BMC Ecology - Latest articles

To sleep or not to sleep: the ecology of sleep in artificial organisms

Alberto Acerbi, Patrick McNamara and Charles L Nunn — 2008-05-14

Background: All animals thus far studied sleep, but little is known about the ecological factors that generate differences in sleep characteristics across species, such as total sleep duration or division of sleep into multiple bouts across 24-hour period (i.e., monophasic or polyphasic sleep activity). Here we address these questions using an evolutionary agent-based model. The model is spatially explicit, with food and sleep sites distributed in two clusters on the landscape. Agents acquire food and sleep energy based on an internal circadian clock coded by 24 traits (one for each hour of the day) that correspond to 'genes' that evolve by means of a genetic algorithm. These traits can assume three different values that specify the agents' behavior: sleep (or search for a sleep site), eat (or search for a food site), or flexibly decide action based on relative levels of sleep energy and food energy. Individuals with higher fitness scores leave more offspring in the next generation of the simulation, and the model can therefore be used to identify evolutionarily adaptive circadian clock parameters under different ecological conditions. Results: We systematically varied input parameters related to the number of food and sleep sites, the degree to which food and sleep sites overlap, and the rate at which food patches were depleted. Our results reveal that: (1) the costs of traveling between more spatially separated food and sleep clusters select for monophasic sleep, (2) more rapid food patch depletion reduces sleep times, and (3) agents spend more time attempting to acquire the 'rarer' resource, that is, the average time spent sleeping is positively correlated with the number of food patches and negatively correlated with the number of sleep patches. 'Flexible' genes, in general, do not appear to be advantageous, though their arrangements in the agents' genome show characteristic patterns that suggest that selection acts on their distribution. Conclusions: Collectively, the output suggests that ecological factors can have striking effects on sleep patterns. Moreover, our results demonstrate that a simple model can produce clear and sensible patterns, thus allowing it to be used to investigate a wide range of questions concerning the ecology of sleep. Quantitative data presently are unavailable to test the model predictions directly, but patterns are consistent with comparative evidence from different species, and the model can be used to target ecological factors to investigate in future research.

Responses of five small mammal species to micro-scale variations in vegetation structure in secondary Atlantic Forest remnants, Brazil

Thomas Puettker, Renata Pardini, Yvonne Meyer-Lucht and Simone Sommer — 2008-05-05

Background: The Brazilian Atlantic Forest is highly endangered and only about 7% of the original forest remains, most of which consists of fragments of secondary forest. Atlantic Forest small mammals are differently affected by this process of fragmentation and conversion of forest into anthropogenic habitats and present distinct ability to occupy the surrounding altered habitats. We investigated the influence of vegetation structure on the micro-scale distribution of five small mammal species in six secondary forest remnants in an Atlantic Forest fragmented landscape. We examined if the occurrence of small mammal species is influenced by vegetation structure, aiming to investigate if species with different degrees of vulnerability to forest fragmentation are associated to distinct vegetation characteristics. Results: Although vegetation structure differed among fragments, micro-scale distribution of most of the species was influenced by vegetation structure in a similar way in different fragments. Among the three species that were previously shown not to be vulnerable to forest fragmentation, A. montensis and G. microtarsus were present at locations with an open canopy and the occurrence of O. nigripes was associated to a low canopy and a dense understory. On the other hand, from the two species that were shown to be vulnerable to fragmentation, M. incanus was captured most often at locations with a closed canopy while the distribution of D. sublineatus was not clearly influenced by micro-scale variation in vegetation structure. Conclusions: Results indicate the importance of micro-scale variation in vegetation structure for the distribution of small mammal species in secondary forest remnants. Species non-vulnerable to fragmentation occurred at locations with vegetation characteristics of more disturbed forest, while species vulnerable to fragmentation were distributed at locations with older forest characteristics. Results suggest that micro-habitat preferences may be an important factor influencing small mammal capacity to occupy altered habitats and, consequently, their vulnerability to forest fragmentation at a larger spatial scale.

The influence of weather conditions on the activity of high-arctic arthropods inferred from long-term observations

Toke T Hoye and Mads C Forchhammer — 2008-05-02

Background: Climate change is particularly pronounced in the High Arctic and a better understanding of the repercussions on ecological processes like herbivory, predation and pollination is needed. Arthropods play an important role in the high-arctic ecosystem and this role is determined by their density and activity. However, density and activity may be sensitive to separate components of climate. Earlier emergence due to advanced timing of snowmelt following climate change may expose adult arthropods to unchanged temperatures but higher levels of radiation. The capture rate of arthropods in passive open traps like pitfall trap integrates density and activity and, therefore, serves as a proxy of the magnitude of such arthropod-related ecological processes. We used arthropod pitfall trapping data and weather data from 10 seasons in high-arctic Greenland to identify climatic effects on the activity pattern of nine arthropod taxa. Results: We were able to statistically separate the variation in capture rates into a non-linear component of capture date (density) and a linear component of weather (activity). The non-linear proxy of density always accounted for more of the variation than the linear component of weather. After accounting for the seasonal phenological development, the most important weather variable influencing the capture rate of flying arthropods was temperature, while surface-dwelling species were principally influenced by solar radiation. Conclusions: Consistent with previous findings, air temperature best explained variation in the activity level of flying insects. An advancement of the phenology in this group due to earlier snowmelt will make individuals appear earlier in the season, but parallel temperature increases could mean that individuals are exposed to similar temperatures. Hence, the effect of climatic changes on the activity pattern in this group may be unchanged. In contrast, we found that solar radiation is a better proxy of activity levels than air temperature in surface-dwelling arthropods. An advancement of the phenology may expose surface-dwelling arthropods to higher levels of solar radiation, which suggest that their locomotory performance is enhanced and their contribution to ecological processes is increased.

Assessing meiofaunal variation among individuals utilising morphological and molecular approaches: an example using the Tardigrada

Chester J Sands, Peter Convey, Katrin Linse and Sandra J McInnes — 2008-04-30

Background: Meiofauna a multicellular animals captured between sieve size 45 and 1000 microns a are a fundamental component of terrestrial, and marine benthic ecosystems, forming an integral element of food webs, and playing a critical roll in nutrient recycling. Most phyla have meiofaunal representatives and studies of these taxa impact on a wide variety of sub-disciplines as well as having social and economic implications. However, studies of variation in meiofauna are presented with several important challenges. Isolating individuals from a sample substrate is a time consuming process, and identification requires increasingly scarce taxonomic expertise. Finding suitable morphological characters in many of these organisms is often difficult even for experts. Molecular markers are extremely useful for identifying variation in morphologically conserved organisms. However, for many species markers need to be developed de novo, while DNA can often only be extracted from pooled samples in order to obtain sufficient quantity and quality. Importantly, multiple independent markers are required to reconcile gene evolution with species evolution. In this primarily methodological paper we provide a proof of principle of a novel and effective protocol for the isolation of meiofauna from an environmental sample. We also go on to illustrate examples of the implications arising from subsequent screening for genetic variation at the level of the individual using ribosomal, mitochondrial and single copy nuclear markers. Results: To isolate individual tardigrades from their habitat substrate we used a non-toxic density gradient media that did not interfere with downstream biochemical processes. Using a simple DNA release technique and nested polymerase chain reaction with universal primers we were able amplify multi-copy and, to some extent, single copy genes from individual tardigrades. Maximum likelihood trees from ribosomal 18S, mitochondrial cytochrome oxidase subunit 1, and the single copy nuclear gene Wingless support a recent study indicating that the family Hypsibiidae is a non-monophyletic group. From these sequences we were able to detect variation between individuals at each locus that allowed us to identify the presence of cryptic taxa that would otherwise have been overlooked. Conclusions: Molecular results obtained from individuals, rather than pooled samples, are a prerequisite to enable levels of variation to be placed into context. In this study we have provided a proof of principle of this approach for meiofaunal tardigrades, an important group of soil biota previously not considered amenable to such studies, thereby paving the way for more comprehensive phylogenetic studies using multiple nuclear markers, and population genetic studies.

A comparison of plotless density estimators using Monte Carlo simulation on totally enumerated field data sets

Neil A White, Richard M Engeman, Robert T Sugihara and Heather W Krupa — 2008-04-17

Background plotless density estimators are those that are based on distance measures rather than counts per unit area (quadrats or plots) to estimate the density of some usually stationary event, e.g. burrow openings, damage to plant stems, etc. These estimators typically use distance measures between events and from random points to events to derive an estimate of density. The error and bias of these estimators for the various spatial patterns found in nature have been examined using simulated populations only. In this study we investigated eight plotless density estimators to determine which were robust across a wide range of data sets from fully mapped field sites. They covered a wide range of situations including animal damage to rice and corn, nest locations, active rodent burrows and distribution of plants. Monte Carlo simulations were applied to sample the data sets, and in all cases the error of the estimate (measured as relative root mean square error) was reduced with increasing sample size. The method of calculation and ease of use in the field were also used to judge the usefulness of the estimator. Estimators were evaluated in their original published forms, although the variable area transect (VAT) and ordered distance methods have been the subjects of optimization studies. Results An estimator that was a compound of three basic distance estimators was found to be robust across all spatial patterns for sample sizes of 25 or greater. The same field methodology can be used either with the basic distance formula or the formula used with the Kendall-Moran estimator in which case a reduction in error may be gained for sample sizes less than 25, however, there is no improvement for larger sample sizes. The variable area transect (VAT) method performed moderately well, is easy to use in the field, and its calculations easy to undertake. Conclusions: Plotless density estimators can provide an estimate of density in situations where it would not be practical to layout a plot or quadrat and can in many cases reduce the workload in the field.

Effects of intraspecific competition on the life cycle of the stonefly, Nemurella pictetii (Plecoptera: Nemouridae)

Reimo Lieske and Peter Zwick — 2008-04-16

Background: Considerable variation of life cycle duration in given insect species has been frequently recorded. Splitting of populations into cohorts with different life cycle lengths may occur, sometimes even between siblings from the same batch. Larval populations of the stonefly Nemurella pictetii in central Europe regularly split into a very fast developing and a normal univoltine cohort, leading to partial multivoltinism. The causes for such variation remain unknown but presumably act on the larval stage in which most of the life cycle is spent. We therefore studied possible effects of intraspecific competition on growth and development of larvae in the laboratory. Results: Intraspecific competition had important influence on growth and development of the larvae. High larval densities led to reduced growth and retarded development through interference, not through exploitative competition. All specimens were negatively affected by frequent encounters and the resulting disturbance. There were no dominant individuals able to grow and develop faster than the rest, at the expense of the others. Conclusion: Differences in life cycle length of Nemurella pictetii may result from different larval densities in different microhabitats and resultant different degrees of interference competition. Although competition alone probably does not cause splitting of populations into cohorts with different life cycle duration differences in size and development caused by other factors are certainly enhanced by intraspecific competition.

Effects of simulated daily precipitation patterns on annual plant populations depend on life stage and climatic region

Martin Köchy — 2008-03-27

Background: To improve the understanding of consequences of climate change for annual plant communities, I used a detailed, grid-based model that simulates the effect of daily rainfall variability on individual plants in five climatic regions on a gradient from 100 to 800 mm mean annual precipitation (MAP). The model explicitly considers moisture storage in the soil. I manipulated daily rainfall variability by changing the daily mean rain (DMR, rain volume on rainy days averaged across years for each day of the year) by ± 20%. At the same time I adjusted intervals appropriately between rainy days for keeping the mean annual volume constant. In factorial combination with changing DMR I also changed MAP by ± 20%. Results: Increasing MAP generally increased water availability, establishment, and peak shoot biomass. Increasing DMR increased the time that water was continuously available to plants in the upper 15 to 30 cm of the soil (longest wet period, LWP). The effect of DMR diminished with increasing humidity of the climate. An interaction between water availability and density-dependent germination increased the establishment of seedlings in the arid region, but in the more humid regions the establishment of seedlings decreased with increasing DMR. As plants matured, competition among individuals and their productivity increased, but the size of these effects decreased with the humidity of the regions. Therefore, peak shoot biomass generally increased with increasing DMR but the effect size diminished from the semiarid to the mesic Mediterranean region. Increasing DMR reduced via LWP the annual variability of biomass in the semiarid and dry Mediterranean regions. Conclusion: More rainstorms (greater DMR) increased the recharge of soil water reservoirs in more arid sites with consequences for germination, establishment, productivity, and population persistence. The order of magnitudes of DMR and MAP overlapped partially so that their combined effect is important for projections of climate change effects on annual vegetation.

Microsatellite data suggest significant population structure and differentiation within the malaria vector Anopheles darlingi in Central and South America

2008-03-26

Background: Anopheles darlingi is the most important malaria vector in the Neotropics. An understanding of A. darlingi's population structure and contemporary gene flow patterns is necessary if vector populations are to be successfully controlled. We assessed population genetic structure and levels of differentiation based on 1,376 samples from 31 localities throughout the Peruvian and Brazilian Amazon and Central America using 5–8 microsatellite loci. Results: We found high levels of polymorphism for all of the Amazonian populations (mean RS = 7.62, mean HO = 0.742), and low levels for the Belize and Guatemalan populations (mean RS = 4.3, mean HO = 0.457). The Bayesian clustering analysis revealed five population clusters: northeastern Amazonian Brazil, southeastern and central Amazonian Brazil, western and central Amazonian Brazil, Peruvian Amazon, and the Central American populations. Within Central America there was low non-significant differentiation, except for between the populations separated by the Maya Mountains. Within Amazonia there was a moderate level of significant differentiation attributed to isolation by distance. Within Peru there was no significant population structure and low differentiation, and some evidence of a population expansion. The pairwise estimates of genetic differentiation between Central America and Amazonian populations were all very high and highly significant (FST = 0.1859 – 0.3901, P < 0.05). Both the DA and FST distance-based trees illustrated the main division to be between Central America and Amazonia. Conclusion: We detected a large amount of population structure in Amazonia, with three population clusters within Brazil and one including the Peru populations. The considerable differences in Ne among the populations may have contributed to the observed genetic differentiation. All of the data suggest that the primary division within A. darlingi corresponds to two white gene genotypes between Amazonia (genotype 1) and Central America, parts of Colombia and Venezuela (genotype 2), and are in agreement with previously published mitochondrial COI gene sequences interpreted as incipient species. Overall, it appears that two main factors have contributed to the genetic differentiation between the population clusters: physical distance between the populations and the differences in effective population sizes among the subpopulations.

Risk-sensitive foraging and the evolution of cooperative breeding and reproductive skew

Hans J Poethke and Jürgen Liebig — 2008-03-18

Background: Group formation and food sharing in animals may reduce variance in resource supply to breeding individuals. For some species it has thus been interpreted as a mechanism of risk avoidance. However, in many groups reproduction is extremely skewed. In such groups resources are not shared equally among the members and inter-individual variance in resource supply may be extreme. The potential consequences of this aspect of group living have not attained much attention in the context of risk sensitive foraging. Results: We develop a model of individually foraging animals that share resources for reproduction. The model allows analyzing how mean foraging success, inter-individual variance of foraging success, and the cost of reproduction and offspring raising influence the benefit of group formation and resource sharing. Our model shows that the effects are diametrically opposed in egalitarian groups versus groups with high reproductive skew. For individuals in egalitarian groups the relative benefit of group formation increases under conditions of increasing variance in foraging success and decreasing cost of reproduction. On the other hand individuals in groups with high skew will profit from group formation under conditions of decreasing variance in individual foraging success and increasing cost of reproduction. Conclusion: The model clearly demonstrates that reproductive skew qualitatively changes the influence of food sharing on the reproductive output of groups. It shows that the individual benefits of variance reduction in egalitarian groups and variance enhancement in groups with reproductive skew depend critically on ecological and life-history parameters. Our model of risk-sensitive foraging thus allows comparing animal societies as different as spiders and birds in a single framework.

Superior infectivity for mosquito vectors contributes to competitive displacement among strains of dengue virus

2008-02-13

Background: Competitive displacement of a weakly virulent pathogen strain by a more virulent strain is one route to disease emergence. However the mechanisms by which pathogens compete for access to hosts are poorly understood. Among vector-borne pathogens, variation in the ability to infect vectors may effect displacement. The current study focused on competitive displacement in dengue virus serotype 3 (DENV3), a mosquito-borne pathogen of humans. In Sri Lanka in the 1980's, a native DENV3 strain associated with relatively mild dengue disease was displaced by an invasive DENV3 strain associated with the most severe disease manifestations, dengue hemorrhagic fever/dengue shock syndrome (DHF/DSS), resulting in an outbreak of DHF/DSS. Here we tested the hypothesis that differences between the invasive and native strain in their infectivity for Aedes aegypti mosquitoes, the primary vector of DENV, contributed to the competitive success of the invasive strain Results: To be transmitted by a mosquito, DENV must infect and replicate in the midgut, disseminate into the hemocoel, infect the salivary glands, and be released into the saliva. The ability of the native and invasive DENV3 strains to complete the first three steps of this process in Aedes aegypti mosquitoes was measured in vivo. The invasive strain infected a similar proportion of mosquitoes as the native strain but replicated to significantly higher titers in the midgut and disseminated with significantly greater efficiency than the native strain. In contrast, the native and invasive strain showed no significant difference in replication in cultured mosquito, monkey or human cells. Conclusion: The invasive DENV3 strain infects and disseminates in Ae. aegypti more efficiently than the displaced native DENV3 strain, suggesting that the invasive strain is transmitted more efficiently. Replication in cultured cells did not adequately characterize the known phenotypic differences between native and invasive DENV3 strains. Infection dynamics within the vector may have a significant impact on the spread and replacement of dengue virus lineages.