tadpole FA levels. Levels of FA in the traits we examined may not be strongly affected by environmental conditions, or may be affected by local variables that were not captured by our landscape-scale measures. Alternatively, populations may be genetically differentiated, affecting how FA levels respond to stress and obscuring the effects of anthropogenic disturbance.     

Fluctuating asymmetry,fecundity and development time in Drosophila: is there an association under optimal and stress conditions?

A number of studies have reported a significant negative association between fluctuating asymmetry (FA) of bilateral morphological traits and individual fitness traits, but almost all of these are unreplicated and based on small sample sizes using single trait estimates of FA. We therefore tested if there was a relationship between the FA of five bilateral traits and fecundity and development time in Drosophila in a multiple replicated experimental design. Stressed treatments were included to increase the variability of traits and to test whether associations among traits were affected by changes in the environment. Significant positive relationships were found between the size of wing characters and mean fecundity for the 5‐day period and this relationship tended to be stronger in the stress treatments. No association was found between FA and mean fecundity for any of the traits measured. Similarly, a significant positive relationship was detected between wing trait size and development time but no association was detected between trait FA and development time. There were no differences between mean fecundity or development time of extreme asymmetry phenotypes compared with modal phenotypes. These results are discussed with reference to suggestions in the literature that FA can be used to estimate individual fitness.     

Effects of experimental manipulations on the life history pattern of Lymnaea stagnalis appressa say (pulmonata: lymnaeidae)

Field sampling of an Iowa population of Lymnaea stagnalis appressa Say indicated an annual generation pattern, with survivorship to maturity of i percent or less. Estimates of adult fecundity ranged from about 300 to 800 eggs.Density and food manipulations were performed to determine whether density dependent limitation of growth rates, maturation, or fecundity occurs in this fresh water pulmonate snail. Addition of a high quality food resource, spinach, accelerated growth rates, but did not drastically accelerate maturity, nor increase fecundity. Density increments lowered growth rates, delayed maturity, and lowered fecundity, and the addition of spinach did not counteract high densities. Adult densities are fairly low in the field population, and adults are randomly dispersed, indicating little density dependent regulation of fecundity in this population. However, the low survivorship to maturity, response in growth rates with food addition, and increasing survivorship with age and size indicate that juvenile mortality may play an important role in structuring life history patterns in this population.     

Ecology and ovarian function among Lese women of the Ituri Forest, Zaire

Ovarian function is examined in 35 Lese women inhabiting the Ituri Forest of northeastern Zaire over a period of 4 months through measurements of progesterone in saliva samples collected twice weekly. Ovulatory frequency is found to be only 56% on average, with a pattern of age variation similar to that observed in western women, though lower in level at each age. Average luteal progesterone levels of the Lese women are lower than those of Boston controls even if only ovulatory cycles are considered. Women with the poorest nutritional status, inferred from longitudinal weight changes and weight for height, show the greatest compromise of ovarian function, and the average ovulatory frequency of the whole sample declines in parallel with a period of weight loss over four months. It is suggested that low ovulatory frequency and luteal insufficiency contribute to the low fecundity of the Lese population and that nutritional status is likely to be one of the ecological factors modulating this effect.     

Temporal structure and some features of stock dynamics of pink salmon <Emphasis Type="Italic">Oncorhynchus gorbuscha</Emphasis> (Salmonidae)

Characteristics of early and late migrants of pink salmon Oncorhynchus gorbuscha that spawned in the rivers of the coast of Aniva Bay (southern Sakhalin) and Iturup Island (southern Kuril Islands) are described. Upstream migration of the late form is characterized by the presence of large exemplars, especially males, and increasing proportion of these fishes in the catches. Absolute fecundity of the females of the early and late forms can be similar (Iturup Island) or different (Aniva Bay), but relative fecundity is lower in the females of the late form in both regions. Production of a larger number of eggs by the females of the early form is associated with a larger level of mortality: the abundance of the early form is comparatively low, and substantial annual variation of the abundance is registered. Relative abundance of pink salmon from the temporal groups depends mainly on reproductive conditions during the freshwater part of their life cycle. Analysis of size composition of the fish conducted over many years shows a trend in the ratio between the body lengths of males and females: unfavorable foraging conditions (and slower growth rate) lead to the appearance of females that are larger than males. To take into account a positive correlation between the body length and fecundity of females, this feature is regarded as a compensatory reaction of the population directed to more intensive reproduction at poor foraging conditions.     

A meta‐analysis of latitudinal variations in life‐history traits of roach,Rutilus rutilus,over its geographical range: linear or non‐linear relationships?

1. We collated information from the literature on life history traits of the roach (a generalist freshwater fish), and analysed variation in absolute fecundity, von Bertalanffy parameters, and reproductive lifespan in relation to latitude, using both linear and non-linear regression models. We hypothesized that because most life history traits are dependent on growth rate, and growth rate is non-linearly related with temperature, it was likely that when analysed over the whole distribution range of roach, variation in key life history traits would show non-linear patterns with latitude.
2. As fecundity depends strongly on length, and the length structure of females varied among populations, latitudinal patterns in fecundity were examined based on residuals from the length–fecundity relationship. The reproductive lifespan of roach was estimated as the difference between age at maturity and maximum age of females in each population.
3. The three life history traits of roach analysed all varied among populations and were non-linearly related to latitude. Only the relationship between reproductive lifespan and latitude was a better fit to a linear that to a quadratic model, although Loess smoothing curves revealed that this relationship was actually closer to biphasic than linear in form. A latitude of 50°N formed a break point in all three life history traits.
4. The negative relationships we have described between (i) fecundity and reproductive lifespan and (ii) fecundity and egg mass suggest that lower fecundity is compensated for by longer lifespan, while lower fecundity is compensated for by an increased egg mass, when analysed independently of location.     

Determinants of fecundity and reproductive success in captive vervet monkeys

Between 1975 and 1983, adult female vervet monkeys (Cercopithecus aethiops sabaeus) over 3.5 years of age, living in two undisturbed social groups in a captive colony in Sepulveda, California, have averaged 1.0 births per female year with a mean interbirth interval of 10.7 months. Increased fecundity did not result in decreased survival rates of offspring in this population. Fecundity was influenced by the mother's age and dominance rank. The primary factor in the age-fecundity relationship was the age at first birth, which varied from three to five years. High-ranking females contributed the most to the high rate of fecundity, with significantly shorter interbirth intervals, more births per female year, and more surviving infants compared to low-ranking females.     

An integrated population model sheds light on the complex population dynamics of a unique colonial breeder

Climate models forecast increasing climatic variation and more extreme events, which could increase the variability in animal demographic rates. More variable demographic rates generally lead to lower population growth and can be detrimental to wild populations, especially if the particular demographic rates affected are those to which population growth is most sensitive. We investigated the population dynamics of a metapopulation of 25 colonies of a semi-arid bird species, the sociable weaver Philetairus socius, and how it was influenced by seasonal weather during 1993–2014. We constructed an integrated population model which estimated population sizes similar to observed population counts, and allowed us to estimate annual fecundity and recruitment. Variance in fecundity contributed most to variance in population growth, which showed no trend over time. No weather variables explained overall demographic variation at the population level. However, a separate analysis of the largest colony showed a clear decline with a high extinction probability (0.05 to 0.33) within 5 years after the study period. In this colony, juvenile survival was lower when summers were hot, and adult survival was lower when winters were cold. Rainfall was also negatively correlated with adult survival. These weather effects could be due to increased physiological demands of thermoregulation and rainfall-induced breeding activity. Our results suggest that the dynamics of the population on the whole are buffered against current weather variation, as individual colonies apparently react in different ways. However, if more and increasingly extreme weather events synchronize colony dynamics, they are likely to have negative effects.     

Identification of age-structured models: cell cycle phase transitions

A methodology is developed that determines age-specific transition rates between cell cycle phases during balanced growth by utilizing age-structured population balance equations. Age-distributed models are the simplest way to account for varied behavior of individual cells. However, this simplicity is offset by difficulties in making observations of age distributions, so age-distributed models are difficult to fit to experimental data. Herein, the proposed methodology is implemented to identify an age-structured model for human leukemia cells (Jurkat) based only on measurements of the total number density after the addition of bromodeoxyuridine partitions the total cell population into two subpopulations. Each of the subpopulations will temporarily undergo a period of unbalanced growth, which provides sufficient information to extract age-dependent transition rates, while the total cell population remains in balanced growth. The stipulation of initial balanced growth permits the derivation of age densities based on only age-dependent transition rates. In fitting the experimental data, a flexible transition rate representation, utilizing a series of cubic spline nodes, finds a bimodal G(0)/G(1) transition age probability distribution best fits the experimental data. This resolution may be unnecessary as convex combinations of more restricted transition rates derived from normalized Gaussian, lognormal, or skewed lognormal transition-age probability distributions corroborate the spline predictions, but require fewer parameters. The fit of data with a single log normal distribution is somewhat inferior suggesting the bimodal result as more likely. Regardless of the choice of basis functions, this methodology can identify age distributions, age-specific transition rates, and transition-age distributions during balanced growth conditions.     

The companion dog as a model for human aging and mortality

Around the world, human populations have experienced large increases in average lifespan over the last 150 years, and while individuals are living longer, they are spending more years of life with multiple chronic morbidities. Researchers have used numerous laboratory animal models to understand the biological and environmental factors that influence aging, morbidity, and longevity. However, the most commonly studied animal species, laboratory mice and rats, do not experience environmental conditions similar to those to which humans are exposed, nor do we often diagnose them with many of the naturally occurring pathologies seen in humans. Recently, the companion dog has been proposed as a powerful model to better understand the genetic and environmental determinants of morbidity and mortality in humans. However, it is not known to what extent the age‐related dynamics of morbidity, comorbidity, and mortality are shared between humans and dogs. Here, we present the first large‐scale comparison of human and canine patterns of age‐specific morbidity and mortality. We find that many chronic conditions that commonly occur in human populations (obesity, arthritis, hypothyroidism, and diabetes), and which are associated with comorbidities, are also associated with similarly high levels of comorbidity in companion dogs. We also find significant similarities in the effect of age on disease risk in humans and dogs, with neoplastic, congenital, and metabolic causes of death showing similar age trajectories between the two species. Overall, our study suggests that the companion dog may be an ideal translational model to study the many complex facets of human morbidity and mortality.     

Climate envelope, life history traits and the resilience of birds facing global change

Few studies have examined how life history traits and the climate envelope influence the ability of species to respond to climate change and habitat degradation. In this study, we test whether 18 species-specific variables, related to the climate envelope, ecological envelope and life history, could predict recent population trends (over 17 years) of 71 common breeding bird species in France. Habitat specialists were declining at a much higher rate than generalists, a sign that habitat quality is decreasing globally. The lower the thermal maximum (temperature at the hot edge of the climate envelope), the more negative are the population trends and the less tolerant these species are climate warming, regardless of the thermal range over which these species occur. The life history trait 'the number of broods per year' was positively related to recent trends, suggesting that single-brooded species might be more sensitive to advances in food peak due to climate change, as it increases the risk of mistiming their single-breeding event. Annual fecundity explained long-term declines, as it is a good proxy for most other demographic rates, with shorter-lived species being more sensitive to global change: individuals of species with higher fecundity might have too short a life to learn to adapt to directional changes in their environment. Finally, there was evidence that natal dispersal was a predictor of recent trends, with species with high natal dispersal experiencing smaller population declines than species with low natal dispersal. This is expected if the higher the natal dispersal, the larger the ability to shift spatially when facing changes in local habitat or climate, in order to track optimal conditions and adapt to global change. Identifying decline-promoting factors allow us to infer mechanisms responsible for observed declines in wild bird populations facing global change, and by doing so allow for a more pre-emptive approach to conservation planning.     

Lower fecundity in parthenogenetic geckos than sexual relatives in the Australian arid zone

Theoretical models of the advantage of sexual reproduction typically assume that reproductive output is equal in sexual and parthenogenetic females. We tested this assumption by comparing fecundity between parthenogenetic and sexual races of gekkonid lizards in the Heteronotia binoei complex, collected across a 1200 km latitudinal gradient through the Australian arid zone. Under laboratory conditions, parthenogenetic geckos had approximately 30% lower fecundity when compared with their sexual progenitors, irrespective of body size. Reflecting clutch-size constraints in gekkonids, this fecundity difference was mainly because of fewer clutches over a shorter period. When parthenogens were compared more broadly with all coexisting sexual races across the latitudinal gradient, parthenogens had lower fecundity than sexuals only when corrected for body size. Differences in fecundity between parthenogens and coexisting sexual races depended on which sexual race was considered. There was no significant relationship between fecundity and parasite (mite) load, despite significantly higher mite loads in parthenogens than in sexual races.     

Kin competition,natal dispersal and the moulding of senescence by natural selection

Most theoretical models for the evolution of senescence have assumed a very large, well mixed population. Here, we investigate how limited dispersal and kin competition might influence the evolution of ageing by deriving indicators of the force of selection, similar to Hamilton (Hamilton 1966 J. Theor. Biol. 12, 12–45). Our analytical model describes how the strength of selection on survival and fecundity changes with age in a patchy population, where adults are territorial and a fraction of juveniles disperse between territories. Both parent–offspring competition and sib competition then affect selection on age-specific life-history traits. Kin competition reduces the strength of selection on survival. Mutations increasing mortality in some age classes can even be favoured by selection, but only when fecundity deteriorates rapidly with age. Population structure arising from limited dispersal however selects for a broader distribution of reproduction over the lifetime, potentially slowing down reproductive senescence. The antagonistic effects of limited dispersal on age schedules of fecundity and mortality cast doubts on the generality of conditions allowing the evolution of ‘suicide genes’ that increase mortality rates without other direct pleiotropic effects. More generally, our model illustrates how limited dispersal and social interactions can indirectly produce patterns of antagonistic pleiotropy affecting vital rates at different ages.     

Patterns of reproduction in two populations of pumpkinseed sunfish,Lepomis gibbosus,with differing food resources

Synopsis The influence of available food resources on growth and patterns of reproduction was studied in two populations of pumpkinseed sunfish in adjacent lakes in southeastern Ontario. Ideally, mature individuals should be long-lived, attain a large size, and maintain a high reproductive effort. However, rarely are environmental conditions ideal and compromises are necessary. In the present study, there were smaller quantities of preferred prey items (gastropods), and total benthos available in one lake. Pumpkinseed in this population ate less in general, and were smaller in body size in the older age classes. These fish matured at a younger age and a smaller size than individuals in the other lake studied, and in populations in this geographic area reported in the literature. In addition, the relationships of both fecundity and ovary weight to total weight were different between lakes. Mean ovary weight, ripe egg size and lifetime fecundity were less in the population of small-bodies fish. Variation in patterns of reproduction between populations suggests that fish may be able to fine-tune responses to local environmental conditions.     

Population dynamics of epiphytic orchids in a metapopulation context

Background and Aims

Populations of many epiphytes show a patchy distribution where clusters of plants growing on individual trees are spatially separated and may thus function as metapopulations. Seed dispersal is necessary to (re)colonize unoccupied habitats, and to transfer seeds from high- to low-competition patches. Increasing dispersal distances, however, reduces local fecundity and the probability that seeds will find a safe site outside the original patch. Thus, there is a conflict between seed survival and colonization.

Methods

Populations of three epiphytic orchids were monitored over three years in a Mexican humid montane forest and analysed with spatially averaged and with spatially explicit matrix metapopulation models. In the latter, population dynamics at the scale of the subpopulations (epiphytes on individual host trees) are based on detailed stage-structured observations of transition probabilities and trees are connected by a dispersal function.

Key Results

Population growth rates differed among trees and years. While ignoring these differences, and averaging the population matrices over trees, yields negative population growth, metapopulation models predict stable or growing populations because the trees that support growing subpopulations determine the growth of the metapopulation. Stochastic models which account for the differences among years differed only marginally from deterministic models. Population growth rates were significantly lower, and extinctions of local patches more frequent in models where higher dispersal results in reduced local fecundity compared with hypothetical models where this is not the case. The difference between the two models increased with increasing mean dispersal distance. Though recolonization events increased with dispersal distance, this could not compensate the losses due to reduced local fecundity.

Conclusions

For epiphytes, metapopulation models are useful to capture processes beyond the level of the single host tree, but local processes are equally important to understand epiphyte population dynamics.