Spatial and temporal variability in a butterfly population

Summary The dynamics of a butterfly (Plebejus argus) population were analysed at two levels, (i) the population as a whole and (ii) sections within the population. Some sections of the population fluctuated out of synchrony with others, such that the variability [SD Log(Density+1)] shown by the population as a whole was less than the variability shown by each part of the population — overall temporal variability was dampened by spatial asynchrony. Since observed population variability depends on the spatial scale that is sampled, comparisons of population variability among taxa should be carried out only with caution. Implications for island biogeography and conservation biology are discussed.     

The fast–slow continuum and mammalian life-history patterns: an empirical evaluation

The fast–slow continuum hypothesis has been proposed to explain the diversity of life-history patterns exhibited by biological populations, but the quantification and population-dynamic consequences of the continuum has remained unclear. I used the ratio of fertility rate to age at first reproduction (F/α ratio) to quantify the tempo of life-history of 138 populations of mammals, and investigated the life-history and population-dynamic consequences of being “fast” or “slow”. “Fast” mammals (F/α>0.60) were characterized by early maturity, short lifespans, low survival rates, and high fertility and projected population growth rate (λ) compared to “slow” (F/α<0.15) mammals. In “fast” populations, λ was overwhelmingly most sensitive to changes in reproductive parameters (age at first reproduction and fertility rates) and relatively insensitive to changes in survival rates. In “slow” populations, λ was very sensitive to changes in juvenile or adult survival rates, and relatively insensitive to changes in reproductive parameters. The pattern of relationships between the F/α ratio and life-history variables, λ, and elasticity of λ to changes in life-history variables persisted even after the effects of body size and phylogeny were statistically removed. These results suggest that fast–slow continuum in mammalian life-history is independent of body size or phylogeny, that the F/α ratio adequately quantifies the position of a population along a fast–slow continuum, and that the tempo of life- histories has substantial population-dynamic consequences.

Zusammenfassung

Die r-K-Kontinuum-Hypothese wurde aufgestellt, um die Diversität von ,,life-history“-Mustern biologischer Populationen zu erklären, aber die Quantifizierung und die Kosnsequenzen für die Populationsdynamik des Kontinuums blieben unklar. Ich benutze das verhältnis der Fortpflanzungsrate zum Fortpflanzungsalter (F/α-Verhältnis) um die Geschwindigkeit der ,,life-history“ von 138 Populationen von Säugetieren zu quantifizieren und untersuchte die Konsequenzen fur die Lebensweise sowie die Populationsdynamik des,,schnell“oder,,langsam “-Seins. ,,Schnelle“Säugetiere (F/α>0.60) waren durch eine frühe Reife, kurze Lebenszeiten, geringe Überlebensraten sowie durch eine große Fertilität und hochgerechnete Populationswachstumsrate (λ) im Vergleich zu ,,langsamen“(F/α<0.15) Säugetieren charakterisiert. In ,,schnellen“ Population reagierte (λ) überwältigend sensibel auf Änderungen in den Fortpflanzungsparametern (Fortpflanzunsalter und Fertilitätsrate) und relativ gering auf Veräanderungen in der Überlebensrate. In ,,langsamen“ Populationen reagierte (λ) sehr sensibel auf Veräanderungen in den reproduktiven Parametern. Das Muster der Beziehung zwischen dem (F/α-Verhältnis) und den Variablen der ,,life-history“,λ, und die Elastizität von λ gegenüber Veränderungen in den variablen der Lebensweise bliev sogar bestehen, nachdem die Effekte von Körpergröße und Phylogenese statistisch eliminiert wurden. Diese Ergebnisse lassen vermuten, dass das r-K-Kontinuum in der ,,life-history“der Säugetiere unabhängig von der Körpergröße und Phylogenie ist, dass das F/α-Verhältnis die Position einer Population im r-K-Kontinuum quantifiziert und dass die Geschwindigkeit der,,life-history“beachtliche konsequenzen fur die Populationsdynamik hat.     

Demography of the demosponge Amphimedon compressa: Evaluation of the importance of sexual versus asexual recruitment to its population dynamics

We used size-based population matrix models to describe the demography of the demosponge Amphimedon compressa. The relative importance of growth, survivorship, and recruitment to population growth (λ) was assessed by performing elasticity and life table response experiment (LTRE) analyses. We also evaluated the relative contribution of sexual and asexual recruitment to λ by analyzing four different scenarios: (1) the combined impact of sexual and asexual recruitment, (2) the impact of only sexual recruits, (3) the impact of only asexual fragmentation, and (4) the impact of no recruitment. Size-based transition matrices were parameterized with field data collected at two sites in the southwest coast of Puerto Rico: Media Luna West (MLW), a reef exposed to high water movement and at Las Pelotas (LP), a reef experiencing low water motion. Estimated λ of 0.8940 and 0.7973 at MLW and LP respectively suggest that both populations are declining. Elasticity analysis indicated that survivorship may be the most important contributor to λ at both sites. However, λ at MLW was influenced more by the survivorship of small individuals whereas survivorship of the large size-class contributed the most to λ at LP. LTRE analysis indicates that the difference in λ between sites was mostly due to difference in survivorship of small sponges. At both sites, λ decreased considerably when sexual recruitment was excluded from the model whereas the absence of asexual recruitment barely changed λ. Therefore, it is suggested that sexual recruitment plays a major role (in comparison to asexual fragmentation) in the population dynamics of this sponge at the studied sites. We conclude that spatial variability in water motion plays an important role in population dynamics of A. compressa by influencing survivorship patterns, including the relative contribution of asexual and sexual recruitment to population growth.     

Risky Business: Temporal and Spatial Variation in Preindustrial Dryland Agriculture

Traditional dryland agriculture in the Pacific island was often labor-intensive and risky, yet settlement and farming in dry areas played an important role in the development of Polynesian societies. We investigate how temporal and spatial climatic fluctuations shape variation in agricultural production across dryland landscapes. We use a model that couples plant growth, climate, and soil organic matter dynamics, together with data from Kohala, Hawai'i, to understand how temperature, rainfall, nitrogen availability, and cropping activity interact to determine yield dynamics through time and space. Due to these interactions, the statistical characterization of rainfall alone is a poor characterization of agricultural yield. Using a simple linear model of human population dynamics, we show that the observed yield variation can affect long-term population growth substantially. Our approach to analyzing spatial and temporal fluctuations in food supply, and to interpreting the population consequences of these fluctuations, provides a quantitative evaluation of agricultural risk and human carrying capacity in dry regions.     

Stochastic density dependence in population size of a benthic clonal invertebrate: the regulating role of fission

The influence of environmental variation on the demography of clonal organisms has been poorly studied. I utilise a matrix model of the population dynamics of the intertidal zoanthid Palythoa caesia to examine how density dependence and temporal variation in demographic rates interact in regulating population size. The model produces realistic simulations of population size, with erratic fluctuations between soft lower and upper boundaries of approximately 55 and 90% cover. Cover never exceeds the maximum possible of 100%, and the population never goes to extinction. A sensitivity analysis indicates that the model’s behaviour is driven by density dependence in the fission of large colonies to produce intermediate sized colonies. Importantly, there is no density-dependent mortality in the model, and density dependence in recruitment, while present, is unimportant. Thus it appears that the main demographic processes which are considered to regulate population size in aclonal organisms may not be important for clonal species. Received: 18 August 1999 / Accepted: 29 October 1999     

Estimating susceptibility of biological control agents to pesticides: influence of life history strategies and population structure

In this study we examined the influences that differing life history strategies and population structures at the time of pesticide exposure have on population susceptibility to pesticides. We used life table data and a matrix projection model to incorporate combinations of mortality (lethal effect) and reductions in fecundity (sublethal effect) into estimates of intrinsic population growth rates (r) for a predator, the seven-spot lady beetle, Coccinella septempunctata L., and its prey, the pea aphid, Acyrthosiphon pisum Harris, and an aphid parasitoid, Diaeretiella rapae (M’Intosh). All three species exhibited differences in key life history variables. The aphid had the highest r and shortest generation time, the ladybeetle had the lowest r and longest generation, while the parasitoid exhibited intermediate life history characteristics. When the model was run with populations started as neonates (aphids) or eggs (lady beetle, parasitoid) for each species, ladybeetle populations were much more susceptible than either aphid or parasitoid populations 30 days after simulated exposure to a pesticide. For example, 50% mortality and a 50% reduction in fecundity resulted in a population headed toward extinction (negative r) for the ladybeetle while the parasitoid population grew exponentially (positive r) even after sustaining 70% mortality and a 70% reduction in fecundity. The aphid species maintained exponential growth after sustaining 80% mortality and an 80% reduction in fecundity. Thus, differences in life history variables accounted for the greater susceptibility of the ladybeetle to a pesticide than its aphid prey or the parasitoid over a set time interval. These differences in susceptibility were greatly reduced when the model was run starting with a mixed age/stage population (the stable age distribution) for each species indicating that population structure at the time of pesticide exposure plays a critical role in population susceptibility. These results suggest that life history attributes as well as population structure at the time of pesticide exposure both play a major role in population susceptibility to pesticides, highlighting the need to explicitly consider differences in life history variables among species when calculating compatibility of pesticides and biological control agents as well as the population structure of beneficial species at the time of pesticide application.     

Empirical and virtual investigation of the population dynamics of an alien plant under the constraints of local carrying capacity: Heracleum mantegazzianum in the Czech Republic

During the last decades, invasive alien species have become a global concern because of their ecological and economic impact. Heracleum mantegazzianum (Apiaceae), is a tall monocarpic perennial native to Caucasus and invasive in Europe since the 1950s. Within an interdisciplinary EU project aimed at assessing suitable management strategy, we analysed the demography and ecology of this species in its invasive range. The monitoring of population dynamics in the Czech Republic led to the result that in the observed sites the species showed decreasing populations. To find an explanation for this unexpected result, two types of models were parameterized, based on the empirical data: (1) a stage-based transition matrix model, which projected a continuous negative development, and (2) a spatially explicit individual-based model (IBM), including individual variation. This second model was able to create a population with steady individual numbers. Analyses of the simulation showed that in more than 54% of the simulated years (n=5000) the growth rate was smaller than one. Still, population increase in the remaining years was sufficient to sustain a population. Nevertheless long-term observations document an invasive behaviour of the observed populations. Hence, we could assume temporal changes in the course of an invasion and thus wanted to evaluate the probability of sampling negative growth in dependence of time since first invasion. By using a method from ‘Virtual Ecology’, we approached the question: first we create an invasive population, based on the empirical data of H. mantegazzianum and second empirical sampling techniques were mimicked using the Virtual Ecologist approach. The results demonstrate how the probability of sampling negative growth increases with time since first invasion. Hence, we assume that the studied populations have already reached a maximum of their local invasive potential and thus stagnate in their size.     

Reproductive fitness,population size and genetic variation in Muscari tenuiflorum (Hyacinthaceae): The role of temporal variation

In plant populations a positive correlation between population size, genetic variation and fitness components is often found, due to increased pollen limitation or reduced genetic variation and inbreeding depression in smaller populations. However, components of fitness also depend on environmental factors which can vary strongly between years. The dry grassland species Muscari tenuiflorum experiences long term habitat isolation and small population sizes. We analyzed seed production of M. tenuiflorum in four years and its dependence on population size and genetic variation. Genetic diversity within populations was high (AFLP: H_e = 0.245; allozymes: H_e = 0.348). An analysis of molecular variance revealed considerable population differentiation (AFLP: 26%; allozyme: 17%). An overall pattern of isolation by distance was found, which, however was not present at distances below 20 km, indicating stronger effects of genetic drift. Genetic diversity was positively correlated to population size. Self pollination reduced seed set by 24%, indicating inbreeding depression. Reproductive fitness was not correlated to genetic diversity and a positive correlation with population size was present in two of four study years. The absence of a general pattern stresses the importance for multi-year studies. Overall, the results show that despite long term habitat isolation M. tenuiflorum maintains seed production in many years independent of population size. The long term persistence of populations is thus expected to depend less on intrinsic genetic or demographic properties affecting seed production but on successful plant establishment and persistence, which latter are based on conservation and protection of suitable habitats.     

Long-term demographic analysis in goshawk <Emphasis Type="Italic">Accipiter gentilis</Emphasis>: the role of density dependence and stochasticity

Density dependence and environmental stochasticity are both potentially important processes influencing population demography and long-term population growth. Quantifying the importance of these two processes for population growth requires both long-term population as well as individual-based data. I use a 30-year data set on a goshawk Accipiter gentilis population from Eastern Westphalia, Germany, to describe the key vital rate elements to which the growth rate is most sensitive and test how environmental stochasticity and density dependence affect long-term population growth. The asymptotic growth rate of the fully age-structured mean matrix model was very similar to the observed one (0.7% vs. 0.3% per annum), and population growth was most elastic to changes in survival rate at age classes 1-3. Environmental stochasticity led only to a small change in the projected population growth rate (between -0.16% and 0.67%) and did not change the elasticities qualitatively, suggesting that the goshawk life history of early reproduction coupled with high annual fertility buffers against a variable environment. Age classes most crucial to population growth were those in which density dependence seemed to act most strongly. This emphasises the importance of density dependence as a regulatory mechanism in this goshawk population. It also provides a mechanism that might enable the population to recover from population lows, because a mean matrix model incorporating observed functional responses of both vital rates to population density coupled with environmental stochasticity reduced long-term extinction risk of 30% under density-independent environmental stochasticity and 60% under demographic stochasticity to zero.     

Numerical algorithms for estimation and calculation of parameters in modeling pest population dynamics and evolution of resistance

Computational simulation models can provide a way of understanding and predicting insect population dynamics and evolution of resistance, but the usefulness of such models depends on generating or estimating the values of key parameters. In this paper, we describe four numerical algorithms generating or estimating key parameters for simulating four different processes within such models. First, we describe a novel method to generate an offspring genotype table for one- or two-locus genetic models for simulating evolution of resistance, and how this method can be extended to create offspring genotype tables for models with more than two loci. Second, we describe how we use a generalized inverse matrix to find a least-squares solution to an over-determined linear system for estimation of parameters in probit models of kill rates. This algorithm can also be used for the estimation of parameters of Freundlich adsorption isotherms. Third, we describe a simple algorithm to randomly select initial frequencies of genotypes either without any special constraints or with some pre-selected frequencies. Also we give a simple method to calculate the “stable” Hardy–Weinberg equilibrium proportions that would result from these initial frequencies. Fourth we describe how the problem of estimating the intrinsic rate of natural increase of a population can be converted to a root-finding problem and how the bisection algorithm can then be used to find the rate. We implemented all these algorithms using MATLAB and Python code; the key statements in both codes consist of only a few commands and are given in the appendices. The results of numerical experiments are also provided to demonstrate that our algorithms are valid and efficient.     

Integral projection models perform better for small demographic data sets than matrix population models: a case study of two perennial herbs

1. Matrix population models are widely used to describe population dynamics, conduct population viability analyses and derive management recommendations for plant populations. For endangered or invasive species, management decisions are often based on small demographic data sets. Hence, there is a need for population models which accurately assess population performance from such small data sets.
2. We used demographic data on two perennial herbs with different life histories to compare the accuracy and precision of the traditional matrix population model and the recently developed integral projection model (IPM) in relation to the amount of data.
3. For large data sets both matrix models and IPMs produced identical estimates of population growth rate (λ). However, for small data sets containing fewer than 300 individuals, IPMs often produced smaller bias and variance for λ than matrix models despite different matrix structures and sampling techniques used to construct the matrix population models.
4. Synthesis and applications . Our results suggest that the smaller bias and variance of λ estimates make IPMs preferable to matrix population models for small demographic data sets with a few hundred individuals. These results are likely to be applicable to a wide range of herbaceous, perennial plant species where demographic fate can be modelled as a function of a continuous state variable such as size. We recommend the use of IPMs to assess population performance and management strategies particularly for endangered or invasive perennial herbs where little demographic data are available.     

Simultaneous identification of growth law and estimation of its rate parameter for biological growth data: a new approach

Scientific formalizations of the notion of growth and measurement of the rate of growth in living organisms are age-old problems. The most frequently used metric, “Average Relative Growth Rate” is invariant under the choice of the underlying growth model. Theoretically, the estimated rate parameter and relative growth rate remain constant for all mutually exclusive and exhaustive time intervals if the underlying law is exponential but not for other common growth laws (e.g., logistic, Gompertz, power, general logistic). We propose a new growth metric specific to a particular growth law and show that it is capable of identifying the underlying growth model. The metric remains constant over different time intervals if the underlying law is true, while the extent of its variation reflects the departure of the assumed model from the true one. We propose a new estimator of the relative growth rate, which is more sensitive to the true underlying model than the existing one. The advantage of using this is that it can detect crucial intervals where the growth process is erratic and unusual. It may help experimental scientists to study more closely the effect of the parameters responsible for the growth of the organism/population under study.     

Spatial and temporal variation in the sediment properties of an intertidal mangrove forest: implications for sampling

Studies of the structure or functioning of intertidal soft sediments often involves collection of biogeochemical data over tidal, diel and seasonal time-scales. Little effort has, however, been made to quantify accurately the time-scales at which these properties vary. Many previous studies collected samples from different sites at different times, potentially resulting in the confounding of spatial and temporal variation. This experiment was designed to determine if time of day or time within the exposure period had any significant effect on measurements of 7 different properties of sediments. Samples of sediment were collected using contact cores at the beginning, middle and end of tidal emersion at each of 9 a.m., 12 p.m. and 3 p.m., with two replicate days of each condition, from mangrove forests fringing Glades Bay, Sydney, Australia. These samples were analysed for water-content, pigments, carbohydrates, grain-size and loss on ignition (LOI), to determine the potential effects of time of day and time within the tidal cycle on these properties of the sediments.Whilst both time of day and time within the tidal cycle were found to occasionally have a significant effect upon the measured properties, most of the variation occurred among sites and between replicate days of each set of conditions. The minimal influence of time of day and time within the tidal cycle show that sampling effort should be preferentially placed into replicating days of sampling and sites because these are the scales with the greatest variation. Differences in the patterns found also depended on whether the data were expressed as content or concentration, the consequences of which are briefly described.     

The population dynamics of Ostertagia gruehneri in reindeer: a model for the seasonal and intensity dependent variation in nematode fecundity

The gastrointestinal nematode Ostertagia gruehneri is a parasite of reindeer that can have a significant impact on host population dynamics. To gain a better understanding of the population dynamics of O. gruehneri, we parameterise a model for its fecundity that describes the observed seasonal and intensity dependent pattern of faecal egg counts well. The faecal egg count model is combined with a model for the seasonal faecal production rate of Svalbard reindeer to obtain quantitative estimates of the fecundity of O. gruehneri. The model is used to evaluate the relative contribution to pasture contamination of variation in the abundance of O. gruehneri and variation in reindeer densities. It is concluded that due to the intensity dependence in nematode fecundity, variation in reindeer population densities is likely to be the most important of these factors for pasture contamination.     

Demographic consequences of age-structure in extreme environments: population models for arctic and alpine ptarmigan

Organisms living in arctic and alpine environments are increasingly impacted by human activities. To evaluate the potential impacts of global change, a better understanding of the demography of organisms in extreme environments is needed. In this study, we compare the age-specific demography of willow ptarmigan (Lagopus lagopus) breeding at arctic and subalpine sites, and white-tailed ptarmigan (L. leucurus) breeding at an alpine site. Rates of egg production improved with age at the alpine and subalpine sites, but the stochastic effects of nest and brood predation led to similar rates of annual fecundity among 1-, 2-, and 3+-year-old females. All populations had short generation times (T<2.7 years) and low net reproductive rates (R ₀<1.2). Stable age distributions were weighted towards 1-year-old females in willow ptarmigan (>59%), and to 3+-year-old females in white-tailed ptarmigan (>47%). High damping ratios (ρ>3.2) indicated that asymptotic estimates were likely to match natural age distributions. Sensitivity and elasticity values indicated that changes in juvenile survival would have the greatest impact on the finite rate of population change (λ) in willow ptarmigan, whereas changes to the survival of 3+-year-old females would have a greater effect in white-tailed ptarmigan. High survivorship buffers white-tailed ptarmigan in alpine environments against the potential effects of climate change on annual fecundity, but may make the species more sensitive to the effects of pollutants or harvesting on adult survival. Conversely, processes that reduce annual fecundity would have a greater impact on the population viability of willow ptarmigan in arctic and subalpine environments. If these same demographic patterns prove to be widespread among organisms in extreme environments, it may be possible to develop general recommendations for conservation of the biological resources of arctic and alpine ecosystems.     

The predictive accuracy of population viability analysis: a test using data from two small mammal species in a fragmented landscape

This study examines the predictive accuracy of the population viability analysis package, ALEX (Analysis of the Likelihood of EXtinction). ALEX was used to predict the probability of patch occupancy for two species of small native Australian mammals (Antechinus agilis and Rattus fuscipes) among 13 patches of suitable habitat in a matrix of plantation pines (Pinus radiata). The study was retrospective, running each simulation from 1900 until 1997, and the model parameterised without knowledge of the 1997 observed field data of patch occupancy. Predictions were made over eight scenarios for each species, allowing for variation in the amount of dispersal between patches, level of environmental stochasticity, and size of habitat patches. Predicted occupancies were compared to the 1997 field data of patch occupancy using logistic regression, testing H _random, that there was no relationship between observed and predicted occupancy, and H _perfect, that there was a perfect, 1:1 relationship between observed and predicted occupancies. Rejection of H _random and failure to reject H _perfect was taken as a good match between observed and predicted occupancies. Such a match was found for one scenario with R. fuscipes, and no scenarios with A. agilis. In general, patch occupancy was underestimated, with field surveys finding that 9 of the 13 patches were occupied by R. fuscipes and 10 by A. agilis. Nonetheless, PVA predictions were in the right direction, whereby patches predicted to have a high probability of occupancy were generally occupied, and vice versa. A post hoc search over additional scenarios found few scenarios with a better match than the original eight. The results of this study support the notion that PVA is best thought of as a relative, rather than absolute predictor of the consequences of management actions in threatened populations.     

Bone histology as an approach to providing data on certain key life history traits in mammals: Implications for conservation biology

The knowledge of the life histories of wild mammals is of crucial importance in the field of conservation management. The endangered status of many species calls for faster data collection that can be used in risk assessment and, ultimately, for designing conservation policies. This study is pioneering the potential of bone histology to provide data on life history traits crucial for conservation biology in long-lived mammals. Long bone cross-sections show pronounced annual cycles of growth arrest allowing application of skeletochronology (counts of lines of arrested growth ‘LAGs’). Consequently, the number of LAGs within the primary fast-growing bone tissue up to the outer cortical slow-growing bone tissue corresponds to the age at first reproduction; whereas the age at death can be estimated by the total number of rest lines throughout the whole of bone cross-section. Furthermore, the diameters of successive growth rings as well as the osteocyte lacuna density may shed light on growth rates. We use the endangered desert dwelling antelope Addax nasomaculatus as a case study. By analyzing different ontogenetic stages in five Addax individuals (three captive and two wild specimens) from a museum collection, we show that bone histology may be a reliable tool for determining certain key life history traits. In our sample, the wild Addax female attained reproductive maturity at three years, whereas the male specimens, both the captive and the wild ones, reached maturity at four years. This is congruent with data from other large antelopes with male-biased size dimorphism, but differs slightly from data on sexual maturity previously published for wild Addax. Moreover, quantification of osteocyte lacunae in both adult males provides a higher cell density in the captive one than in the wild one suggesting the strong effect of constant resources supply in individuals from zoos on growth rates. While age at first reproduction and longevity are essentials parameters to carry out demographic models, growth rates may allow evaluation of the health status of wild populations. This approach may provide useful data on life history traits when applied to bones collected in the wild.