An experimental examination of the effects of habitat quality on the dispersal and local abundance of the butterfly Parnassius smintheus

Abstract 1. Nectar flower abundance was manipulated through flower removal, and sex ratio was manipulated by moving individual butterflies within a series of nine alpine meadows. The movement and abundance of the butterfly Parnassius smintheus in the meadows were monitored using mark–release–recapture methods.
2. A total of 937 butterflies, 698 males and 239 females, was captured. There were 223 observed between-meadow movements. Fifty-two per cent of males and 35% of females moved among meadows.
3. The immigration of male butterflies was related positively to nectar flowers, host plant abundance, and female butterflies. Male emigration was not affected by any of the treatments. The number of males captured was related positively to nectar flowers and host plants but not affected by sex ratio. The number of resident male butterflies was greater in meadows containing flowers and was related positively to host plant abundance, but unaffected by sex ratio.
4. Flower removal, sex ratio, and abundance of Sedum had no significant effect on the abundance, movement, or residence time for female butterflies, in part due to small sample size.
5. The fact that males immigrate to higher quality meadows suggests that male butterflies are assessing meadow quality, either by sampling meadows or potentially from a distance using olfactory cues.     

Temporal autocorrelation can enhance the persistence and abundance of metapopulations comprised of coupled sinks

In spatially heterogeneous landscapes, some habitats may be persistent sources, providing immigrants to sustain populations in unfavorable sink habitats (where extinction is inevitable without immigration). Recent theoretical and empirical studies of source-sink systems demonstrate that temporally variable local growth rates in sinks can substantially increase average abundance of a persisting population, provided that the variation is positively autocorrelated--in effect, temporal variation inflates average abundance. Here we extend these results to a metapopulation in which all habitat patches are sinks. Using numerical studies of a population with discrete generations (buttressed by analytic results), we show that temporal variation and moderate dispersal can jointly permit indefinite persistence of the metapopulation and that positive autocorrelation both lowers the magnitude of variation required for persistence and increases the average abundance of persisting metapopulations. These effects are weakened--but not destroyed--if variation in local growth rates is spatially synchronized and dispersal is localized. We show that the inflationary effect is robust to a number of extensions of the basic model, including demographic stochasticity and density dependence. Because ecological and environmental processes contributing to temporally variable growth rates in natural populations are typically autocorrelated, these observations may have important implications for species persistence.     

Evaluating pollen flow indicators for an insect-pollinated plant species

For insect-pollinated plant species, reproductive success and genetic exchange via the transfer of pollen between flowers depends (i.a.) on the efficiency, abundance and behaviour of floral visitors. These in turn are expected to respond to plant population size and flower density. High floral densities for example usually attract large numbers of pollinators that visit more flowers per plant or patch, which increases pollen deposition at short distances. Thus, population characteristics might serve as indicators for pollen dispersal patterns and help to identify suitable habitat size and quality for conservation measures. To test this hypothesis, we observed floral visitors of a generalist, entomophilous species, Comarum palustre, and compared their abundance and visitation rates in populations of different sizes and flower densities. At the same time, we mimicked pollen flow using fluorescent dye. In the large and dense populations, pollinator abundance and visitation rates were high and dye was dispersed to the edges of the populations (up to 200 m). In the medium-sized population with high flower density, insect abundance and visitation rates were unexpectedly low and dye dispersal declined very quickly. On the contrary, in the smallest population with scattered flowers, especially bumble bee abundance was similar to the large populations and dye dispersal mirrored this high bumble bee activity. Thus, our results indicate that in smaller habitat fragments, the mere size of a population might be insufficient to suggest pollen flow for a plant species. Instead, the abundance of its major pollinators should be considered.     

Potential metapopulation structure and the effects of habitat quality on population size of the endangered False Ringlet butterfly

The False Ringlet (Coenonympha oedippus) is a European butterfly species, endangered due to the severe loss and fragmentation of its habitat. In Hungary, two remaining populations of the butterfly occur in lowland Purple Moorgrass meadows. We studied a metapopulation occupying twelve habitat patches in Central Hungary. Our aim was to reveal what measures of habitat quality affect population size and density of this metapopulation, estimate dispersal parameters and describe phenology of subpopulations. Local population sizes and dispersal parameters were estimated from an extensive mark–release–recapture dataset, while habitat quality was characterized by groundwater level, cover of grass tussocks, bush cover, height of vegetation and grass litter at each habitat patch. The estimated size of the metapopulation was more than 3,000 individuals. We estimated a low dispersal capacity, especially for females, indicating a very low probability of (re)colonization. Butterfly abundance and density in local populations increased with higher grass litter, lower groundwater level and larger area covered by tussocks. We suppose that these environmental factors affect butterfly abundance by determining the microclimatic conditions for both larvae and adult butterflies. Our results suggest that the long-term preservation of the studied metapopulation needs the maintenance of high quality habitat patches by appropriate mowing regime and water regulation. Management also should facilitate dispersal to strengthen metapopulation structure with creating stepping-stones or gradually increase habitat quality in present matrix.     

Observational evidence that maladaptive gene flow reduces patch occupancy in a wild insect metapopulation

Theory predicts that dispersal throughout metapopulations has a variety of consequences for the abundance and distribution of species. Immigration is predicted to increase abundance and habitat patch occupancy, but gene flow can have both positive and negative demographic consequences. Here, we address the eco‐evolutionary effects of dispersal in a wild metapopulation of the stick insect Timema cristinae, which exhibits variable degrees of local adaptation throughout a heterogeneous habitat patch network of two host‐plant species. To disentangle the ecological and evolutionary contributions of dispersal to habitat patch occupancy and abundance, we contrasted the effects of connectivity to populations inhabiting conspecific host plants and those inhabiting the alternate host plant. Both types of connectivity should increase patch occupancy and abundance through increased immigration and sharing of beneficial alleles through gene flow. However, connectivity to populations inhabiting the alternate host‐plant species may uniquely cause maladaptive gene flow that counters the positive demographic effects of immigration. Supporting these predictions, we find the relationship between patch occupancy and alternate‐host connectivity to be significantly smaller in slope than the relationship between patch occupancy and conspecific‐host connectivity. Our findings illustrate the ecological and evolutionary roles of dispersal in driving the distribution and abundance of species.     

Abundance�Coccupancy relationships in metapopulations: examples of rock pool Daphnia

Intraspecific positive relationships between abundance and occupancy are observed for many species, suggesting that the same processes drive local and regional species dynamics. Two main groups of mechanisms explain this relationship: spatiotemporal variation in local population growth rates due to variation in habitat quality, or dispersal effects that increase occupancy of a species when locally abundant. Several studies show that spatiotemporal variation in population growth rates causes positive abundance?Coccupancy relationships, but few have shown dispersal effects. It is believed that such effects should be more evident for species whose dispersal is limited, e.g. metapopulations, but those studies are limited. This study investigates abundance?Coccupancy relationships in three Daphnia metapopulations in rock pools and the degree to which dispersal or habitat quality affect their local abundances and occurrence. Daphnia longispina and Daphnia magna showed positive abundance?Coccupancy relationships, but not Daphnia pulex. No single ecological factor could explain the abundance?Coccupancy relationships of any given species. Instead, dispersal processes and rock pool quality (mainly salinity and depth) seem to act together to shape the abundance?Coccupancy relationships. Such a conclusion is also supported by an immigration experiment in natural rock pools. This study suggests that although positive abundance?Coccupancy relationships may be commonly found for metapopulations, both dispersal processes and variation in habitat quality can be factors determining the abundance?Coccupancy relationship of metapopulations experiencing habitat heterogeneity.     

Integrating distance sampling survey data with population indices to separate trends in abundance and temporary immigration

Managers rely on accurate estimators of wildlife abundance and trends for management decisions. Despite the focus of contemporary wildlife science on developing methods to improve inference from wildlife surveys, legacy datasets often rely on index counts that lack information about the detection process. Data integration can be a useful tool for combining index counts with data collected under more rigorous designs (i.e., designs that account for the detection process), but care is required when datasets represent different population processes or are mismatched in space and time. This can be particularly problematic in cases where animals aggregate in response to a spatially or temporally limited resource because individuals may temporarily immigrate from outside the study area and be included in the abundance index. Abundance indices based on brown bear (Ursus arctos) feeding aggregations within coastal meadows in early summer in Lake Clark National Park and Preserve, Alaska, USA, are one such example. These indices reflect the target population (brown bears residing within the park) and temporary immigrants (i.e., bears drawn from outside the park boundary). To properly account for the effects of temporary immigration, we integrated the index data with abundance data collected via park-wide distance sampling surveys, the latter of which properly addressed the detection process. By assuming that the distance data provide inference on abundance and the index counts represent some combination of abundance and temporary immigration processes, we were able to decompose the relative contribution of each to overall trend. We estimated that the density of brown bears within our study area was 38–54 adults/1,000 km² during 2003–2019 and that abundance increased at a rate of approximately 1.4%/year. The contribution of temporary immigrants to overall trend in the index was low, so we created 3 hypothetical scenarios to more fully demonstrate how the integrated approach could be useful in situations where the composite trend in meadow counts may obscure trends in abundance (e.g., opposing trends in abundance and temporary immigration). Our work represents a conceptual advance supporting the integration of legacy index data with more rigorous data streams and is broadly applicable in cases where trends in index values may represent a mixture of population processes.     

Relationships among nectar‐dwelling yeasts,flowers and ants: patterns and incidence on nectar traits

Nectar‐dwelling yeasts are emerging as widely distributed organisms playing a potentially significant and barely unexplored ecological role in plant pollinator mutualisms. Previous efforts at understanding nectar–pollinator–yeast interactions have focused on bee‐pollinated plants, while the importance of nectarivorous ants as vectors for yeast dispersal remains unexplored so far. Here we assess the abundance and composition of the nectar fungal microbiota of the ant‐pollinated plant Cytinus hypocistis, study whether yeast transmission is coupled with ant visitation, and discern whether ant‐ transported yeasts promote changes in nectar characteristics. Our results show that a high percentage of flowers (77%) and plants (94%) contained yeasts, with yeast cell density in nectar reaching up to 6.2 × 10⁴ cells mm^?3, being the highest densities associated with the presence of the nectar‐specialist yeast Metschnikowia reukaufii. The establishment of fungal microbiota in nectar required flower visitation by ants, with 70% of yeast species transported by them being also detected in nectar. Ant‐vectored yeasts diminished the nutritional quality of nectar, with flowers exposed to pollinators and yeasts containing significantly lower nectar sugar concentration than virgin flowers (13.4% and 22.8%, respectively). Nectar of flowers that harbored M. reukaufii showed the lowest quality, with nectar concentration declining significantly with increasing yeast density. Additionally, yeasts modified patterns of interpopulation variation in nectar traits, homo genizing differences between populations in some nectar attributes. We show for the first time that the outcome of the tripartite pollinator–flower–yeast interaction is highly dependent on the identity and inherent properties of the participants, even to the extent of influencing the species composition of this ternary system, and can be mediated by ecological characteristics of plant populations. Through their influence on plant functional traits, yeasts have the potential to alter nectar consumption, pollinator foraging behavior and ultimately plant reproduction.     

Metapopulation structure and habitat quality in modelling dispersal in the butterfly Iolana iolas

Quantifying dispersal is fundamental to understanding the effects of fragmentation on populations. Although it has been shown that patch and matrix quality can affect dispersal patterns, standard metapopulation models are usually based on the two basic variables, patch area and connectivity. In 2004 we studied migration patterns among 18 habitat patches in central Spain for the butterfly Iolana iolas, using mark–release–recapture methods. We applied the virtual migration (VM) model and estimated the parameters of emigration, immigration and mortality separately for males and females. During parameter estimation and model simulations, we used original and modified patch areas accounting for habitat quality with three different indices. Two indices were based on adult and larval resources (flowers and fruits) and the other one on butterfly density. Based on unmodified areas, our results showed that both sexes were markedly different in their movements and mortality rates. Females emigrated more frequently from patches, but males that emigrated were estimated to move longer daily dispersal distances and suffer higher mortality than females during migration. Males were more likely to emigrate from small than from large patches, but patch area had no significant effect on female emigration. The effects of area on immigration rate and the within-patch mortality were similar in both sexes. Based on modified areas, the estimated parameter values and the model simulation results were similar to those estimated using the unmodified patch areas. One possible reason for the failure to significantly improve the parameter estimates of the VM model is the fact that resource quantity and butterfly population sizes were strongly correlated with patch area. Our results suggest that the standard VM modelling approach, based on patch area and connectivity, can provide a realistic picture of the movement patterns of I. iolas .     

Interacting maternal and spatial cues influence natal – dispersal out of social groups

Proximate cues for animal dispersal are complex and varied. Multiple cues may provide information about different aspects of habitat quality, and these aspects may interact with each other, as well as with population density in different ways. We examined how individuals incorporate multiple cues in their decisions to emigrate and immigrate in the colonial orb‐weaving spider, Cyrtophora citricola. We manipulated maternal feeding as a cue for prey abundance and measured the size of the maternal web, which provides a limited space for philopatric offspring and a second potential dispersal cue. In addition, we recorded all immigration events to determine dispersal distances and the cues juveniles may use in settlement. Dispersal increased when mothers were poorly fed, web sizes were small and clutch sizes were large. In addition to these overall effects, maternal feeding also interacted with web size, indicating that offspring from well‐fed mothers were more tolerant of high sibling densities. We also detected a threshold for the effect of clutch size on dispersal for the first egg sac: below 20 offspring, there was no effect of clutch size, but dispersal increased with clutch size for larger clutches. Dispersal distances were often short, and immigrants preferred sheltered trees and those occupied by adult females. Dispersal not only depended on multiple cues, but these cues interacted, and the importance of web size suggested that saturation of the natal web might force dispersal, at least for spiders with poorly‐fed mothers. How one aspect of habitat quality influences dispersal can therefore depend on the state of other aspects of habitat quality. In particular, some natal resources, such as a nest or territory, may become saturated and limit group size, but this limit will also depend on other factors, such as prey availability.     

Evolutionary consequences of asymmetric dispersal rates

We study the consequences of asymmetric dispersal rates (e.g., due to wind or current) for adaptive evolution in a system of two habitat patches. Asymmetric dispersal rates can lead to overcrowding of the "downstream" habitat, resulting in a source-sink population structure in the absence of intrinsic quality differences between habitats or can even cause an intrinsically better habitat to function as a sink. Source-sink population structure due to asymmetric dispersal rates has similar consequences for adaptive evolution as a source-sink structure due to habitat quality differences: natural selection tends to be biased toward the source habitat. We demonstrate this for two models of adaptive evolution: invasion of a rare allele that improves fitness in one habitat but reduces it in the other and antagonistic selection on a quantitative trait determined by five additive loci. If a habitat can sustain a population without immigration, the conditions for adaptation to that habitat are most favorable if there is little or no immigration from the other habitat; the influence of emigration depends on the magnitude of the allelic effects involved and other parameters. If, however, the population is initially unable to persist in a given habitat without immigration, our model predicts that the population will be most likely to adapt to that habitat if the dispersal rates in both directions are high. Our results highlight the general message that the effect of gene flow upon local adaptation should depend profoundly on the demographic context of selection.     

Linking dispersal, immigration and scale in the neutral theory of biodiversity

In the classic spatially implicit formulation of Hubbell's neutral theory of biodiversity a local community receives immigrants from a metacommunity operating on a relatively slow timescale, and dispersal into the local community is governed by an immigration parameter m . A current problem with neutral theory is that m lacks a clear biological interpretation. Here, we derive analytical expressions that relate the immigration parameter m to the geometry of the plot defining the local community and the parameters of a dispersal kernel. Our results facilitate more rigorous and extensive tests of the neutral theory: we conduct a test of neutral theory by comparing estimates of m derived from fits to empirical species abundance distributions to those derived from dispersal kernels and find acceptable correspondence; and we generate a new prediction of neutral theory by investigating how the shapes of species abundance distributions change theoretically as the spatial scale of observation changes. We also discuss how our main analytical results can be used to assess the error in the mean-field approximations associated with spatially implicit formulations of neutral theory.     

High dispersal ability and low genetic differentiation in the widespread butterfly species <Emphasis Type="Italic">Melanargia galathea</Emphasis>

Agricultural intensification caused a fragmentation of flower-rich extensively used meadows which resulted in the reduction of the abundance of species of these habitats. The abundance pattern and the dispersal behaviour of species influence the connectivity of local populations. In this context exchange rates can be directly measured by mark release recapture (MRR) studies and indirectly by genetic analysis. Both approaches are used in our study in a comparative way. As a test species to investigate the influence of habitat interconnectivity on a local and regional scale, we selected the butterfly Melanargia galathea (Lepidoptera, Nymphalidae), a widely distributed species, which is common on flower-rich meadows in our study area in western Germany. We marked 3,175 individuals of four neighbouring sites in a mark-release-recapture study and analysed 18 allozyme loci for 644 individuals sampled over 17 sites. Only 3.3% of the total genetic variance was found among samples, thus supporting the detected between-patch movements. Both approaches revealed a high exchange rate among local populations. Moderate between- and high within-patch movements were recorded by MRR analysis, dependent on the geographical distance. The two analytical tools showed high estimated effective population sizes for all populations. In the light of conservation biology, the combination of the MRR and allozyme data support the assumption that high dispersal ability and habitat interconnectivity countervail genetic differentiation and enable the maintenance of a high level of genetic differentiation.     

Relative contributions of local and regional factors to species richness and total density of butterflies and moths in semi-natural grasslands

Metapopulation theory predicts that species richness and total population density of habitat specialists increase with increasing area and regional connectivity of the habitat. To test these predictions, we examined the relative contributions of habitat patch area, connectivity of the regional habitat network and local habitat quality to species richness and total density of butterflies and day-active moths inhabiting semi-natural grasslands. We studied butterflies and moths in 48 replicate landscapes situated in southwest Finland, including a focal patch and the surrounding network of other semi-natural grasslands within a radius of 1.5 km from the focal patch. By applying the method of hierarchical partitioning, which can distinguish between independent and joint contributions of individual explanatory variables, we observed that variables of the local habitat quality (e.g. mean vegetation height and nectar plant abundance) generally showed the highest independent effect on species richness and total density of butterflies and moths. Habitat area did not show a significant independent contribution to species richness and total density of butterflies and moths. The effect of habitat connectivity was observed only for total density of the declining butterflies and moths. These observations indicate that the local habitat quality is of foremost importance in explaining variation in species richness and total density of butterflies and moths. In addition, declining butterflies and moths have larger populations in well-connected networks of semi-natural grasslands. Our results suggest that, while it is crucial to maintain high-quality habitats by management, with limited resources it would be appropriate to concentrate grassland management and restoration to areas with well-connected grassland networks in which the declining species currently have their strongest populations. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

What can genetics tell us about population connectivity?

Genetic data are often used to assess ‘population connectivity’ because it is difficult to measure dispersal directly at large spatial scales. Genetic connectivity, however, depends primarily on the absolute number of dispersers among populations, whereas demographic connectivity depends on the relative contributions to population growth rates of dispersal vs. local recruitment (i.e. survival and reproduction of residents). Although many questions are best answered with data on genetic connectivity, genetic data alone provide little information on demographic connectivity. The importance of demographic connectivity is clear when the elimination of immigration results in a shift from stable or positive population growth to negative population growth. Otherwise, the amount of dispersal required for demographic connectivity depends on the context (e.g. conservation or harvest management), and even high dispersal rates may not indicate demographic interdependence. Therefore, it is risky to infer the importance of demographic connectivity without information on local demographic rates and how those rates vary over time. Genetic methods can provide insight on demographic connectivity when combined with these local demographic rates, data on movement behaviour, or estimates of reproductive success of immigrants and residents. We also consider the strengths and limitations of genetic measures of connectivity and discuss three concepts of genetic connectivity that depend upon the evolutionary criteria of interest: inbreeding connectivity, drift connectivity, and adaptive connectivity. To conclude, we describe alternative approaches for assessing population connectivity, highlighting the value of combining genetic data with capture‐mark‐recapture methods or other direct measures of movement to elucidate the complex role of dispersal in natural populations.     

Differential effects of spatial network structure and scale on population size and genetic diversity of the ninespine stickleback in a remnant wetland system

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Butterfly Density and Behaviour in Uncut Hay Meadow Strips: Behavioural Ecological Consequences of an Agri-Environmental Scheme

Sparing zones from mowing has been proposed, and applied, to improve local conditions for survival and reproduction of insects in hay meadows. However, little is known about the efficiency of refuge zones and the consequences for local populations. We studied population densities of butterflies before and after mowing in the refuge zone of 15 meadows in 2009 and 2011. We also studied the behaviour of the meadow brown (Maniola jurtina) comparing nectar use, interactions and flights in the refuge zone before and after mowing. Densities of grassland butterflies in this zone doubled on average after mowing. The density of females of M. jurtina increased on average fourfold, while males showed a more modest increase. In line with the idea of increased scramble competition in the refuge zone after mowing, M. jurtina increased the time spent on nectar feeding, the preferred nectar source was visited more frequently, and females made more use of non-preferred nectar sources. Maniola jurtina did not interact more with conspecifics after mowing, but interactions lasted longer. Flight tracks did not change in linearity, but were faster and shorter after mowing. After mowing, only a part of the local grassland butterflies moved to the uncut refuge zone. The resulting concentration effect alters the time allocated to different activities, nectar use and movements. These aspects have been largely ignored for agri-environmental schemes and grassland management in nature reserves and raise questions about optimal quantities and quality of uncut refuge sites for efficient conservation of grassland arthropods in agricultural landscapes.     

Do species population parameters and landscape characteristics affect the relationship between local population abundance and surrounding habitat amount?

In landscape ecology, correlational approaches are typically used to analyse links between local population abundance, and the surrounding habitat amount to estimate biologically-relevant landscape size (extent) for managing endangered or pest populations. The direction, strength, and spatial extent of the correlations are then sometimes interpreted in terms of species population parameters. Here we simulated the population dynamics of generalized species across spatially explicit landscapes that included two distinct habitat types. We investigated how characteristics of a landscape (structure), including the variation in habitat quality and spatial aggregation of the habitat, and the precise population-dynamic properties of the simulated species (dispersal and growth rates) affect the correlation between population abundance and amount of surrounding favourable habitat in the landscape. To evaluate these spatial extents of correlation, proportions of favourable habitat were calculated within several circles of increasing diameter centred on sampling patches of favourable habitat where population abundance was recorded.We found that the value of the correlation coefficients between population abundance and amount of surrounding favourable habitat depended on both population dynamic parameters and landscape characteristics. Coefficients of correlation increased with the variation in habitat quality and the aggregation of favourable habitat in the landscape, but decreased with the dispersal distance. The distance at which the correlation was maximized was sensitive to an interaction between the level of aggregation of the habitat and the dispersal distance; whereas the greatest distance at which a significant correlation occurred was more sensitive to the variation in habitat quality. Our results corroborate the view that correlational analyses do provide information on the local population dynamics of a species in a given habitat type and on its dispersal rate parameters. However, even in simplified, model frameworks, direct relationships are often difficult to disentangle and global landscape characteristics should be reported in any studies intended to derive population-dynamic parameters from correlations. Where possible, replicated landscapes should be examined in order to control for the interaction between population dynamics and landscape structure. Finally, we recommend using species-specific, population-dynamic modelling in order to interpret correctly the observed patterns of correlation in the landscape.     

Dispersal and mortality of prairie voles (Microtus ochrogaster) in fragmented landscapes: a field experiment

We conducted a field experiment that manipulated landscapes by mowing so that the amount of unfavorable habitat (low cover) for prairie voles ( Microtus ochrogaster ) increased while the number and size of favorable patches (high cover) remained constant. Distance between favorable patches increased as the amount of unfavorable habitat increased, so we could test two current hypotheses concerning the effect of habitat fragmentation on local populations: 1) increased distance between favorable habitat patches reduces successful per capita dispersal (emigration and immigration) because dispersers suffer greater exposure to predators (the predation hypothesis); and 2) per capita dispersal is inversely density dependent in voles because increased aggression at higher density inhibits movements (the social fence hypothesis). As predicted by the predation hypothesis, increased distance between favorable habitat patches led to decreased successful dispersal among patches and increased per capita mortality, particularly among subadult and adult males (the categories of voles most likely to emigrate). As predicted by the social fence hypothesis, dispersal was inversely density dependent, and dispersing voles displayed a greater frequency of wounding (an indication of increased aggressive interactions) than did residents. The amount of wounding in general did not increase with density, however, and, as distance between patches increased to 60 m, successful dispersal became rare and erratic. Nevertheless, our overall results supported current hypotheses regarding the effects of increased habitat fragmentation on patterns of dispersal and mortality. Examining the impact of these effects on local population dynamics within different landscapes will require longer periods of observation.