The relative importance to population increase of fluctuations in mortality,fecundity and the time variables of the reproductive schedule

Summary Previous authors have used simple models to investigate the relative importance to population increase of variations in the total and age-specific reproductive rates. But while acknowledging that the latter were the product of the age specific birth and death rates, they have used their models only to investigate changes in total or age-specific birth rates and have not been concerned with variations in death rates. This paper extends the use of Lewontin's (1965) model, to a wide range of values of r, the exponential rate of population increase. It shows how the relative importance of changes in certain life-history features can change with r and be reversed when r is near to zero. It is also shown that variations in mortality rate are not necessarily best expressed in analogous terms to variations in birth rate. If more suitable terms are used it is seen that changes in mortality rate can be of varying importance depending on the existing mortality rate. They can be overwhelmingly important when the mortality rate is high.     

Changes in Demographic Parameters of <Emphasis Type="Italic">Macaca fuscata</Emphasis> at Takasakiyama in Relation to Decrease of Provisioned Foods

Though researchers have studied population dynamics extensively in many free-ranging primate populations, information on the relationship between food abundance and demographic parameters based on long-term data is sparse. We examined changes in demographic parameters in relation to decreased provisioning of foods based on data gathered for >50 yr in a provisioned, free-ranging Japanese macaque (Macaca fuscata) population at Takasakiyama, Oita Prefecture, in the southern region of Japan. At Takasakiyama, population size increased linearly because of heavy provisioning in the 1950s and 1960s. The provisioning of food to macaques decreased from 1965. We examined changes in the physique index of full adult females, primiparous age, birth percentage, infant mortality rate, population density, and annual population growth rate relative to the decrease of provisioned foods between the 1970s and 1990s. As a result of the drastic decrease in the amount of foods provisioned, the physique index of full adult females decreased, primiparous age increased, birth percentage decreased, and infant mortality rate increased, such that annual population growth rate diminished substantially. Ours is the first study to demonstrate quantitatively the relationship between provisioned food abundance and demographic parameters based on long-term data.     

Age‐specific mortality analysis of the dry forest kissing bug,Rhodnius neglectus

Age‐specific mortality patterns can be very different across insects with different life histories. Some holometabolous insects (like mosquitoes, fruit flies) show a pattern where mortality rate decelerates at older ages, whereas other holometabolous insects (bruchid beetles) and hemimetabolous insects (cotton stainers, milkweed bugs, and kissing bugs) show an age‐specific mortality pattern that increases through all ages. Kissing bugs are strictly hematophagous and are vectors of Trypanosoma cruzi Chagas, the etiologic agent of Chagas disease. Here, we tested whether cohort data from the dry forest kissing bug, Rhodnius neglectus Lent (Hemiptera: Reduviidae), supports an increase of mortality rate that decelerates with age. We analyzed the age‐specific mortality pattern of a cohort of 250 individuals of R. neglectus. We used a suite of seven models with different degrees of complexity, to model age‐dependent forms of change in mortality rate increase in R. neglectus in the laboratory. We used the Akaike model selection criterion to choose between models that consider absence or presence of mortality deceleration. Five of the seven models (logistic, Gavrilovs, Gompertz, DeMoivre, and exponential) showed a statistically significant fit to the mortality rate. Weak late‐age mortality deceleration in R. neglectus was supported by the best fit (logistic model), and this result is consistent with predictions of the disposable soma theory of senescence.     

Dynamics of an age‐structured population drawn from a random numbers table

I constructed age‐structured populations by drawing numbers from a random numbers table, the constraints being that within a cohort each number be smaller than the preceding number (indicating that some individuals died between one year and the next) and that the first two‐digit number following 00 or 01 ending one cohort’s life be the number born into the next cohort. Populations constructed in this way showed prolonged existence with total population numbers fluctuating about a mean size and with long‐term growth rate (r) ≈ 0. The populations’ birth rates and growth rates and the females’ per capita fecundity decreased significantly with population size, whereas the death rates showed no significant relationship to population size. These results indicate that age‐structured populations can persist for long periods of time with long‐term growth rates of zero in the absence of negative‐feedback loops between a population’s present or prior density and its birth rate, growth rate, and fecundity, contrary to the assumption of density‐dependent regulation hypotheses. Thus, a long‐term growth rate of zero found in natural populations need not indicate that a population’s numbers are regulated by density‐dependent factors.     

Seasonality,Population Dynamics and Production of Daphnia longispina in the Subtropical Lake Bhimtal (U.P.), India

Seasonal variation in population density, birth rate, mortality and production of a daphnid was studied in a high, altitude sub-tropical lake. A bimodal growth curve with maxima during summer and autumn was observed. A positive correlation was reported between fecundity and death rate, probably due to a difference in age specific death rate combined with a high relative frequency of juveniles in a growing population. The production of Daphnia longispina correlates better with temperature than with food. The P/B ratio was higher during summer and autumn months.     

Age- and density-dependent population dynamics in static and variable environments

We consider a general model of a single-species population with age- and density-dependent per capita birth and death rates. In a static environment we show that if the per capita death rate is independent of age, then the local stability of any stationary state is guaranteed by the requirement that, in the region of the steady state, the density dependence of the birth rate should be negative and that of the death rate positive. In a variable environment we show that, provided the system is locally stable, small environmental fluctuations will give rise to small age structure and population fluctuations which are related to the driving environmental fluctuations by a simple “transfer function.” We illustrate our general theory by examining a model with a per capita death rate which is age and density independent and a per capita birth rate which is zero up to some threshold age a₀, adopts a finite density-dependent value up to a maximum age a_o + α, and is zero thereafter. We conclude from this model that resonance due specifically to single-species age-structure effects will only be of practical importance in populations whose members have a life cycle consisting of a long immature phase followed by a short burst of intense reproductive effort (α a_o).     

A case study of the influence of local weather on Aedes aegypti (L.) aging and mortality

The survival rate of mosquitoes is an important topic that affects many aspects of decision‐making in mosquito management. This study aims to estimate the variability in the survival rate of Ae. aegypti, and climate factors that are related to such variability. It is generally assumed that the daily probability of mosquito survival is independent of natural environment conditions and age. To test this assumption, a three‐year fieldwork (2005–2007) and experimental study was conducted at Fortaleza‐CE in Brazil with the aim of estimating daily survival rates of the dengue vector Aedes aegypti under natural conditions in an urban city. Survival rates of mosquitoes may be age‐dependent and statistical analysis is a sensitive approach for comparing patterns of mosquito survival. We studied whether weather conditions occurring on a particular day influence the mortality observed on that particular day. We therefore focused on the impact of daily meteorological fluctuations around a given climate average, rather than on the influence of climate itself. With regard to survival time, multivariate analyses using the stepwise logistic regression model, adjusted for daily temperature, relative humidity, and saturated vapor pressure deficit (SVPD), suggest that age, the seasonal factor, and the SVPD were the most dependent mortality factors. Similar results were obtained using the Cox proportional hazard model, which explores the relationships between the survival and explanatory variables.     

Allometric relationship between body mass and aerobic metabolism in zebrafish Danio rerio

The relationship between body mass (M) and metabolic rate was investigated through the assessment of active (R_A) and standard (R_S) metabolic rate at different life stages in zebrafish Danio rerio (5 day‐old larvae, 2 month‐old juveniles and 6 month‐old adults). Scaling exponents and constants were assessed for standard (R_S = 0·273M^0·965 in mgO₂ g^?1 h^?1) and active metabolic rate (R_A = 0·799M^0·926 in mgO₂ g^?1 h^?1). These data provide the basis for further experiments regarding the effects of environmental factors on aerobic metabolism throughout the life cycle of this species.     

Nonlinear processes in seagrass colonisation explained by simple clonal growth rules

The development of single clones of the seagrass Cymodocea nodosa was analysed using a growth model based on the formation of structures limited by diffusive aggregation. The model implemented the measured clonal growth rules (i.e. rhizome elongation and branching rates, branching angle, and spacer length between consecutive shoots) and shoot mortality rate for C. nodosa at Alfacs Bay (Spain). The simulated patches increased their size nonlinearly with time displaying two different domains of growth. Young patches showed a rapid increase with time of the length of rhizome network and the number of living shoots, which depended on rhizome branching rate, and increased the radial patch size (R_g) algebraically with the number of living shoots as R_g ∝ N_s^1/Df, being D_f the fractal dimension of the patch structure. Patches older than 4 years increased the production of rhizome network and the number of living shoots much more slowly, while their radial patch size behaved as R_g ∝ N_s^0.5 resulting from an internal patch compactation. Moreover, the linear growth rate of the simulated patches changed up to 30 fold during patch development, increasing with increasing patch size until patches reached an intermediate size. The modelled patch development was found to closely reproduce the observed patch structure for the species at the Alfacs Bay (Spain). Hence, the growth of C. nodosa patches initially proceeds with a growth mode controlled by the branching pattern (branching frequency and angle) of the species, producing sparse and elongated patches. Once patches exceed 4–5 years of age and contained >500 shoots, becoming dense and circular, they shifts to a growth model typical of compact structures. These results explain previously unaccounted evidence of the emergence of nonlinear patch growth from simple clonal growth rules, and highlight the importance of branching frequency and angles as critical determinants of the space occupation rate of seagrasses and probably other clonal plants.     

Density‐dependent regulation of the critically endangered black rhinoceros population in Ithala Game Reserve,South Africa

Ensuring the persistence of populations of endangered species requires an understanding of, and response to, the causes of population declines. Species occurring in small populations are vulnerable to stochastic problems that are environmental, demographic, or genetic in nature and can reduce survival as much as the deterministic threats of habitat degradation. Critically endangered black rhinoceros (Diceros bicornis) populations declined throughout Africa since 1960, with numbers steadily increasing at a continental level, but remaining lower than three generations ago. However, size, demographics, trends, and factors affecting these, are poorly known. We used 18 years (1990–2008) of long‐term sightings data from Ithala Game Reserve, KwaZulu‐Natal, South Africa, to determine population estimates, growth rate and fecundity over time, as well as sex and age structure and age‐specific probabilities of survival. Calf survivorship between the ages of 0 and 1 year was 74% for females and 94% for males. Age‐specific survivorship for both sexes was significantly higher from yearling to subadult age‐classes (1–6 years) than for adults (7–30 years). The most frequent cause of mortality was attributed to unknown causes while fighting injuries was recorded as the second most common cause of mortality, particularly among subadult and adult males. There was no significant difference in the sex ratios at birth, although the proportion of females in the population was 0.58. There was strong evidence for density‐dependent regulation, with density in conception year a key driver of population performance (birth rate). The population does not appear to be at ecological carrying capacity; however, social effects are delaying conception. To mitigate density‐dependent social effects, we recommend an adaptive management strategy of pre‐selecting individuals for removal from the reserve, so as to maintain stability in the social organization of the population.     

The influence of local environment on the aging and mortality ofAedes aegypti (L.): Case study in Fortaleza‐CE,Brazil

It is generally assumed that the daily probability of survival of mosquitoes is independent of age. To test this assumption we have conducted a three‐year experimental fieldwork study (2005–2007) at Fortaleza‐CE in Brazil, determining daily survival rates of the dengue vector Aedes aegypti (L.). Survival rates of adult Ae. aegypti may be age‐dependent and the statistical analysis is a sensitive approach for comparing patterns of mosquito survival. The mosquito survival data were better fit by a Weibull survival function than by the more traditionally used Gompertz or logistic survival functions. Gompertz, Weibull, or logistic survival functions often fit the survival, and the tails of the survival curves usually appear to fall between the values predicted by the three functions. We corroborate that the mortality of Ae. aegypti in semi‐natural conditions may no more be considered as a constant phenomenon during the life of adult mosquitoes but varies according to the age and environmental conditions under a tropical climate. This study estimates the variability in the survival rate of Ae. aegypti and environmental factors that are related to such variability. The statistical analysis shows that the fitting ability, concerning the hazard function, was in decreasing order: Seasonal Cox, the three‐parameter Gompertz, and the three‐parameter Weibull, that was similar to the three‐parameter logistic. The advantage of using the Cox model is that it is convenient for exploring the relationship between survival and several explanatory variables. The Cox model has the advantage of preserving the variable in its original quantitative form and of using a maximum of information. The survival analyses indicate that mosquito mortality is both age‐ and environment‐dependent.     

Using viable eggs to accurately determine the demographic and predation potential of Harmonia dimidiata (Coleoptera: Coccinellidae)

Life table parameters and predation rate of the coccinellid predator Harmonia dimidiata F. fed on Aphis gossypii Glover were determined at 25°C using the age‐stage, two‐sex life table. When the total number of eggs were included in our calculations, the intrinsic rate of increase (r), finite rate (λ), net reproductive rate (R₀) and mean generation time (T) for H. dimidiata were 0.1354/day, 1.1450/day, 280.8 offspring and 41.6 day, respectively. These values were significantly different, however, only when viable (=hatchable) H. dimidiata eggs were counted (0.0909/day, 1.0952/day, 67.6 offspring and 46.3 day, respectively). The values obtained using all eggs did not realistically reflect the effect of variable hatch rate and true population parameters of H. dimidiata. We therefore excluded unhatched eggs from our data and demonstrated mathematically that in future demographic studies, it should be a standard procedure to exclude all unhatched eggs when analysing hatch rates that vary with maternal age. A mathematical proof was derived in this study to substantiate this. To observe and quantify variations that occur in the predation rate due to the age and stage of the predator, the daily number of A. gossypii consumed by individual H. dimidiata was analysed using the age‐stage, two‐sex life table. The net predation rate of H. dimidiata on A. gossypii was 14 804 aphids. The transformation rate Q_p showed that the predator needs to consume an average of 219.1 aphids to produce one viable egg. The finite predation rate of H. dimidiata was 125.7 when only hatchable eggs were included in the analysis. Because the age‐stage, two‐sex life table takes both of the sexes and the variable predation rate occurring among stages into consideration, it becomes possible to use the population projection to quantify and time biological control procedures.     

Growth and stomatal responses of wheat seedlings to spatial and temporal variations in soil strength of bi-layered soils

Leaf growth and stomatal behaviour are sensitive to variations in soil mechanical resistance to penetration (Rs). That resistance is strongly influenced by soil water constant, density and texture. As such it is therefore an inherently variable and changing characteristic of the roots natural environment. Leaf responses to spatial and temporal variations in Rs were analysed in wheat using two kinds of simplified model systems: (a) bi-layered soils made of either a low Rs layer on top of a high Rs layer, or the converse, (b) soils where, after enduring high Rs, the whole root system was suddenly exposed to lower Rs by raising soil water content.Both leaf expansion rate and stomatal conductance responded to some roots meeting a new soil layer and also to a step change in impedance to the bulk of roots. These responses could not be ascribed to variations in water or nutrient status per se and strengthen the case for the involvement of some kind of chemical signalling of Rs to leaf cells. Moreover, a striking and novel feature of these responses is that they were always detected with a significant time-lag after the change in Rs had first been experienced. It is concluded that leaf biological age is a paramount factor in explaining such a lag. These data reveal that leaf sensitivity to Rs is mostly confined to early developmental stages preceding blade emergence. However, they also point to the contribution of additional factors, raising the questions of the role of root parts behind the tip and of threshold-type leaf responses to stress induced root signals.Keywords: Soil strength, root impedance, bi-layered soil, wheat, leaf expansion, stomatal conductance, critical biological age.      

Diurnal and seasonal variations in stomatal conductance of rice at elevated atmospheric CO2 under fully open‐air conditions

Understanding of leaf stomatal responses to the atmospheric CO₂ concentration, [CO₂], is essential for accurate prediction of plant water use under future climates. However, limited information is available for the diurnal and seasonal changes in stomatal conductance (g_s) under elevated [CO₂]. We examined the factors responsible for variations in g_s under elevated [CO₂] with three rice cultivars grown in an open‐field environment under flooded conditions during two growing seasons (a total of 2140 individual measurements). Conductance of all cultivars was generally higher in the morning and around noon than in the afternoon, and elevated [CO₂] decreased g_s by up to 64% over the 2 years (significantly on 26 out of 38 measurement days), with a mean g_s decrease of 23%. We plotted the g_s variations against three parameters from the Ball‐Berry model and two revised versions of the model, and all parameters explained the g_s variations well at each [CO₂] in the morning and around noon (R² > 0.68), but could not explain these variations in the afternoon (R² < 0.33). The present results provide an important basis for modelling future water use in rice production.     

The effects of productivity and seasonality on life history: comparing age at maturity among moose (Alces alces) populations

The productivity hypothesis in respect of an animal species’ geographical range predicts that whereas higher productivity at the equatorial periphery of a species’ range favours superior competitors, lower productivity at the centre of a species’ range favours high reproduction and reduced competitive traits. I test whether life‐history patterns follow this hypothesis, using demographic data from 15 Canadian moose (Alces alces) populations. Two models are contrasted; the first assumes that intraspecific variation in age at maturity is explained proximately by density and juvenile mortality. Age at maturity was found to increase with decreasing juvenile mortality (P = 0.01) and increasing density (P = 0.006). To test the productivity hypothesis, the second model additionally included primary productivity and seasonality as geographical explanatory variables that would ultimately influence age at maturity via juvenile mortality and density. Path analysis indicated that including productivity and seasonality improved the model predictions of variation in age at maturity (R_a² 0.56 and 0.85). In bivariate comparisons, seasonality was negatively associated (P = 0.01) with age at maturity. In the best model, however, primary productivity was the environmental variable that explained 25% of the variance in age at maturity, and forest cover replaced seasonality as an explanatory variable. The positive association between primary productivity and age at maturity is consistent with the productivity hypothesis. Relative to populations that lived at the centre of the species’ range (51°N), moose populations living in relatively high productivity and low seasonality environments (equatorial periphery of species’ range; 48°N) experienced less juvenile mortality, more variable year‐to‐year density, higher relative density and slower life history (slower growth rate, later age at maturity, lower fecundity).     

Non-uniform stomatal closure and the apparent convexity of the photosynthetic photon flux density response curve

The photosynthetic response of leaves to photosynthetic photon flux density (PPFD) may be described by parameters for the rate of dark respiration (R), the initial slope (Ф), the PPFD-saturated rate of net photosynthesis (A_mQ) and the apparent convexity (θ_a). We tested the hypothesis that non-uniformity in stomatal aperture across a leaf results in a clearly lower θ_a because PPFD saturation will occur at different irradiances in different regions of the leaf. A computer model was constructed to simulate the effects of bell-shaped and other distributions of stomatal conductance for CO₂ (g_s) across a model leaf. In the model, Ф and A_mQ decreased by up to 50% whereas θ_a decreased by at most 8%, essentially negating the hypothesis. The relationship between photosynthetic rate and g_s determined the size of the responses of θ_a and Ф to non-uniform g_s. In support of the model, experiments on sunflower leaves showed that Ф and A_mQ decreased by 32 and 52%, respectively, while no measurable change in θ_a occurred, when abscisic acid was used to induce patchiness in g_s. Although not all possible patterns of non-uniform stomatal conductance have been tested, it appears that if large variations in θ_a occur in nature they do not result from non-uniform g_s.     

Population dynamics and production of Daphnia hyalina Leydig and Daphnia cucullata Sars in Tjeukemeer

As part of a research programme on the food chains in Tjeukemeer, the Daphnia hyalina and Daphnia cucullata populations were studied for three successive years. To analyse the factors regulating the production of these two species, their population parameters (density, size distribution, fecundity) and population dynamics (birth rate, mortality rate) were studied and related to environmental factors. Since Daphnia in Tjeukemeer shows continuous recruitment, the population dynamics model INSTAR was developed and used to integrate field data with laboratory data on development rates and length-weight relationships. The dynamics of the Daphnia species are mainly regulated by temperature and fish predation, the latter affects both birth rate and mortality. Total annual Daphnia production was 3.1–6.9 g org. dry wt M^–2, and annual P/B ratio ranged from 25 to 40 for D. cucullata and from 45 to 49 for D. hyalina.     

Population dynamics of a thistle-feeding lady beetle Epilachna niponica (Coccinellidae: Epilachninae) in Kanazawa, Japan. 1. Adult demographic traits and population stability

 Population dynamics of a thistle-feeding univoltine lady beetle, Epilachna niponica Lewis (Coleoptera: Coccinellidae), was studied from 1996 to 1999 in Yuwaku, Kanazawa, Japan. The lady beetles often reached such a high density level that food was depleted. The Jolly–Seber method was used for adult marking, release, and recapture data to estimate population parameters of adult number, daily resident rate, longevity, reproductive rate (R, the number of new adults produced per overwintered adults), and survival rate of new adults to the reproductive seasons (S _w). These estimates were compared with those of the Asiu, Kutsuki (A and F), and Kyoto populations, which were previously studied with similar methods and have similar intensities. Asiu and Kutsuki F populations remained at a rather low density with a low R, while Kutsuki A and Kyoto populations reached a high density where food depletion occurred with a high R value. The Yuwaku population often reached a food-depleting level as in the Kutsuki A and Kyoto populations. It also shared the short life span of overwintered adults (13.5 days) of other high-density populations; however, it showed much shorter longevity of new adults (36.6 days), much lower R (1.0–2.5), and higher S _w (43%–53%). In some traits the Yuwaku population was similar to the Asiu population: low R, high S _w, and low population variability (SD of log densities; 0.103 and 0.115 for overwintered and new adults, respectively, which were lowest among the populations). Received: July 26, 2001 / Accepted: May 21, 2002     

Mortality limits used in wind energy impact assessment underestimate impacts of wind farms on bird populations

1. The consequences of bird mortality caused by collisions with wind turbines are increasingly receiving attention. So‐called acceptable mortality limits of populations, that is, those that assume that 1%–5% of additional mortality and the potential biological removal (PBR), provide seemingly clear‐cut methods for establishing the reduction in population viability.
2. We examine how the application of these commonly used mortality limits could affect populations of the Common Starling, Black‐tailed Godwit , Marsh Harrier, Eurasian Spoonbill, White Stork, Common Tern, and White‐tailed Eagle using stochastic density‐independent and density‐dependent Leslie matrix models.
3. Results show that population viability can be very sensitive to proportionally small increases in mortality. Rather than having a negligible effect, we found that a 1% additional mortality in postfledging cohorts of our studied populations resulted in a 2%–24% decrease in the population level after 10 years. Allowing a 5% mortality increase to existing mortality resulted in a 9%–77% reduction in the populations after 10 years.
4. When the PBR method is used in the density‐dependent simulations, the proportional change in the resulting growth rate and carrying capacity was species‐independent and largely determined by the recovery factor (F_r). When F_r = 1, a value typically used for robust populations, additional mortality resulted in a 50%–55% reduction in the equilibrium density and the resulting growth rate. When F_r = 0.1, used for threatened populations, the reduction in the equilibrium density and growth rate was about 5%.
5. Synthesis and applications. Our results show that by allowing a mortality increase from wind farm collisions according to both criteria, the population impacts of these collisions can still be severe. We propose a simple new method as an alternative that was able to estimate mortality impacts of age‐structured stochastic density‐dependent matrix models.

     

Microgeographic structure of Anopheles gambiae in western Kenya based on mtDNA and microsatellite loci

The population genetic structure of the Anopheles gambiae in western Kenya was studied using length variation at five microsatellite loci and sequence variation in a 648-nt mtDNA fragment. Mosquitoes were collected from houses in villages spanning up to 50 km distance, The following questions were answered, (i) Are mosquitoes in a house more related genetically to each other than mosquitoes between houses? (ii) What degree of genetic differentiation occurs on these geographical scales? (iii) How consistent are the results obtained with both types of genetic markers? At the house level, no differentiation was detected by F_ST and R_ST, and the band sharing index test revealed no significant associations of alleles across loci. Likewise, indices of kinship based on mtDNA haplotypes in houses were even lower than in the pooled sample. Therefore, the hypothesis that mosquitoes in a house are more related genetically was rejected. At increasing geographical scales, microsatellite allele distributions were similar among all population samples and no subdivision of the gene pool was detected by F_ST or R_ST. Likewise, estimates of haplotype divergence of mtDNA between populations were not higher than the within population estimates, and mtDNA-based F_ST values were not significantly different from zero. That sequence variation in mtDNA provided matching results with microsatellite loci (while high genetic variation was observed in all loci), suggested that this pattern represents the whole genome. The minimum area associated with a deme of A. gambiae in western Kenya is therefore larger than 50 km in diameter.