Detecting Lévy walks without turn designation

Many organisms have been reported to have movement patterns that are well approximated as Lévy walks. This is typically because distributions of straight line distances between consecutive significant turns in movement paths have heavy power law tails. This diagnostic tool has been called into question because there is currently no standard, unambiguous way to identify significant turns. Even if such a way could be found, statistical analyses based on significant turns cannot account for actual movements made between turns and as a consequence cannot distinguish between true Lévy walks and other fractal random walks such as Lévy modulated correlated random walks where organisms randomly meander rather than move in straight lines between consecutive reorientation events. Here, I show that structure functions (i.e. moments of net displacements made across fixed time intervals) can distinguish between different kinds of Lévy walks and between Lévy walks and random walks with a few scales such as composite correlated random walks and correlated random walks. Distinguishing between these processes will lead to a better understanding of how and why animals perform Lévy walks and help bridge the apparent divide between correlated random walks and Lévy walks. Structure functions do not require turn identification and instead take account of entire movement paths. Using this diagnostic tool, I bolster previous claims that honeybees use a movement strategy that can be approximated by Lévy walks when searching for their hive. I also show how structure functions can be used to establish the extent of self-similar behaviour in meandering Lévy walks.     

Scale-invariant movements of fishermen: the same foraging strategy as natural predators.

We analyzed the movement of fishing vessels during fishing trips in order to understand how fishermen behave in space while searching for fish. For that purpose we used hourly geo-referenced positions of vessels, provided by a satellite vessel monitoring system, for the entire industrial fleet (809 vessels) of the world's largest single species fishery (Peruvian anchovy, Engraulis ringens) from December 1999 to March 2003. Observed trajectories of fishing vessels are well modeled by Lévy random walks, suggesting that fishermen use a stochastic search strategy which conforms to the same search statistics as non-human predators. We show that human skills (technology, communication, or others) do not result in the fishermen's spatial behavior being fundamentally different from that of animal predators. With respect to probability of prey encounter, our results suggest that fishermen, on average, evolved an optimal movement pattern (mu = 2.00) among the family of Lévy random walks. This Lagrangian approach opens several perspectives in terms of operational management of the pelagic fish stock.     

Are Lévy flight patterns derived from the Weber–Fechner law in distance estimation?

Honeybees (Apis mellifera) are regularly faced with the task of navigating back to their hives from remote food sources. They have evolved several methods to do this, including compass-directed “vector” flights and the use of landmarks. If these hive-centered mechanisms are disrupted, bees revert to searching for the hive, using an optimal Lévy flight searching strategy. The same strategy is adopted when a food source at a known location ceases to be available. Here, we show that the programming for this Lévy strategy does not need to be very sophisticated or clever on the bee’s part, as Lévy flight patterns can be derived from the Weber–Fechner law in a bee’s odometer. Odometry errors of a different kind occur in desert ants (Cataglyphis spp., Melophorus bagoti). The searching behaviors of these ants are very similar in overall structure to that of honeybees but do not display any Lévy flight characteristics. We suggest that errors in the estimation of distance can be implicitly involved in shaping the structure of systematic search behavior and should not be regarded as merely deficiencies in the odometer.     

Lévy flight patterns are predicted to be an emergent property of a bumblebees’ foraging strategy

Bumblebees forage uninterrupted for long periods of time because they are not distracted by sex or territorial defense and have few predators. This has led to a long running debate about whether bumblebees forage optimally. This debate has been enriched by the possibility that bumblebees foraging within clover patches have flight patterns that can be approximated by Lévy flights. Such flight patterns optimise the success of random searches. Bumblebees foraging within a flower patch tend to approach the nearest flower but then often depart without landing or probing it if it has been visited previously; unvisited flowers are not rejected in this manner. Here, this foraging behaviour has been replicated in numerical simulations. Lévy flight patterns are found to be an inconsequential emergent property of a bumblebees’ foraging behaviour. Lévy flights are predicted to emerge when bees reject at least 99% of previously visited flowers. A foraging bumblebee can certainly empty a clover flower head of nectar in one visit, but lower rates of rejection are observed for many other flowers. These findings suggest that Lévy flight patterns in foraging bumblebees are rare and specific to a few flower species and that if they exist, then they are not part of an innate, evolved optimal searching strategy.     

Spatial synchrony in coral reef fish populations and the influence of climate

We investigated spatial patterns of synchrony among coral reef fish populations and environmental variables over an eight-year period on the Great Barrier Reef, Australia. Our aims were to determine the spatial scale of intra- and interspecific synchrony of fluctuations in abundance of nine damselfish species (genus Pomacentrus) and assess whether environmental factors could have influenced population synchrony. All species showed intraspecific synchrony among populations on reefs separated by < or =100 km, and interspecific synchrony was also common at this scale. At greater spatial scales, only four species showed intraspecific synchrony, over distances ranging from 100-300 km to 500-800 km, and no cases of interspecific synchrony were recorded. The two mechanisms most likely to cause population synchrony are dispersal and environmental forcing through regionally correlated climate (the Moran effect). Dispersal may have influenced population synchrony over distances up to 100 km as this is the expected spatial range for ecologically significant reef fish dispersal. Environmental factors are also likely to have synchronized population fluctuations via the Moran effect for three reasons: (1) dispersal could not have caused interspecific synchrony that was common over distances < or =100 km because dispersal cannot link populations of different species, (2) variations in both sea surface temperature and wind speed were synchronized over greater spatial scales (>800 km) than fluctuations in damselfish abundance (< or =800 km) and were correlated with an index of global climate variability, the El Ni?o-Southern Oscillation (ENSO), and (3) synchronous population fluctuations of most damselfish species were correlated with ENSO; large population increases often followed ENSO events. We recorded regional variations in the strength of population synchrony that we suspect are due to spatial differences in geophysical, oceanographic, and population characteristics, which act to dilute or enhance the effects of synchronizing mechanisms. We conclude that synchrony is common among Pomacentrus populations separated by tens of kilometers but less prevalent at greater spatial scales, and that environmental variation linked to global climate is likely to be a driving force behind damselfish population synchrony at all spatial scales on the Great Barrier Reef.     

Breeding success of tree swallows along a gradient of agricultural intensification

The intensification of agricultural practices has been identified as the main cause of population decline in farmland birds since the 1960s in both Europe and North America. Although the links between species richness or abundance and various components of agricultural intensification are well established, the mechanisms underlying these trends have rarely been addressed along a gradient of intensification or have been quantified at only one spatial scale. Here we quantified the influence of landscape structure on the nest box occupancy and breeding success of Tree Swallows (Tachycineta bicolor) at seven spatial scales (1, 2, 3, 4, 5, 10, and 20 km radii) over a 10 200-km2 gradient of agricultural intensification in southern Québec, Canada. A network of 400 nest boxes distributed among 40 farms was visited every two days over three breeding seasons, 2004-2006. Nest box occupancy decreased with the proportion of intensive cultures (maize, cereals, and soybeans) in the landscape, especially when manure heaps and tanks were abundant, and was also determined by local variables (i.e., nest box clearance, interspecific competition) and by previous-year fledging success. Clutch size decreased as the breeding season progressed and with the proportion of intensive cultures in the landscape, with no consistent variation across spatial scales. Hatching success was not related to any landscape variables but increased with clutch size. Both the number of fledglings and fledging probability increased with the proportion of extensive cultures (hayfields, pastures, and fallows). These effects increased with spatial scale and reached a plateau at the 5 km radius: the maximum distance from the nest reached by foraging Tree Swallows. Our results can likely be attributed to lower food availability in intensive cultures compared to extensive ones. This study suggests that several components of breeding that impact on population structure and dynamics of insectivorous birds will be negatively affected by agricultural intensification.     

Spatial scaling of avian population dynamics: population abundance, growth rate, and variability

Synchrony in population fluctuations has been identified as an important component of population dynamics. In a previous study, we determined that local-scale (<15-km) spatial synchrony of bird populations in New England was correlated with synchronous fluctuations in lepidopteran larvae abundance and with the North Atlantic Oscillation. Here we address five questions that extend the scope of our earlier study using North American Breeding Bird Survey data. First, do bird populations in eastern North America exhibit spatial synchrony in abundances at scales beyond those we have documented previously? Second, does spatial synchrony depend on what population metric is analyzed (e.g., abundance, growth rate, or variability)? Third, is there geographic concordance in where species exhibit synchrony? Fourth, for those species that exhibit significant geographic concordance, are there landscape and habitat variables that contribute to the observed patterns? Fifth, is spatial synchrony affected by a species' life history traits? Significant spatial synchrony was common and its magnitude was dependent on the population metric analyzed. Twenty-four of 29 species examined exhibited significant synchrony in population abundance: mean local autocorrelation (rho)= 0.15; mean spatial extent (mean distance where rho=0) = 420.7 km. Five of the 29 species exhibited significant synchrony in annual population growth rate (mean local autocorrelation = 0.06, mean distance = 457.8 km). Ten of the 29 species exhibited significant synchrony in population abundance variability (mean local autocorrelation = 0.49, mean distance = 413.8 km). Analyses of landscape structure indicated that habitat variables were infrequent contributors to spatial synchrony. Likewise, we detected no effects of life history traits on synchrony in population abundance or growth rate. However, short-distance migrants exhibited more spatially extensive synchrony in population variability than either year-round residents or long-distance migrants. The dissimilarity of the spatial extent of synchrony across species suggests that most populations are not regulated at similar spatial scales. The spatial scale of the population synchrony patterns we describe is likely larger than the actual scale of population regulation, and in turn, the scale of population regulation is undoubtedly larger than the scale of individual ecological requirements.     

Spatial changes in the distributions of deep-sea “Cerviniidae” (Harpacticoida, Copepoda) and their associations with environmental factors in the bathyal zone around Sagami Bay, Japan

To estimate species turnover rates on scales of several tens of km in deep-sea benthic animals, we analyzed spatial and inter-annual changes in species diversity and composition of cerviniids, a typical group of deep-sea harpacticoids, at stations in and around Sagami Bay, central Japan. Associations with environmental factors were also investigated. Generally, bathymetrical patterns in diversity of benthos are unimodal and peak at depths of 2,000–3,000 m. In Sagami Bay, cerviniid diversity did not follow this trend; both species richness and evenness were negatively correlated with water depth. Multivariate analyses [detrended correspondence analysis (DCA) and non-metric multi-dimensional scaling] suggested that temporal changes in species composition of cerviniids are smaller than spatial changes that occur on horizontal scales of several tens of km. Community structure does not change completely on these scales in the bathyal zone around Sagami Bay. DCA also showed that bathymetrical changes in species composition can be regulated by certain factors associated with water depth.     

Displaced honey bees perform optimal scale-free search flights

Honey bees (Apis mellifera) are regularly faced with the task of navigating back to their hives from remote food sources. They have evolved several methods to do this, including compass-directed "vector" flights and the use of landmarks. If these hive-centered mechanisms are disrupted, bees revert to searching for the hive, but the nature and efficiency of their searching strategy have hitherto been unknown. We used harmonic radar to record the flight paths of honey bees that were searching for their hives. Our subsequent analysis of these paths revealed that they can be represented by a series of straight line segments that have a scale-free, Lévy distribution with an inverse-square-law tail. We show that these results, combined with the "no preferred direction" characteristic of the segments, demonstrate that the bees were flying an optimal search pattern. Lévy movements have already been identified in a number of other animals. Our results are the best reported example where the movements are mostly attributable to the adoption of an optimal, scale-free searching strategy.     

Scale-Dependent Effects of Habitat Disturbance on Species Richness in Tropical Forests

Abstract: Despite growing concern, no consensus has emerged over the effects of habitat modification on species diversity in tropical forests. Even for comparatively well-studied taxa such as Lepidoptera, disturbance has been reported to increase and decrease diversity with approximately equal frequency. Species diversity within landscapes depends on the spatial scale at which communities are sampled, and the effects of disturbance in tropical forests have been studied at a wide range of spatial scales. Yet the question of how disturbance affects diversity at different spatial scales has not been addressed. We reanalyzed data from previous studies to examine the relationship between spatial scale and effects of disturbance on tropical-forest Lepidoptera. Disturbance had opposite effects on diversity at large and small scales: as scale decreased, the probability of a positive effect of disturbance on diversity increased. We also explicitly examined the relationship between spatial scale and the diversity of butterflies in selectively logged and unlogged forest in Maluku Province, Indonesia. Species richness increased with spatial scale in both logged and unlogged forest, but at a significantly faster rate in unlogged forest, whereas species evenness increased with scale in unlogged forest but did not increase with scale in logged forest. These data indicate that the effects of habitat modification on species diversity are heavily scale-dependent. As a result, recorded effects of disturbance were strongly influenced by the spatial scale at which species assemblages were sampled. Future studies need to account for this by explicitly examining the effects of disturbance at a number of different spatial scales. A further problem arises because the relationship between scale and diversity is likely to differ among taxa in relation to mobility. This may explain to some extent why the measured effects of disturbance have differed between relatively mobile and immobile taxa.     

城乡建设用地景观格局多尺度差异

以郑州市域内1989、2000和2009年的典型城市与农村样地（25 km×25 km）为例,应用空间自相关、半方差、以及九项轨迹方差分析方法,对城市与农村建设用地景观多尺度上的空间格局特征与动态差异进行研究,结果表明,（1）城市建设用地景观聚集强度较高,斑块规模较大,而农村建设用地景观在大部分尺度上呈随机或均匀分布,斑块规模较小。（2）随着尺度的增加,城市及农村建设用地景观分布状态都由聚集向均匀过渡,但前者过渡较为平缓,而后者在小尺度范围（0~2km）变化速度较快,在较大尺度（3~16 km）上其分布状态的波动较小。（3）1989-2009年,城市与农村建设用地景观动态特征有较大差异,并表现出尺度依赖性。城市较小尺度上（0~4 km）建设用地景观的分布状态表现出聚集-分散的趋势,在较大尺度上（8~14 km）表现出分散-聚集的变化趋势。农村小尺度上（0~4 km）建设用地景观的分布状态也表现出聚集-分散的趋势,但大尺度（4~16 km）上状态变化很小且没有规律。     

Spatial patterns of spawning in the coral-reef damselfish Dascyllus albisella

Reproductive patterns of the endemic Hawaiian damselfish Dascyllus albisella were examined around the island of Oahu for 5 m in 1992. Daily variation in spawning was studied at four spatial scales: individual territories; reef transects; whole patch reefs; and locations separated by>20 km. A 6 d, non-lunar nesting cycle was found for individual males, and a synchronized, 6 d, nonhinar spawning cycle was found within reef transects, whole patch reefs, and locations. Spawning synchronicity was maintained among reefs within a location. However, spawning among locations was found to be asynchronous, limiting the spatial scale of reproductive synchrony to>1 km but<20 km. Inherent benefits of synchronous reproduction are discussed for D. albisella, but only the localized swamping of planktonic predators may be an inherent benefit of synchronous spawning for this species. Alternatively, synchronous spawning among populations may exist from adults tracking local, favorable environmental conditions rather than having evolved towards an inherent benefit of synchronization.     

Honeybees use a Lévy flight search strategy and odour-mediated anemotaxis to relocate food sources

The availability of food resources changes over time and space, and foraging animals are constantly faced with choices about how to respond when a resource becomes depleted. We hypothesise that flying insects like bees discover new food sources using an optimal Lévy flight searching strategy and odour-mediated anemotaxis, as well as visual cues. To study these searching patterns, foraging honeybees were trained to a scented feeder which was then removed. Two new unrewarding feeders, or ‘targets’, were then positioned up- and downwind of the original location of the training feeder. The subsequent flight patterns of the bees were recorded over several hundred metres using harmonic radar. We show that the flight patterns constitute an optimal Lévy flight searching strategy for the location of the training feeder, a strategy that is also optimal for the location of alternative food sources when patchily distributed. Scented targets that were positioned upwind of the original training feeder were investigated most with the numbers of investigations declining with increasing distance from the original feeder. Scented targets in downwind locations were rarely investigated and unscented targets were largely ignored, despite having the same visual appearance as the rewarding training feeder.     

Combining fishing and acoustic monitoring data to evaluate the distribution and movements of spotted ratfish <Emphasis Type="Italic">Hydrolagus colliei</Emphasis>

Direct and indirect methods have been used to describe patterns of movement of fishes, but few studies have compared these methods simultaneously. We used 20 years of trawl survey data and 1 year of acoustic telemetry data to evaluate the vertical and horizontal movement patterns of spotted ratfish Hydrolagus colliei in Puget Sound, WA, USA. Densities of large ratfish (≥30 cm) were higher at the deepest depths trawled (70 m) during daylight hours, whereas densities were similar across depth zones (to 10 m) at night. Acoustic tracking of ratfish showed distinct diel patterns of movement and activity level; ratfish moved into shallow, nearshore habitats at night from deeper, offshore habitats during the day and made ~3 times more moves at night than day in shallow habitats. Broader spatial patterns depended on where ratfish were tagged: one tag group remained in one general location with few excursions, whereas a second tag group moved within a 20-km band with some individuals moving >90 km. These data will help inform food web models’ abilities to quantify interspecific interactions between ratfish and other components of their community.     

Scale-dependent changes in the importance of larval supply and habitat to abundance of a reef fish

While there is great interest in the degree to which local interactions "scale-up" to predict regional patterns of abundance, few studies in marine systems have simultaneously examined patterns of abundance at both the large scale (tens of kilometers) typical of larval movement and the small scale (meters) typical of post-settlement interactions. We addressed this gap by monitoring larval supply, adult survivorship, and giant kelp (Macrocystis pyrifera, a primary habitat-forming species) abundance for 13 populations of kelp bass (Paralabrax clathratus) spread over approximately 200 km in the Santa Barbara Channel, California, USA. At the small, within-site scale, both recruitment and adult survivorship of kelp bass were density-dependent and positively related to kelp abundance. At the larger, among-site scale, the spatial pattern of adult kelp bass abundance was predicted well by the pattern of kelp bass larval supply, but there was a consistent negative spatial relationship between kelp abundance and kelp bass larval supply despite the positive effects of kelp on kelp bass at the smaller spatial scale. This large-scale negative relationship was likely a product of a channel-wide spatial mismatch between oceanographic conditions that favor kelp survival and those that concentrate and distribute fish larvae. These results generally support the recruit-adult hypothesis: kelp bass populations are limited by recruitment at low recruit densities but by density-dependent competition for food resources and/or predator refuges at high recruit densities. At the same time, spatial variation in kelp abundance produced substantial spatiotemporal heterogeneity in kelp bass demographics, which argues for a multispecies, metacommunity approach to predicting kelp bass dynamics.     

GPS tracking a marine predator: the effects of precision,resolution and sampling rate on foraging tracks of African Penguins

We used a prototype GPS logger to track the movements of breeding African Penguins (Spheniscus demersus). The loggers also recorded temperature and water depth, which allowed us to reconstruct foraging tracks in three dimensions, although GPS signals are interrupted when the birds dive. Here we report the loggers performance in the field and assess the effects of GPS error, resolution and sampling rate on estimates of foraging track length and speed. There is a trade-off between sampling rate and battery lifespan. We tested loggers at sampling intervals of 1 s, 10 s, 1 min, 2 min and 10 min. Sampling less frequently increases the chance of tracking an entire foraging trip, but it slows uplink times, slightly decreases the accuracy of positional fixes, and significantly reduces the ability to measure fine-scale aspects of foraging behaviour. Compared with radio or satellite tracking, GPS loggers offer unprecedented detail about animal movements. The results of our analysis suggest that techniques that sample relatively infrequently, such as satellite tracking, underestimate actual track lengths by up to 50%. However, caution is needed when interpreting fine-scale sampling for relatively slow-moving organisms. Re-sampling 1-s tracks suggests that c. 35% of apparent movements at this scale are due to measurement error and, more importantly, the limited spatial resolution of GPS (1.85×1.54 m at the study area). We recommend that researchers use a 1-s sampling rate for fine-scale studies, but resample at less frequent intervals to remove spurious noise for slow-moving animals. At current levels of resolution, animals should move at least 4 m per sampling interval. We provide empirical correction factors to compare inferred track length sampled at different rates, but caution that these are idiosyncratic and strongly dependent on the animals behaviour. Overall, GPS loggers offer a significant advance for studies of fine-scale animal movement patterns.Communicated by O. Kinne, Oldendorf/Luhe     

Highly Differentiated Populations of the Narrow Endemic Plant Maguire Primrose (Primula maguirei)

Maguire primrose (Primula maguirei) is a geographically restricted plant species, known only from a 19-km stretch of Logan Canyon in northern Utah (U.S.A.). We examined variation at 13 isozyme loci from 25 individuals of P. maguirei at each of eight sites. At individual loci we detected no statistically significant deviations from Hardy-Weinberg proportions within sites (subpopulations). However, some loci were almost fixed for different alleles at the upper reaches of the species' range relative to populations approximately 10 km away. The total mean gene diversity among loci was 0.22, of which 55% was partitioned within subpopulations, 0.7% among subpopulations within populations (100 m spatial scale), 3% among populations separated by about 1 km, and 41% between an Upper Canyon group of populations and a Lower Canyon group (10-km scale). We detected no gametic disequilibria among loci within subpopulations (and populations). Two hypotheses are proposed to explain the results: (1) past genetic bottlenecks and (2) genetic divergence as a by-product of local adaptations to different habitats. Regardless of the causes of allozymic differentiation, our results suggest that plans for artificial establishment or reestablishment of P. maguirei populations should use source populations within 1 km of the establishment site. This study emphasizes the potential use of data on population genetic structure for managing and monitoring rare species.     

Spectrum of selection: new approaches to detecting the scale-dependent response to habitat

Detecting habitat selection depends on the spatial scale of analysis, but multi-scale studies have been limited by the use of a few, spatially variable, hierarchical levels. We developed spatially explicit approaches to quantify selection along a continuum of scales using spatial (coarse-graining) and geostatistical (variogram) pattern analyses at multiple levels of habitat use (seasonal range, travel routes, feeding areas, and microsites). We illustrate these continuum-based approaches by applying them to winter habitat selection by woodland caribou (Rangifer tarandus caribou) using two key habitat components, Cladina lichens and snow depth. We quantified selection as the reduction in variance in used relative to available sites, thus avoiding reliance on correlations between organism and habitat, for which interpretation can be impeded by cross-scale correlations. By consistently selecting favorable habitat features, caribou experienced reduced variance in these features. The degree to which selection was accounted for by the travel route, feeding area, or microsite levels varied across the scale continuum. Caribou selected for Cladina within a 13-km scale domain and selected shallower snow at all scales. Caribou responded most strongly at the dominant scales of patchiness, implicating habitat heterogeneity as an underlying cause of multi-scale habitat selection. These novel approaches enable a spatial understanding of resource selection behavior.     

Variations in mortality of a coral-reef fish: links with predator abundance

The mortality rates of a pomacentrid Acanthochromis polyacanthus were examined in relation to the abundance of large predatory fish (>200 mm total length, TL) at two spatial scales. Survivorship was negatively related to patterns of predator abundance at a large spatial scale (hundreds of metres) over 3 yr, but not at a small spatial scale (tens of metres) over 2 yr. On the large scale, mortality was consistently greater (14 to 30%) in locations where there were greater numbers of predators, and lower in locations where predators occurred in smaller numbers. Among these locations, spatial differences in rank abundance of surviving juveniles were primarily due to mortality, whereas temporal differences in rank abundance were primarily due to initial juvenile abundance. These data suggest that impacts of large predatory fish were likely to have been greater in space than time and at the large spatial scale than the small spatial scale.     

A methodological framework for integrating computational fluid dynamics and ecological models applied to juvenile freshwater mussel dispersal in the Upper Mississippi River

Interdisciplinary research in hydraulics and ecology for river management and restoration must integrate processes that occur over a wide range of spatial and temporal scales, which presents a challenge to ecohydraulics modelers. Computational fluid dynamics (CFD) models are being more widely used to determine flow fields for ecohydraulics applications. In the Upper Mississippi River (UMR), the mussel dynamics model was developed as a tool for management and conservation of freshwater mussels (Unionidae), which are benthic organisms, imperiled in North America, that are inextricably linked with the hydraulics of river flow. We updated the juvenile dispersal component of the mussel dynamics model by using stochastic Lagrangian particle tracking in a three dimensional flow field output from CFD models of reaches in the UMR. We developed a methodological framework to integrate hydrodynamic data with the mussel dynamics model, and we demonstrate the use of the juvenile dispersal model employed within the methodological framework in two reaches of the UMR. The method was used to test the hypothesis that impoundment affects the relationship of some hydraulic parameters with juvenile settling distribution. Simulation results were consistent with this hypothesis, and the relationships of bed shear stress and Froude number with juvenile settling were altered by impoundment most likely through effects on local hydraulics. The methodological framework is robust, integrates Eulerian and Lagrangian reference frameworks, and incorporates processes over a wide range of temporal and spatial scales, from watershed scale hydrologic processes (decades), to reach scale (km) processes that occur over hours or days, and turbulent processes on spatial scales of meter to millimeter and times scales of seconds. The methods are presently being used to assess the impacts of pre- and early post-settlement processes on mussel distributions, including the effects of bed shear stress, and the sensitivity of the location of the host fish when juveniles excyst, on juvenile settling distribution.