Density, climate and varying return points: an analysis of long-term population fluctuations in the threatened European tree frog

Experimental research has identified many putative agents of amphibian decline, yet the population-level consequences of these agents remain unknown, owing to lack of information on compensatory density dependence in natural populations. Here, we investigate the relative importance of intrinsic (density-dependent) and extrinsic (climatic) factors impacting the dynamics of a tree frog (Hyla arborea) population over 22 years. A combination of log-linear density dependence and rainfall (with a 2-year time lag corresponding to development time) explain 75% of the variance in the rate of increase. Such fluctuations around a variable return point might be responsible for the seemingly erratic demography and disequilibrium dynamics of many amphibian populations.     

Temporal and spatial variation of larval parasitism in non-outbreaking populations of a folivorous moth

In order to assess the role of parasitoids in the regulation of non-outbreaking populations of Epirrita autumnata, a geometrid lepidopteran with outbreaking populations in northern Europe, we examined the temporal and spatial variation of larval parasitism in southwestern Finland during 6 successive years. The study was carried out on two spatial scales, among trees within sites of about 1 ha and among sites separated by distances of 2–10 km, using experimental and observational approaches respectively. The overall percent parasitism was independent of host density on both spatial scales, while temporally it fluctuated only little. Of the two main parasitoids, the commoner one, Protapanteles immunis, showed a variable response to host density on the larger spatial scale and negative density dependence on the smaller scale. Temporally, parasitism caused by this species was independent of host density. Another parasitoid, Phobocampe bicingulata, showed positive density dependence on the smaller spatial scale and had a variable response on the larger scale, but exhibited negative density dependence over time. The results of this study caution against drawing conclusions concerning population regulation on the grounds of spatial density dependence alone. Larval parasitoids apparently do not maintain low densities in the E. autumnata populations studied. However, they may suppress E. autumnata densities to a level low enough for density-dependent mortality factor(s) to become regulating. Among other mortality factors of E. autumnata, pupal predation has been found to be temporally positively density-dependent. Received: 19 October 1999 / Accepted: 10 January 2000     

Early survival in roe deer: causes and consequences of cohort variation in two contrasted populations

Time- and sex-specific summer survival of roe deer fawns was estimated using capture-mark-recapture methods in two enclosed populations living in contrasting conditions. The population of Trois Fontaines (eastern France) was roughly constant in size throughout the study period, while in Chizé (western France), the population experienced frequent summer droughts and numbers decreased continuously during the study. Early survival of fawns was low and highly variable over the years at both Chizé and Trois Fontaines, and demonstrated marked variations between cohorts that need to be taken into account when modelling roe deer population dynamics. In Trois Fontaines, fawn survival was positively correlated with early body growth and total rainfall in May and June. In Chizé, fawn survival decreased with increasing density and tended to increase with increasing rainfall in May and June and adult female body mass. These factors explained more than 75% of the variability in early survival observed in both populations. Variation between cohorts had different consequences for the two populations. At Trois Fontaines, cohort variation was limited to a numerical effect on early survival. However at Chizé, cohort variation was long-lasting and affected the phenotypic quality of survivors at later ages, and thereby future survival and breeding abilities (both numerical and quality effects). Male and female fawns had similar survival over their first summer in both populations. This result contrasts with the lower survival of young males often observed in ungulates. Two ultimate causes can be proposed to account for the low and variable survival of roe deer fawns over the first summer: the high energy expenditures incurred by does during each breeding attempt and/or the low absolute body size of newborn roe deer fawns. Received: 28 April 1997 / Accepted: 14 July 1997     

Population and prehistory II: space-limited human populations in constant environments

We present a population model to examine the forces that determined the quality and quantity of human life in early agricultural societies where cultivable area is limited. The model is driven by the non-linear and interdependent relationships between the age distribution of a population, its behavior and technology, and the nature of its environment. The common currency in the model is the production of food, on which age-specific rates of birth and death depend. There is a single non-trivial equilibrium population at which productivity balances caloric needs. One of the most powerful controls on equilibrium hunger level is fertility control. Gains against hunger are accompanied by decreases in population size. Increasing worker productivity does increase equilibrium population size but does not improve welfare at equilibrium. As a case study we apply the model to the population of a Polynesian valley before European contact.     

An experimental study of population regulation in the salamander,Notophthalmus viridescens dorsalis (Urodela: Salamandridae)

Summary The roles of density-dependent larval survival and cannibalism of larvae as potential mechanisms of population regulation in the newt (Notophthalmus viridescens dorsalis) were evaluated in laboratory and field experiments. In laboratory containers, adults cannibalized larvae and large larvae cannibalized smaller larvae. In artificial ponds, larval survival did not depend on initial larval density. No cannibalism could be demonstrated in the complex environment, although the experiment was powerful enough to detect an ecologically relevant difference in survival. Adult growth was negatively correlated with the final biomass of larval newts, suggesting that the two life stages competed for resources. Larval growth rates were negatively correlated with final larval density, suggesting that larvae competed with each other. The proportion of larvae that became sexually mature at age 7 months (paedomorphs and adults that skipped the eft stage) varied inversely with larval density. Therefore, the potential regulatory mechanisms identified in this study are competition within and between life stages.     

From population ecology to metabolism: growth of Trypanosoma evansi,and implications of glucose depletion,in a live host

Little attention has been paid to the potential top-down effects of population ecology on metabolism. Because it is capable of a fast growth that dramatically modifies its blood environment, the parasitic protozoan, Trypanosoma evansi, is a promising model for the study of density-dependent metabolic processes. We assess the in vivo growth rate, doubling time, biomass yield, glycolytic flux, and enzyme expression of T. evansi. Then, we explore the metabolic changes likely occurring during its growth. At low T. evansi densities, most host glucose is used to produce energy, which results in the release of pyruvate. Part of glucose is used for NADPH production through the pentose phosphate pathway. At high T. evansi densities, fructose, mannose, and glycerol become additional energy sources. Part of host glucose is used for biosynthesis and for NADPH production through alternative metabolic pathways, which results in the release of succinate, alanine, and acetate. The ability of organisms to adjust to resource changes is crucial to their survival. Irrespective if the triggering mechanism is direct (nutrient limitation) or indirect (a pheromone-like factor), nutrient availability is necessarily the main evolutionary factor responsible for the density-dependent metabolic properties of trypanosome populations.     

The effect of the rate of successful dispersal of a Phytoseiid mite,Phytoseiulus persimilis Athias-Henriot (Acarina: Phytoseiidae) on the persistence in the interactive system between the predator and its prey

Dispersal behaviour was studied on a predacious phytoseiid mite, Phytoseiulus persimilisAthias-Henriot in response to the density of its prey, Tetranychus kanzawaiKishida . And the effect of the change in the rate of successful dispersal of the predators among patches was tested on the persistence of the predator-prey system. The results of the study are summarized as follows:

1. With a severe decline in prey density available per individual predator, the predators exhibited a marked behavioural change and dispersed to other areas.
2. When two neighboring host plants did not touch with each other and the dispersal of the predators was possible only through a 20cm plywood “bridge” which connected the two plants, the rate of successful dispersal of the predators was only 20 to 25%, whereas it was 95% when the plants touched with each other.
3. In the system where 32 host plants touched with each other, the predators succeeded in immigrating into all the plants by the 16th day, and they completely eliminated all the prey by the 33rd day.
4. In another system, the 32 plants were equally divided into eight “patches” which were connected with each other with two of the bridges and the predators could move between them only through the bridges as mentioned above. In such a system the predators and their prey coexisted about three times longer than in the other one.
5. It is suggested that for a longer continued coexistence of the predators and prey, it would be necessary to introduce more physical barrier against the dispersal of the predators and to provide more chances for the prey to disperse.

     

The effect of the rate of successful dispersal of a phytoseiid mite,Phytoseiulus persimilis Athias-Henriot (Acarina: Phytoseiidae) on the persistence in the interactive system between the predator and its prey

Dispersal behaviour was studied on a predacious phytoseiid mite, Phytoseiulus persimilisAthias-Henriot in response to the density of its prey, Tetranychus kanzawaiKishida . And the effect of the change in the rate of successful dispersal of the predators among patches was tested on the persistence of the predator-prey system. The results of the study are summarized as follows:

1. With a severe decline in prey density available per individual predator, the predators exhibited a marked behavioural change and dispersed to other areas.
2. When two neighboring host plants did not touch with each other and the dispersal of the predators was possible only through a 20cm plywood “bridge” which connected the two plants, the rate of successful dispersal of the predators was only 20 to 25%, whereas it was 95% when the plants touched with each other.
3. In the system where 32 host plants touched with each other, the predators succeeded in immigrating into all the plants by the 16th day, and they completely eliminated all the prey by the 33rd day.
4. In another system, the 32 plants were equally divided into eight “patches” which were connected with each other with two of the bridges and the predators could move between them only through the bridges as mentioned above. In such a system the predators and their prey coexisted about three times longer than in the other one.
5. It is suggested that for a longer continued coexistence of the predators and prey, it would be necessary to introduce more physical barrier against the dispersal of the predators and to provide more chances for the prey to disperse.

     

Predicting fluctuations of reintroduced ibex populations: the importance of density dependence, environmental stochasticity and uncertain population estimates

1. Development of population projections requires estimates of observation error, parameters characterizing expected dynamics such as the specific population growth rate and the form of density regulation, the influence of stochastic factors on population dynamics, and quantification of the uncertainty in the parameter estimates. 2. Here we construct a Population Prediction Interval (PPI) based on Bayesian state space modelling of future population growth of 28 reintroduced ibex populations in Switzerland that have been censused for up to 68 years. Our aim is to examine whether the interpopulation variation in the precision of the population projections is related to differences in the parameters characterizing the expected dynamics, in the effects of environmental stochasticity, in the magnitude of uncertainty in the population parameters, or in the observation error. 3. The error in the population censuses was small. The median coefficient of variation in the estimates across populations was 5.1%. 4. Significant density regulation was present in 53.6% of the populations, but was in general weak. 5. The width of the PPI calculated for a period of 5 years showed large variation among populations, and was explained by differences in the impact of environmental stochasticity on population dynamics. 6. In spite of the high accuracy in population estimates, the uncertainty in the parameter estimates was still large. This uncertainty affected the precision in the population predictions, but it decreased with increasing length of study period, mainly due to higher precision in the estimates of the environmental variance in the longer time-series. 7. These analyses reveal that predictions of future population fluctuations of weakly density-regulated populations such as the ibex often become uncertain. Credible population predictions require that this uncertainty is properly quantified.     

Induction of embryonic dormancy in the calanoid copepod Acartia hudsonica: heritability and phenotypic plasticity in two geographically separated populations

The planktonic copepod Acartia hudsonica apparently requires summer diapause or dormancy to persist in Narragansett Bay, RI. In estuaries to the north, however, active A. hudsonica populations are present year-round and may not express dormancy. Using a full-sibling rearing design and analysis of variance (ANOVA), I assessed the importance of phenotypic and genetic sources of variation in two geographically separate populations, one from Maine and one from Rhode Island. Both populations showed phenotypic plasticity in the percentage of dormant eggs produced. Moreover, experiments revealed significant sibship-environment interactions in both populations, indicating that the phenotypic plasticity has a genetic component. Both populations also revealed a significant amount of genetic variation in the percentage of dormant eggs produced. For the Maine population, broad sense heritability was high (0.91, 1.10) in two short-day (12L:12D) treatments and near zero (0.08) in the long-day treatment (15L:9D). For a Rhode Island population, broad sense heritability was higher (0.95) in a low-temperature short-day treatment (13.5 °C, 12L:12D) than in a high-temperature short-day treatment (17.5 °C, 12L:12D; h²=0.25).     

Variation in plant quality and the population dynamics of herbivores: there is nothing average about aphids

In the attempt to use results from small-scale studies to make large-scale predictions, it is critical that we take into account the greater spatial heterogeneity encountered at larger spatial scales. An important component of this heterogeneity is variation in plant quality, which can have a profound influence on herbivore population dynamics. This influence is particularly relevant when we consider that the strength of density dependence can vary among host plants and that the strength of density dependence determines the difference between exponential and density- dependent growth. Here, we present some simple models and analyses designed to examine the impact of variable plant quality on the dynamics of insect herbivore populations, and specifically the consequences of variation in the strength of density dependence among host plants. We show that average values of herbivore population growth parameters, calculated from plants that vary in quality, do not predict overall population growth. Furthermore, we illustrate that the quality of a few individual plants within a larger plant population can dominate herbivore population growth. Our results demonstrate that ignoring spatial heterogeneity that exists in herbivore population growth on plants that differ in quality can lead to a misunderstanding of the mechanisms that underlie population dynamics.     

Dynamics of springtail and mite populations: the role of density dependence, predation, and weather

Abstract 1. Ecological theory suggests that density‐dependent regulation of organism abundance will vary from exogenous to endogenous factors depending on trophic structure. Changes in abundance of soil arthropods were investigated at three trophic levels, springtails (Collembola), predaceous mites (Acari), and macro‐arthropods (spider, adult and larval beetles, centipedes). Predictions were that springtails are predator regulated and mites are food limited according to the Hairston et al. (1960) model, which predicts alternating regulation by competition and predation from fungi to springtails to mites to macro‐arthropods. The alternate hypothesis was based on the bottom‐up model of trophic dynamics, which predicts that each trophic level is regulated by competition for resources. 2. The relative contributions to springtail and mite population dynamics of endogenous (i.e. density‐dependent population growth related to food availability) and exogenous (i.e. predation and weather) factors were tested using time‐series analysis and experimental manipulation of water conditions. Box patterns were distributed within an aspen forest habitat located in the Canadian prairies and surveyed weekly from May to September 1997–1999. Each box depressed the leaf litter, creating a microhabitat island for soil arthropods that provided counts of invertebrates located immediately beneath the boxes. 3. Strong evidence was found for endogenous control of springtail and mite numbers, indicated by a reduction in population growth related to density in the previous week. Contrary to predictions, no evidence was found for regulation of springtail numbers by mites, or for regulation of mite numbers by macro‐arthropods. Springtail population growth rate was related positively to current springtail density (8 and 23% variation explained) and related negatively to 1‐week lagged density (85 and 58%), and related negatively to temperature (5 and 5%) for time‐series data and for experimental addition of water respectively. Mite population growth rate was related positively to current mite density (54%) and temperature (4%), and negatively to 1‐week lagged mite density (20%) and precipitation (6%) for time‐series analysis. For experimental addition of water, mite growth rate was related positively to current mite density (44%) and temperature (5%), and negatively to 1‐week lagged density (11%). Results differed from the Hairston et al. (1960) model predictions but were consistent with a bottom‐up view that springtail and mite populations were regulated intrinsically by competition for food and secondarily by temperature as a function of reproduction.     

A new system for understanding the biodiversity in different nature reserves: capacity, connectivity and quality of biodiversity

In this paper, we propose a new system for understanding the biodiversity in different conservation areas. It includes three aspects: the capacity, the connectivity and the quality. The capacity refers to the numbers of biodiversity, including absolute and relative richness of the vegetation types N _v and D _v =(N _v −1)/lnA, species numbers S and richness of species d _Gl =(S − 1)/lnA, and germ plasm resources within a nature reserve, and also the potential biological living space offered by the natural resource. It comprises the total biological resources in a nature reserve. The connectivity refers to the flux of biodiversity, including similarity and connected status of the vegetation types SI _Li =2z/(x + y) and species numbers SI _C =2z/(x + y) among different nature reserves. The quality refers to the stability of biodiversity, including relative species richness index RS _Li =d/d _max, relative vegetation richness index RV _Li =D _v /D _maxv , fastness to invasion species ƒ _Li =1−S _i /S _t , weighted values, representativeness and vulnerability of special vegetations, special species, CITES species and rare species as the protected targets.     

A new system for understanding the biodiversity in different nature reserves:capacity,connectivity and quality of biodiversity

In this paper,we propose a new system for understanding the biodiversity in different conservation areas.It includes three aspects:the capacity,the connectivity and the quality.The capacity refers to the numbers of biodiversity,including absolute and relative richness of the vegetation types Nv and Dv = (Nv-1)/lnA,species numbers S and richness of species dGI = (S- 1)/lnA,and germ plasm resources within a nature reserve,and also the potential biological living space offered by the natural resource.It comprises the total biological resources in a nature reserve.The connectivity refers to the flux of biodiversity,including similarity and connected status of the vegetation types SILi = 2z/(x + y) and species numbers SIc = 2z/(x + y) among different nature reserves.The quality refers to the stability of biodiversity,including relative species richness index RSLi = d/dmax,relative vegetation richness index RVLi =Dv/Dmaxv,fastness to invasion species fLi = 1-Si/St,weighted values,representativeness and vulnerability of special vegetations,special species,CITES species and rare species as the protected targets.     

Breeding ecology and male spawning success in two hill-stream populations of the freshwater goby,Padogobius martensi

Synopsis This investigation was carried out at two study sites to examine the influence of shelter availability, nest site quality and male body size on spawning success of malePadogobius martensi. At Stirone Stream (first study site) individuals were not randomly distributed with respect to usable stones; larger males and females were consistently found to occupy larger stones. Laboratory experiments showed that the bottom surface area of the shelter was the relevant cue for the choice. The number of nesting males settled in a given area increased with increasing availability of local stone cover; inter-nest distance was directly correlated with male size. At Rivarolo Stream (second study site), which was characterized by lower nest density and a greater shortage of large stones, the pattern of shelter occupation was similar. These results suggest the presence of competition among individuals for the possession of larger-sized shelter sites. At both sites, the number of eggs in the nest increased as stone size increased; however, at the first study site, male body size played a major role in male spawning success as it concerns both the numbers of females mated and the number of eggs, while at the second study site, male body size was not shown to influence male spawning success. Factors responsible for between-site differences in male spawning success are discussed.     

Comparison of male adult population densities of the oriental and artocarpus fruit flies,Dacus spp. (Diptera: Tephritidae), in two nearby villages in Penang,Malaysia

The populations of native male adult oriental fruit fly Dacus dorsalis (Hendel ) and artocarpus fruit fly D. umbrosus (F.) in two selected site (BU and SD) were estimated weekly by the capture-recapture technique using live traps baited with methyl eugenol. In BU where many varieties of fruit trees were grown, the estimated population densities of D. dorsalis were between 980 and 3100 male flies per ha between May and July, 1984. During the same period, in SD where there were fewer number and varieties of fruit trees, the estimated population densities were between 300 and 1000 flies per ha. The estimated population densities of D. umbrosus over the same period were between 570 and 1290 flies per ha in BU; and between 5 and 95 flies per ha in SD. Of a total 6828 marked D. dorsalis flies released only one fly (released 6 weeks earlier in BU) was caught in a different site.     

Making the most of mitochondrial genomes--markers for phylogeny, molecular ecology and barcodes in Schistosoma (Platyhelminthes: Digenea)

An increasing number of complete sequences of mitochondrial (mt) genomes provides the opportunity to optimise the choice of molecular markers for phylogenetic and ecological studies. This is particularly the case where mt genomes from closely related taxa have been sequenced; e.g., within Schistosoma. These blood flukes include species that are the causative agents of schistosomiasis, where there has been a need to optimise markers for species and strain recognition. For many phylogenetic and population genetic studies, the choice of nucleotide sequences depends primarily on suitable PCR primers. Complete mt genomes allow individual gene or other mt markers to be assessed relative to one another for potential information content, prior to broad-scale sampling. We assess the phylogenetic utility of individual genes and identify regions that contain the greatest interspecific variation for molecular ecological and diagnostic markers. We show that variable characters are not randomly distributed along the genome and there is a positive correlation between polymorphism and divergence. The mt genomes of African and Asian schistosomes were compared with the available intraspecific dataset of Schistosoma mansoni through sliding window analyses, in order to assess whether the observed polymorphism was at a level predicted from interspecific comparisons. We found a positive correlation except for the two genes (cox1 and nad1) adjoining the putative control region in S. mansoni. The genes nad1, nad4, nad5, cox1 and cox3 resolved phylogenies that were consistent with a benchmark phylogeny and in general, longer genes performed better in phylogenetic reconstruction. Considering the information content of entire mt genome sequences, partial cox1 would not be the ideal marker for either species identification (barcoding) or population studies with Schistosoma species. Instead, we suggest the use of cox3 and nad5 for both phylogenetic and population studies. Five primer pairs designed against Schistosoma mekongi and Schistosoma malayensis were tested successfully against Schistosoma japonicum. In combination, these fragments encompass 20-27% of the variation amongst the genomes (average total length approximately 14,000bp), thus providing an efficient means of encapsulating the greatest amount of variation within the shortest sequence. Comparative mitogenomics provides the basis of a rational approach to molecular marker selection and optimisation.     

Density-dependent growth and reproduction of the apple snail, Pomacea canaliculata: a density manipulation experiment in a paddy field

To examine density dependence in the survival, growth, and reproduction of Pomacea canaliculata, we conducted an experiment in which snail densities were manipulated in a paddy field. We released paint-marked snails of 15–20 mm shell height into 12 enclosures (pens) of 16 m² at one of five densities – 8, 16, 32, 64, or 128 snails per pen. The survival rate of released snails was 95% and was independent of snail density. The snail density had a significant effect on the growth and egg production of individual snails. This density dependence may have been caused by reduced food availability. The females at high density deposited fewer and smaller egg masses than those at low density, and consequently produced fewer eggs. The females at densities 8 and 16 deposited more than 3000 eggs per female, while the females at density 128 oviposited only 414 eggs. The total egg production per pen was, however, higher at higher snail density. The survival rates of juvenile snails were 21%–37% and were independent of adult density. The juvenile density was positively correlated with the total egg production per pen and hence was higher at higher adult density. However, the density of juveniles larger than 5 mm in shell height, i.e., juveniles that can survive an overwintering period, was not significantly different among density treatments. These results suggest that snail density after the overwintering period is independent of the density in the previous year. Thus, density dependence in growth and reproduction might regulate the population of P. canaliculata in paddies. Received: October 23, 1998 / Accepted: July 16, 1999