Estimating density dependence from population time series using demographic theory and life-history data

For populations with a density-dependent life history reproducing at discrete annual intervals, we analyze small or moderate fluctuations in population size around a stable equilibrium, which is applicable to many vertebrate populations. Using a life history having age at maturity alpha, with stochasticity and density dependence in adult recruitment and mortality, we derive a linearized autoregressive equation with time lags from 1 to alpha yr. Contrary to current interpretations, the coefficients corresponding to different time lags in the autoregressive dynamics are not simply measures of delayed density dependence but also depend on life-history parameters. The theory indicates that the total density dependence in a life history, D, should be defined as the negative elasticity of population growth rate per generation with respect to change in population size, [Formula: see text], where lambda is the asymptotic multiplicative growth rate per year, T is the generation time, and N is adult population size. The total density dependence in the life history, D, can be estimated from the sum of the autoregression coefficients. We estimate D in populations of seven vertebrate species for which life-history studies and unusually long time series of complete population censuses are available. Estimates of D were statistically significant and large, on the order of 1 or higher, indicating strong density dependence in five of the seven species. We also show that life history can explain the qualitative features of population autocorrelation functions and power spectra and observations of increasing empirical variance in population size with increasing length of time series.     

Expression and regulation of delta5-desaturase,delta6-desaturase,stearoyl-coenzyme A (CoA) desaturase 1, and stearoyl-CoA desaturase 2 in rat testis

In mammalian cells, essential polyunsaturated fatty acids (PUFAs) are converted to longer PUFAs by alternating steps of elongation and desaturation. In contrast to other PUFA-rich tissues, the testis is continuously drained of these fatty acids as spermatozoa are transported to the epididymis. Alteration of the germ cell lipid profile from spermatogonia to condensing spermatids and mature spermatozoa has been described, but the male gonadal gene expression of the desaturases, responsible for the PUFA-metabolism, is still not established. The focus of this study was to characterize the expression and regulation of stearoyl-CoA desaturase 1 (SCD1), stearoyl-CoA desaturase 2 (SCD2), and Delta5- and Delta6-desaturase in rat testis. Desaturase gene expression was detected in testis, epididymis, and separated cells from seminiferous tubulus using Northern blot analysis. For the first time, SCD1 and SCD2 expression is demonstrated in rat testis and epididymis, both SCDs are expressed in epididymis, while testis mainly contains SCD2. Examination of the testicular distribution of Delta5- and Delta6-desaturase and SCD1 and SCD2 shows that all four desaturases seem to be localized in the Sertoli cells, with far lower expression in germ cells. In light of earlier published results showing that germ cells are richer in PUFAs than Sertoli cells, this strengthens the hypothesis of a lipid transport from the Sertoli cells to the germ cells. As opposed to what is shown in liver, Delta5- and Delta6-desaturase mRNA levels in Sertoli cells are up-regulated by dexamethasone. Furthermore, dexamethasone induces SCD2 mRNA. Insulin also up-regulates these three genes in the Sertoli cell, while SCD1 mRNA is down-regulated by both insulin and dexamethasone. Delta5- and Delta6-desaturase, SCD1, and SCD2 are all up-regulated by FSH. A similar up-regulation of the desaturases is observed when treating Sertoli cells with (Bu)2cAMP, indicating that the desaturase up-regulation observed with FSH treatment results from elevated levels of cAMP. Finally, testosterone has no influence on the desaturase gene expression. Thus, FSH seems to be a key regulator of the desaturase expression in the Sertoli cell.     

Regulation of foraging trips and costs of incubation shifts in the Antarctic petrel (Thalassoica antarctica)

In species where incubation is shared by both parents, the mate'sability to fast on the nest may constrain the time availablefor foraging. The decision to return to the nest should thereforebe a compromise between an animal's own foraging success andits mate's ability to fast on the nest. To examine how the bodyconditions of incubating Antarctic petrels, Thalassoica antarctica,influence both the length of foraging trips and incubation shifts,we experimentally handicapped females by increasing their flightcosts during a foraging trip by adding lead weights to theirlegs. Handicapped females spent more time at sea and had lowerbody conditions at arrival to the colony than controls, and,moreover, females in poor body condition at arrival to the colonyspent generally more time at sea than those with higher bodycondition. The prolonged time period spent at sea by handicappedfemales was associated with higher desertion rates than amongcontrols. The time the incubating mates fasted increased withtheir body condition at arrival to the colony, suggesting thata high body condition of the incubating bird may reduce theprobability of nest desertion. Accordingly, our results suggestthat the time spent foraging is adjusted to the body conditionsof both the foraging and incubating mate.     

Finite metapopulation models with density-dependent migration and stochastic local dynamics

The effects of small density-dependent migration on the dynamics of a metapopulation are studied in a model with stochastic local dynamics. We use a diffusion approximation to study how changes in the migration rate and habitat occupancy affect the rates of local colonization and extinction. If the emigration rate increases or if the immigration rate decreases with local population size, a positive expected rate of change in habitat occupancy is found for a greater range of habitat occupancies than when the migration is density-independent. In contrast, the reverse patterns of density dependence in respective emigration and immigration reduce the range of habitat occupancies where the metapopulation will be viable. This occurs because density-dependent migration strongly influences both the establishment and rescue effects in the local dynamics of metapopulations.     

Effective size of fluctuating populations with two sexes and overlapping generations

We derive formulas that can be applied to estimate the effective population size N(e) for organisms with two sexes reproducing once a year and having constant adult mean vital rates independent of age. Temporal fluctuations in population size are generated by demographic and environmental stochasticity. For populations with even sex ratio at birth, no deterministic population growth and identical mean vital rates for both sexes, the key parameter determining N(e) is simply the mean value of the demographic variance for males and females considered separately. In this case Crow and Kimura's generalization of Wright's formula for N(e) with two sexes, in terms of the effective population sizes for each sex, is applicable even for fluctuating populations with different stochasticity in vital rates for males and females. If the mean vital rates are different for the sexes then a simple linear combination of the demographic variances determines N(e), further extending Wright's formula. For long-lived species an expression is derived for N(e) involving the generation times for both sexes. In the general case with nonzero population growth and uneven sex ratio of newborns, we use the model to investigate numerically the effects of different population parameters on N(e). We also estimate the ratio of effective to actual population size in six populations of house sparrows on islands off the coast of northern Norway. This ratio showed large interisland variation because of demographic differences among the populations. Finally, we calculate how N(e) in a growing house sparrow population will change over time.     

Multilocus heterozygosity and inbreeding depression in an insular house sparrow metapopulation

Inbreeding causes reduction of genetic variability that may have severe fitness consequences. In spite of its potentially huge impact on viability and evolutionary processes especially in small populations, quantitative demonstrations of genetic and demographic effects of inbreeding in natural populations are few. Here, we examine the relationship between individual inbreeding coefficients (F) and individual standardized multilocus heterozygosity (H) in an insular metapopulation of house sparrows (Passer domesticus) in northern Norway in order to evaluate whether H is a good predictor for F. We then relate variation in fitness (i.e. the probability of surviving from fledging to recruitment) to F and H, which enables us to examine whether inbreeding depression is associated with a reduction in genetic variability. The average level of inbreeding in the house sparrow metapopulation was high, and there was large inter-individual variation in F. As expected, standardized multilocus heterozygosity decreased with the level of inbreeding. The probability of recruitment was significantly negatively related to F, and, accordingly, increased with H. However, H explained no significant additional variation in recruitment rate than was explained by F. This suggests that H is a good predictor for F in this metapopulation, and that an increase in F is likely to be associated with a general increase in the level of homozygosity on loci across the genome, which has severe fitness consequences.     

Predicting the time to quasi-extinction for populations far below their carrying capacity

Populations threatened by extinction are often far below their carrying capacity. A population collapse or quasi-extinction is defined to occur when the population size reaches some given lower density. If this density is chosen to be large enough for the demographic stochasticity to be ignored compared to environmental stochasticity, then the logarithm of the population size may be modelled by a Brownian motion until quasi-extinction occurs. The normal-gamma mixture of inverse Gaussian distributions can then be applied to define prediction intervals for the time to quasi-extinction in such processes. A similar mixture is used to predict the population size at a finite time for the same process provided that quasi-extinction has not occurred before that time. Stochastic simulations indicate that the coverage of the prediction interval is very close to the probability calculated theoretically. As an illustration, the method is applied to predict the time to extinction of a declining population of white stork in southwestern Germany.     

Seasonal cycles of species diversity and similarity in a tropical butterfly community

1. Studies of seasonality in ecological diversity rarely extend over more than a few years, and few studies of seasonal diversity have explicitly investigated the influence of environmental factors on seasonal community composition, especially in tropical communities. 2. Our 10 years of monthly sampling in Amazonian Ecuador yielded 20 996 individuals of 137 fruit-feeding butterfly species. Seasonal cycles of rainfall drive annual cycles in species diversity and community similarity. Undetermined processes operating most strongly during the dry season maintain species diversity and high community similarity across years. 3. Seasonal cycles in community diversity and similarity are superimposed on a gradual decline in similarity between community samples on a decadal time-scale because of long-term changes in species abundances. 4. Monitoring and analysis of changes in community composition over a range of time-scales can be used to refine models of community dynamics by incorporating environmental factors necessary to predict the ecological impact of future climate change.     

Hanocladius,a new orthoclad genus from China (Diptera: Chironomidae)

Hanocladius longipes gen. nov., sp. nov. from Oriental China is described from the male imago. The long metatarsus, the very broad, scale-like virga and the conspicuously scalpellate acrostichals distinguish the genus from all of the known genera in Orthocladiinae.