Interactions between density-dependent processes, population dynamics and control of an invasive plant species, Tripleurospermum perforatum (scentless chamomile)

Tripleurospermum perforatum is an invasive weedy species which exhibits strong over-compensating density dependence. Interactions between density-dependent survival, probability of flowering and fecundity were modelled and their impact on the population dynamics were examined. When only fecundity was density-dependent, the dynamics were similar to those observed in the model containing all three density-dependent terms. Density-dependent survival was a stabilizing process when acting in combination with density-dependent fecundity and probability of flowering; removing density-dependent survival from the model produced two-point cycles. The addition of a seed bank was also stabilizing. Simulations of control strategies at different life-history stages indicated that full control would be difficult due to the strong over-compensating density dependence, with severe reductions in fecundity and late season survival necessary in order to reduce equilibrium seed density and biomass.     

Density-dependent parameters and demographic equilibrium in open populations

Populations near their equilibrium are expected to show density-dependence through a negative feedback on at least one demographic parameter, e.g. survival and/or productivity. Nevertheless, it is not always clear which vital rate is affected the most, and even less whether this dependence holds in open populations in which immigration and emigration are also important. We assessed the relative importance of population density in the variation of local survival, recruitment, proportion of transients (emigrants) and productivity through the analysis of detailed life-histories of 4286  seabirds from a colony that reached an apparent demographic equilibrium after a period of exponential increase. We provide evidence that the role of population density and resource availability changes according to the demographic parameter considered. Estimates indicated that transients increased from 5% to 20% over the study period, suggesting an average turnover of about 1400 individuals per year. The parameters most influenced by population density alone were local survival and probability of transience. Recruitment was negatively associated with population density during the increasing phase but unexpected high values were also recorded at high population levels. These high values were explained by a combination of population size and food availability. Mean productivity varied with food availability, independently from population variations. The population density alone explained up to a third of the yearly variation of the vital rates considered, suggesting that open populations are equally influenced by stochastic and density-independent events (such as environmental perturbations) than by intrinsic (i.e. density-dependent) factors.     

Exploring density-dependent relationships in demographic parameters in populations of birds at a large spatial scale

The importance of density-dependent processes in natural populations is widely accepted, but the issue of the shape of density-dependent relationships (such as influenced by vagueness, or time-delay) remains unresolved. We explored the density-dependent relationships in demographic parameters for 12 species of birds in Britain using large-scale, long-term data sets. We predicted that a negative relation between density and demographic parameters should be observed for the stable species, whereas the decreasing or increasing species should display a positive relation if the environment changes progressively through time bringing about a continuous change in density dependence. Our prediction was verified for nine species out of 12; however, we observed, for the three remaining species, a significant decrease of survival rates through time that seems to be involved in a long-term population decline. In all cases where a density-dependent relation was found, we observed an important variance around the relation. In one case, we showed that this variance increased significantly with density. We found evidence for time-delayed effects of density dependence both for survival and breeding performance. In two species, our results suggest the existence of complex interactions (compensatory mechanisms) between survival and breeding performance or between the different components of breeding performance.     

Density-Dependent Growth in Invasive Lionfish (Pterois volitans)

Direct demographic density dependence is necessary for population regulation and is a central concept in ecology, yet has not been studied in many invasive species, including any invasive marine fish. The red lionfish (Pterois volitans) is an invasive predatory marine fish that is undergoing exponential population growth throughout the tropical western Atlantic. Invasive lionfish threaten coral-reef ecosystems, but there is currently no evidence of any natural population control. Therefore, a manipulative field experiment was conducted to test for density dependence in lionfish. Juvenile lionfish densities were adjusted on small reefs and several demographic rates (growth, recruitment, immigration, and loss) were measured throughout an 8-week period. Invasive lionfish exhibited direct density dependence in individual growth rates, as lionfish grew slower at higher densities throughout the study. Individual growth in length declined linearly with increasing lionfish density, while growth in mass declined exponentially with increasing density. There was no evidence, however, for density dependence in recruitment, immigration, or loss (mortality plus emigration) of invasive lionfish. The observed density-dependent growth rates may have implications for which native species are susceptible to lionfish predation, as the size and type of prey that lionfish consume is directly related to their body size. The absence of density-dependent loss, however, contrasts with many native coral-reef fish species and suggests that for the foreseeable future manual removals may be the only effective local control of this invasion.     

Density-dependent reproduction in the European rabbit: a consequence of individual response and age-dependent reproductive performance

Density dependence of reproduction has generally been proposed to be caused by habitat heterogeneity and by the individual response of reproductive output. However, a further mechanism might generate density dependence of average reproductive rates. High density situations might be associated with a high proportion of first-season breeders which often show a principally lower reproductive performance. We tested for the existence of the latter mechanism as well as for density-dependent individual changes of reproductive effort in a population of European rabbits living in a homogeneous grassland habitat. The study was conducted over a period of eleven years. Overall, a strong relationship between mean reproductive rates and the breeding density of females was apparent. All necessary conditions for the presence of a density-dependent effect caused by age-dependent reproduction were fulfilled: Fluctuations of breeding density were paralleled by variations in the proportion of one-year-old females. These one-year-old, first-season breeders showed a consistently lower reproductive performance than older females, which might be caused by their lower body mass and their lower social rank. However, we also found strong evidence for density-dependent response of individual reproductive effort: Individual changes in fecundity over successive years were explained by changes in the breeding density of females. The results suggest that density dependence of reproduction in European rabbits is due to an interaction of age-dependent reproductive performance together with short-term fluctuations in breeding density, and a density-dependent, individual based response of reproductive rates. We further conclude that the lower reproductive performance of first-season breeders in age-structured animal populations may contribute substantially to interannual, and under particular circumstances to density-dependent variations of mean reproductive rates.     

The relative roles of density and climatic variation on population dynamics and fecundity rates in three contrasting ungulate species

The relative influences of density-dependent and -independent processes on vital rates and population dynamics have been debated in ecology for over half a century, yet it is only recently that both processes have been shown to operate within the same population. However, generalizations on the role of each process across species are rare. Using a process-orientated generalized linear modelling approach we show that variations in fecundity rates in populations of three species of ungulates with contrasting life histories are associated with density and winter weather in a remarkably similar manner. However, there are differences and we speculate that they are a result of differences in size between the species. Much previous research exploring the association between vital rates, population dynamics and density-dependent and -independent processes has used pattern-orientated approaches to decompose time-series into contributions from density-dependent and -independent processes. Results from these analyses are sometimes used to infer associations between vital rates, density and climatic variables. We compare results from pattern-orientated analyses of time-series with process-orientated analyses and report that the two approaches give different results. The approach of analysing relationships between vital rates, density and climatic variables may detect important processes influencing population dynamics that time-series methodologies may overlook.     

Geographical variation in the influence of density dependence and climate on the recruitment of Norwegian moose

The effects of variation in climate on population dynamics are likely to differ within the distributional range of a species, yet the consequences of such regional variation on demography and population dynamics are rarely considered. Here we examine how density dependence and different climate variables affect spatio-temporal variation in recruitment rates of Norwegian moose using data collected over a large geographical area during the hunting season. After accounting for observation error by a Bayesian Markov chain Monte Carlo technique, temporal variation in recruitment rates was relatively independent of fluctuations in local population size. In fact, a positive relationship was as common as a density-dependent decrease in fecundity rates. In general, high recruitment rates were found during autumn 1 year after years with a warm February, and after a warm May or cold June in year t − 1 or in year t. Large regional variation was also found in the effects of some of the weather variables, especially during spring. These patterns demonstrate both direct and delayed effects of weather on the recruitment of moose that possibly operate through an effect of body mass on the proportion of the females that sexually mature as 1.5 or 2.5 years old.     

Recruitment limitation: A theoretical perspective

Abstract A theoretical analysis of the concept of recruitment limitation leads to the conclusion that most populations should he regarded as jointly limited by recruitment and interactions between individuals after recruitment. The open nature of local marine systems does not permit avoidance of density-dependent interactions; it simply may make them more difficult to detect. Local populations consisting of settled organisms may not experience density-dependent interactions under some circumstances, but the entire species population consisting of the collection of local populations and their planktonic larvae must have density-dependent dynamics. Any local population of settled individuals can escape density dependence if sufficient density dependence occurs among planktonic larvae or within other local populations. Common conceptions of density dependence are too narrow, leading too often to the conclusion that it is absent from a system. It is equally wrong to expect that density-dependent interactions after settlement determine local population densities independently of recruitment. Special circumstances allowing density dependence to act strongly and quickly are needed before density dependence can neutralize the effects of recruitment. Recruitment limitation and density-dependent interactions therefore should not be regarded as alternatives but as jointly acting to determine the densities of marine benthic populations. Moreover, the interaction between fluctuating recruitment and density dependence is potentially the most interesting feature of recruitment limitation. For example, this interaction may be an important diversity-maintaining mechanism for marine systems.     

Density-dependent effects on physical condition and reproduction in North American elk: an experimental test

Density dependence plays a key role in life-history characteristics and population ecology of large, herbivorous mammals. We designed a manipulative experiment to test hypotheses relating effects of density-dependent mechanisms on physical condition and fecundity of North American elk (Cervus elaphus) by creating populations at low and high density. We hypothesized that if density-dependent effects were manifested principally through intraspecific competition, body condition and fecundity of females would be lower in an area of high population density than in a low-density area. Thus, we collected data on physical condition and rates of pregnancy in each experimental population. Our manipulative experiment indicated that density-dependent feedbacks affected physical condition and reproduction of adult female elk. Age-specific pregnancy rates were lower in the high-density area, although there were no differences in pregnancy of yearlings or in age at peak reproduction between areas. Age-specific rates of pregnancy began to diverge at 2 years of age between the two populations and peaked at 6 years old. Pregnancy rates were most affected by body condition and mass, although successful reproduction the previous year also reduced pregnancy rates during the current year. Our results indicated that while holding effects of winter constant, density-dependent mechanisms had a much greater effect on physical condition and fecundity than density-independent factors (e.g., precipitation and temperature). Moreover, our results demonstrated effects of differing nutrition resulting from population density during summer on body condition and reproduction. Thus, summer is a critical period for accumulation of body stores to buffer animals against winter; more emphasis should be placed on the role of spring and summer nutrition on population regulation in large, northern herbivores.     

Contributions of forage competition, harvest, and climate fluctuation to changes in population growth of northern white-tailed deer

Recently there has been considerable interest in determining the relative roles of endogenous (density-dependent) and exogenous (density-independent) factors in driving the population dynamics of free-ranging ungulates. We used time-series analysis to estimate the relative contributions of density-dependent forage competition, climatic fluctuation, and harvesting on the population dynamics of white-tailed deer (Odocoileus virginianus) in Nova Scotia, Canada, from 1983 to 2000. A model incorporating the population density 2 years previous, an interaction term for the harvest of females and population density 2 years previous, and the total snowfall during the previous 2 winters explained 80% of the variation in inter-annual population growth rate. Natality of adult females was negatively related to deer density during the present winter, whereas that of yearlings may have been correlated with the snowfall of three winters previous. Natality of fawns was related to deer density and total snowfall during the previous winter. Coyotes (Canis latrans) prey extensively on deer fawns in northeastern North America and the annual harvest of snowshoe hares (Lepus americanus), the major alternate prey of coyotes, explained 48% of the inter-annual variation in fawn recruitment. The proportions of fawn, yearling, and adult deer suffering from severe malnutrition during late winter were all correlated with deer density during the present winter. We conclude that the limiting effects of winter weather on over-winter survival of deer may be cumulative over two consecutive winters. During the late 1980s, density dependence and winter severity acted in concert to effect substantial declines in deer population growth both by effecting winter losses directly and by exacerbating predation by coyotes. During this period liberal harvesting did not relieve density-dependent forage competition and probably accelerated the decline.     

Survival-variation within and between functional categories of the African multimammate rat

1. By identifying ecological factors specific to functional categories of individuals, it may be possible to understand the mechanisms underlying life-history evolution and population dynamics. While empirical analyses within the field of population biology have focused on changes in population size, theoretical models assuming differential sensitivities of population growth rate or fitness to demographic parameters have mostly been untested, particularly against data on small mammals.
2. Statistical modelling of capture–mark–recapture data on the multimammate rat ( Mastomys natalensis ) from Tanzania shows that: (i) females survive slightly better than males and subadults survive much better than adults; (ii) temporal variation of survival of all individuals is similarly related to the rainfall of the month; (iii) subadults exhibit a strongly density-dependent low persistence rate in the population immediately after their first capture; (iv) subadults survival in later months is, however, positively related to density; and (v) adult survival shows negative density-dependence.
3. Both density-dependent and density-independent factors simultaneously determine stage-dependent survival variation of the multimammate rat. Whereas environmental factors in this population seem to affect survival rates of all individuals in a similar manner, density-dependent relationships are more complex.
4. The patterns of survival variation in small mammals may be different from those observed in large mammals.
5. Further studies of demography in small mammals should aim at understanding how much of the variability in population growth rate is accounted for by the variability of the demographic rates resulting from limiting (density-independent) and regulating (density-dependent) factors, respectively. This study emphasizes the use of robust and accurate statistical methods as well as stage- or age-structured population modelling.     

Lottery coexistence models extended to plants with disjoint generations

Abstract. Neither conventional niche theory nor current lottery models offer a satisfactory theoretical scope for modelling coexistence of species with disjoint generations. South-African fynbos and Australian kwongan include many species which are killed by, and recruit only after, fire. We propose a density-dependent lottery model which accommodates the unusual demographics of these species. We show that coexistence requires density dependence in recruitment. The result applies to a wider class of populations than the one considered here. It is applied to non-resprouting species in fynbos and kwongan. We show that the lottery assumption of recruitment in proportion to propagules is often satisfied, while the production of such propagules is often density-dependent, and we discuss some evidence of mechanisms whereby this may occur.     

Growth and decline of a penguin colony and the influence on nesting density and reproductive success

Colonial breeding is characteristic of seabirds but nesting at high density has both advantages and disadvantages and may reduce survival and fecundity. African penguins (Spheniscus demersus) initiated breeding at Robben Island, South Africa in 1983. The breeding population on the island increased in the late 1990s and early 2000s before decreasing rapidly until 2010. Before the number breeding peaked, local nest density in the areas where the colony was initiated plateaued, suggesting that preferred nests sites were mostly occupied, and the area used by breeding birds expanded. However, it did not contract again as the population decreased, so that nesting density varied substantially. Breeding success was related positively to the prey available to the breeding birds and negatively to local nest density, particularly during the chick-rearing period, suggesting a density-dependence operating through social interactions in the colony, possibly exacerbated by poor prey availability when the breeding population was large. Although nest density at Robben Island was not high, nesting burrows, which probably reduce the incidence of aggressive encounters in the colony, are scarce and our results suggest that habitat alteration has modified the strength of density-dependent relationships for African penguins. Gaining a better understanding of how density dependence affects fecundity and population growth rates in colonial breeders is important for informing conservation management of the African penguin and other threatened taxa.     

Climate warming and the evolution of morphotypes in a reptile

Climate warming is known to have effects on population dynamics through variations in survival, fecundity and density. However, the impacts of climate change on population composition are still poorly documented. Morphotypes are powerful markers to track changes in population composition. In the common lizard, Lacerta vivipara , individuals display two types of dorsal patterns: reticulated (R individuals) and linear (L individuals). We examined how local warming affected intrapopulation frequencies of these morphotypes across 11 years. We observed changes in morph frequency of dorsal patterns across years, paralleling the rise of spring temperatures. The proportion of R individuals increased with June temperatures in juveniles, yearlings, and adult males and females. Three mechanisms could explain these changes: phenotypic plasticity, microevolution and/or dispersal between populations. We investigated the ontogenetic determinism, fitness and recruitment rates associated with dorsal morphotypes. Dorsal pattern ontogeny showed temperature dependence but this relationship was not associated with the warming trend during this study. We found variation by morphotype in survival and clutch size, but these factors did not explain R frequency increases. Among all the parameters considered in this study, only a decrease of immigration, which was more pronounced in the L morphotype, could explain the change in population composition. To our knowledge, this provides the first evidence of the impact of climate warming on population composition due to its effects on immigration.     

Density dependence and the control of helminth parasites

1. The transient dynamics and stability of a population are determined by the interplay between species density, its spatial distribution and the positive and negative density-dependent processes regulating population growth. 2. Using the human-helminth parasite system as an example, we propose that the life-stage upon which negative density dependence operates will influence the rate of host reinfection following anthelmintic chemotherapy, and the likely success of control programmes. 3. Simple deterministic models are developed which highlight how a parasite species whose population size is down-regulated by density-dependent establishment will reinfect a host population at a faster rate than a species with density-dependent parasite fecundity. 4. Different forms of density dependence can produce the same equilibrium behaviour but different transient dynamics. Under-representing the nature and magnitude of density-dependent mechanisms, and in particular those operating upon establishing life-stages, may cause the resilience of the parasite population to a control perturbation to be underestimated.     

Population regulation and demography in a harvested freshwater crayfish from Madagascar

Despite advances in statistical techniques for investigating population dynamics based on mark–recapture data, the majority of our understanding about demography and regulation comes from relatively few taxa. Most proposed generalisations about the association between demography and variation in population size are based on data from vertebrates, there are few sufficiently detailed invertebrate studies to examine whether these generalisations are widely supported. The population biology of freshwater invertebrates is especially poorly known. We present a large-scale mark–recapture study of an endemic freshwater crayfish from Madagascar ( Astacoides granulimanus ). Variation in density, caused by difference in fishing pressure due to local taboos, allowed us to investigate density-dependent regulation. We found evidence of density dependence in fecundity operating through the proportion of reproductive females by size but no significant evidence of density dependence in growth. Using a prospective analysis based on the elasticities from a size-structured matrix model, we found that both recruitment rates and survival rates of large individuals were strongly associated with deterministic population growth – a result that differs from generalisations drawn from vertebrate studies. A central assumption in mark–recapture studies is that handling does not affect mortality. By treating the number of times an individual was captured as an individual covariate, easily done using the freeware program MARK, we were able to test for, and take account of, handling-induced mortality. Our results show interesting similarities, and important differences, to generalisations based on vertebrate studies and emphasise the importance of population studies on poorly known taxa.     

Patterns and causes of demographic variation in a harvested moose population: evidence for the effects of climate and density-dependent drivers

1.?Better understanding of the mechanisms affecting demographic variation in ungulate populations is needed to support sustainable management of harvested populations. While studies of moose Alces alces L. populations have previously explored temporal variation in demographic processes, managers responsible for populations that span large heterogeneous landscapes would benefit from an understanding of how demography varies across biogeographical gradients in climate and other population drivers. Evidence of thresholds in population response to manageable and un-manageable drivers could aid resource managers in identifying limits to the magnitude of sustainable change. 2.?Generalized additive models (GAMs) were used to evaluate the relative importance of population density, habitat abundance, summer and winter climatic conditions, primary production, and harvest intensity in explaining spatial variation in moose vital rates in Ontario, Canada. Tree regression was used to test for thresholds in the magnitudes of environmental predictor variables that significantly affected population vital rates. 3.?Moose population growth rate was negatively related to moose density and positively related to the abundance of mixed deciduous habitat abundant in forage. Calf recruitment was negatively related to a later start of the growing season and calf harvest. The ratio of bulls to cows was related to male harvest and hunter access, and thresholds were evident in predictor variables for all vital rate models. 4.?Findings indicate that the contributions of density-dependent and independent factors can vary depending on the scale of population process. The importance of density dependence and habitat supply to low-density ungulate populations was evident, and management strategies for ungulates may be improved by explicitly linking forest management and harvest. Findings emphasize the importance of considering summer climatic influences to ungulate populations, as recruitment in moose was more sensitive to the timing of vegetation green-up than winter severity. The efficacy of management decisions for harvested ungulates may require regional shifts in targets where populations span bioclimatic gradients. The use of GAMs in combination with recursive partitioning was demonstrated to be an informative analytical framework that captured nonlinear relationships common in natural processes and thresholds that are relevant to population management in diverse systems.     

Plant performance in central and northern peripheral populations of the widespread Plantago coronopus

Peripheral populations have long been predicted to show lower vital rates, higher demographic fluctuations, and lower densities than central populations. However, recent research has questioned the existence of clear patterns across species’ ranges. To test these hypotheses, we monitored five central and six northern peripheral populations of the widespread herb Plantago coronopus along the European Atlantic coast during 5 yr. We estimated population density, and calculated mean values and temporal variability of four vital rates (survival, individual growth, fecundity and recruitment) in hundreds of plants in permanent plots. Central populations showed higher fecundity, whereas peripheral populations had higher recruitment per reproductive plant, indicating a higher overall reproductive success in the periphery. Central populations showed a marginally significant tendency for higher growth, and there were no differences between range positions in survival. Fecundity and growth were affected by intraspecific competition, and recruitment was affected by precipitation, highlighting the importance of local environmental conditions for population performance. Central and peripheral populations showed no significant differences in temporal variability of vital rates. Finally, density was significantly higher in peripheral than in central populations, in discrepancy with the abundant‐centre model. Density was correlated to seedling recruitment, which would counterbalance in peripheral populations the lower fecundity and the tendency for lower growth of established plants. Such compensations among vital rates might be particularly common in widespread plants, and advise against simplistic assumptions of population performance across ranges. The whole species’ life cycle should be considered, since different arrangements of vital rates are expected to maximize fitness in local environments. Our results show also the importance of discerning between geographical periphery and ecological marginality. In a context of climate‐induced range shifts, these considerations are crucial for the reliability of niche‐models and the management of plant peripheral populations.     

Fluctuations in numbers in box populations of Drosophila and selective genetic mechanism of their regulation

Box populations of Drosophila melanogaster are characterized by two types of periodical fluctuations of numbers: with low and high frequency. High frequency fluctuations are determined by existence of preimago and imago stages and subsequent delay in density-dependent limitation of imago reproduction, duration of which is determined by time of preimago stage. The period of these fluctuations should be limited within two generation, that is confirmed by experimental data. Low frequency fluctuations with the period of 13-15 generations are the result of ecological density-dependent effect. In this case during pick density one can observe continuous degradation of population (i.e. decrease in fecundity and life time of imago) and following decrease in numbers. Temporary changes in fecundity of females and their offspring of the second generation are positively correlated with low frequency fluctuations in numbers. Such relationships show the possibility of density-dependent, cyclic, genetic changes in fecundity connected with fluctuations in numbers. It means that at the phase of growth in numbers when the density is still low, the selection is directed to the individuals with high fecundity sensible to overpopulation. The phase of decline in numbers is connected with high density and selection directed to the individuals with low fecundity in low density populations. The changes in genetic structure of fluctuating population lead to the weakening of this fluctuations and to the maintaining of population under such conditions.     

Capitulum density-dependent effects generate peak seed yield at an intermediate density of a Tibetan lotus

Aims Theory suggests that species perform best at intermediate densities, where density-dependent facilitation and antagonism are balanced, but empirical evidence is scarce, particularly in plants. In a self-incompatible perennial herb (Saussurea nigrescens), whose recruitment heavily relies on seed output, we test whether both intraspecific facilitation and antagonism significantly affect seed production, resulting in highest seed yield at an intermediate capitulum density.Methods Plots with different S. nigrescens densities were sampled in an Eastern Tibetan meadow during the growing season of 2012 to investigate the relationships between capitulum density and pollinator visitation rate, seed set ratio, parasite ratio, seed damage ratio, and capitulum size. Both simple linear and quadratic models were employed to determine the shape of relationships.Important findings In line with general theory, hump-shaped relationships of capitulum density versus seed set ratio and number of florets per capitulum indicate intraspecific facilitation in sparse populations, which can be attributed to positive density-dependent pollinator visitation and the amelioration of detrimental physical factors. However, the proportion of seeds damaged by pre-dispersal predators increased monotonically with capitulum density, which may have—in combination with increased intraspecific competition for light and soil nutrients—resulted in density-dependent antagonism. Both positive and negative density-dependent agents acted simultaneously throughout the density range investigated and led to the highest seed yield at intermediate density levels in the Tibetan lotus. More efforts concurrently exploring the two effects are needed to facilitate understanding species abundance and community structure.