Scaling laws of ambush predator 'waiting' behaviour are tuned to a common ecology

The decisions animals make about how long to wait between activities can determine the success of diverse behaviours such as foraging, group formation or risk avoidance. Remarkably, for diverse animal species, including humans, spontaneous patterns of waiting times show random ‘burstiness’ that appears scale-invariant across a broad set of scales. However, a general theory linking this phenomenon across the animal kingdom currently lacks an ecological basis. Here, we demonstrate from tracking the activities of 15 sympatric predator species (cephalopods, sharks, skates and teleosts) under natural and controlled conditions that bursty waiting times are an intrinsic spontaneous behaviour well approximated by heavy-tailed (power-law) models over data ranges up to four orders of magnitude. Scaling exponents quantifying ratios of frequent short to rare very long waits are species-specific, being determined by traits such as foraging mode (active versus ambush predation), body size and prey preference. A stochastic–deterministic decision model reproduced the empirical waiting time scaling and species-specific exponents, indicating that apparently complex scaling can emerge from simple decisions. Results indicate temporal power-law scaling is a behavioural ‘rule of thumb’ that is tuned to species’ ecological traits, implying a common pattern may have naturally evolved that optimizes move–wait decisions in less predictable natural environments.

Rationalizing spatial exploration patterns of wild animals and humans through a temporal discounting framework

Understanding the exploration patterns of foragers in the wild provides fundamental insight into animal behavior. Recent experimental evidence has demonstrated that path lengths (distances between consecutive turns) taken by foragers are well fitted by a power law distribution. Numerous theoretical contributions have posited that “Lévy random walks”—which can produce power law path length distributions—are optimal for memoryless agents searching a sparse reward landscape. It is unclear, however, whether such a strategy is efficient for cognitively complex agents, from wild animals to humans. Here, we developed a model to explain the emergence of apparent power law path length distributions in animals that can learn about their environments. In our model, the agent’s goal during search is to build an internal model of the distribution of rewards in space that takes into account the cost of time to reach distant locations (i.e., temporally discounting rewards). For an agent with such a goal, we find that an optimal model of exploration in fact produces hyperbolic path lengths, which are well approximated by power laws. We then provide support for our model by showing that humans in a laboratory spatial exploration task search space systematically and modify their search patterns under a cost of time. In addition, we find that path length distributions in a large dataset obtained from free-ranging marine vertebrates are well described by our hyperbolic model. Thus, we provide a general theoretical framework for understanding spatial exploration patterns of cognitively complex foragers.

Rationalizing spatial exploration patterns of wild animals and humans through a temporal discounting framework

Understanding the exploration patterns of foragers in the wild provides fundamental insight into animal behavior. Recent experimental evidence has demonstrated that path lengths (distances between consecutive turns) taken by foragers are well fitted by a power law distribution. Numerous theoretical contributions have posited that “Lévy random walks”—which can produce power law path length distributions—are optimal for memoryless agents searching a sparse reward landscape. It is unclear, however, whether such a strategy is efficient for cognitively complex agents, from wild animals to humans. Here, we developed a model to explain the emergence of apparent power law path length distributions in animals that can learn about their environments. In our model, the agent’s goal during search is to build an internal model of the distribution of rewards in space that takes into account the cost of time to reach distant locations (i.e., temporally discounting rewards). For an agent with such a goal, we find that an optimal model of exploration in fact produces hyperbolic path lengths, which are well approximated by power laws. We then provide support for our model by showing that humans in a laboratory spatial exploration task search space systematically and modify their search patterns under a cost of time. In addition, we find that path length distributions in a large dataset obtained from free-ranging marine vertebrates are well described by our hyperbolic model. Thus, we provide a general theoretical framework for understanding spatial exploration patterns of cognitively complex foragers.

English Channel towed sledge seabed images. Phase 1: scoping study and example analysis

During the 1970’s and 1980’s, the late Dr Norman Holme undertook extensive towed sledge surveys in the English Channel and some in the Irish Sea. Only a minority of the resulting images were analysed and reported before his death in 1989 but logbooks, video and film material has been archived in the National Marine Biological Library (NMBL) in Plymouth. A scoping study was therefore commissioned by the Joint Nature Conservation Committee and as a part of the Mapping European Seabed Habitats (MESH) project to identify the value of the material archived and the procedure and cost to undertake further work. The results of the scoping study are: 1. NMBL archives hold 106 videotapes (reel-to-reel Sony HD format) and 59 video cassettes (including 15 from the Irish Sea) in VHS format together with 90 rolls of 35 mm colour transparency film (various lengths up to about 240 frames per film). These are stored in the Archive Room, either in a storage cabinet or in original film canisters. 2. Reel-to-reel material is extensive and had already been selectively copied to VHS cassettes. The cost of transferring it to an accepted ‘long-life’ medium (Betamax) would be approximately £15,000. It was not possible to view the tapes as a suitable machine was not located. The value of the tapes is uncertain but they are likely to become beyond salvation within one to two years. 3. Video cassette material is in good condition and is expected to remain so for several more years at least. Images viewed were generally of poor quality and the speed of tow often makes pictures blurred. No immediate action is required. 4. Colour transparency films are in good condition and the images are very clear. They provide the best source of information for mapping seabed biotopes. They should be scanned to digital format but inexpensive fast copying is problematic as there are no between-frame breaks between images and machines need to centre the image based on between-frame breaks. The minimum cost to scan all of the images commercially is approximately £6,000 and could be as much as £40,000 on some quotations. There is a further cost in coding and databasing each image and, all-in-all it would seem most economic to purchase a ‘continuous film’ scanner and undertake the work in-house. 5. Positional information in ships logs has been matched to films and to video tapes. Decca Chain co-ordinates recorded in the logbooks have been converted to latitude and longitude (degrees, minutes and seconds) and a further routine developed to convert to degrees and decimal degrees required for GIS mapping. However, it is unclear whether corrections to Decca positions were applied at the time the position was noted. Tow tracks have been mapped onto an electronic copy of a Hydrographic Office chart. 6. The positions of start and end of each tow were entered to a spread sheet so that they can be displayed on GIS or on a Hydrographic Office Chart backdrop. The cost of the Hydrographic Office chart backdrop at a scale of 1:75,000 for the whole area was £458 incl. VAT. 7. Viewing all of the video cassettes to note habitats and biological communities, even by an experienced marine biologist, would take at least in the order of 200 hours and is not recommended. English Channel towed sledge seabed images. Phase 1: scoping study and example analysis. 6 8. Once colour transparencies are scanned and indexed, viewing to identify seabed habitats and biological communities would probably take about 100 hours for an experienced marine biologist and is recommended. 9. It is expected that identifying biotopes along approximately 1 km lengths of each tow would be feasible although uncertainties about Decca co-ordinate corrections and exact positions of images most likely gives a ±250 m position error. More work to locate each image accurately and solve the Decca correction question would improve accuracy of image location. 10. Using codings (produced by Holme to identify different seabed types), and some viewing of video and transparency material, 10 biotopes have been identified, although more would be added as a result of full analysis. 11. Using the data available from the Holme archive, it is possible to populate various fields within the Marine Recorder database. The overall ‘survey’ will be ‘English Channel towed video sled survey’. The ‘events’ become the 104 tows. Each tow could be described as four samples, i.e. the start and end of the tow and two areas in the middle to give examples along the length of the tow. These samples would have their own latitude/longitude co-ordinates. The four samples would link to a GIS map. 12. Stills and video clips together with text information could be incorporated into a multimedia presentation, to demonstrate the range of level seabed types found along a part of the northern English Channel. More recent images taken during SCUBA diving of reef habitats in the same area as the towed sledge surveys could be added to the Holme images.

Mechanisms of long-term decline in size of lesser sandeels in the North Sea explored using a growth and phenology model

The lesser sandeel Ammodytes marinus is a key species in the North Sea ecosystem, transferring energy from planktonic producers to top predators. Previous studies have shown a long-term decline in the size of 0-group sandeels in the western North Sea, but they were unable to pinpoint the mechanism (later hatching, slower growth or changes in size-dependent mortality) or cause. To investigate the first 2 possibilities we combined 2 independent time series of sandeel size, namely data from chick-feeding Atlantic puffins Fratercula arctica and from the Continuous Plankton Recorder (CPR), in a novel statistical model implemented using Markov Chain Monte Carlo (MCMC). The model estimated annual mean length on 1 July, as well as hatching date and growth rate for sandeels from 1973 to 2006. Mean length-at-date declined by 22% over this period, corresponding to a 60% decrease in energy content, with a sharper decline since 2002. Up to the mid-1990s, the decline was associated with a trend towards later hatching. Subsequently, hatching became earlier again, and the continued trend towards smaller size appears to have been driven by lower growth rates, particularly in the most recent years, although we could not rule out changes in size-dependent mortality. Our findings point to major changes in key aspects of sandeel life history, which we consider are most likely due to direct and indirect temperature-related changes over a range of biotic factors, including the seasonal distribution of copepods and intra- and inter-specific competition with planktivorous fish. The results have implications both for the many predators of sandeels and for age and size of maturation in this aggregation of North Sea sandeels.

Long-Term Retrospective Analysis of Mackerel Spawning in the North Sea: A New Time Series and Modeling Approach to CPR Data

We present a unique view of mackerel (Scomber scombrus) in the North Sea based on a new time series of larvae caught by the Continuous Plankton Recorder (CPR) survey from 1948-2005, covering the period both before and after the collapse of the North Sea stock. Hydrographic backtrack modelling suggested that the effect of advection is very limited between spawning and larvae capture in the CPR survey. Using a statistical technique not previously applied to CPR data, we then generated a larval index that accounts for both catchability as well as spatial and temporal autocorrelation. The resulting time series documents the significant decrease of spawning from before 1970 to recent depleted levels. Spatial distributions of the larvae, and thus the spawning area, showed a shift from early to recent decades, suggesting that the central North Sea is no longer as important as the areas further west and south. These results provide a consistent and unique perspective on the dynamics of mackerel in this region and can potentially resolve many of the unresolved questions about this stock

Zooplankton associations in a Mediterranean long-term time-series

Coastal zooplankton have been investigated since 1984 at a Long Term Ecological Research station MC (LTER-MC) in the inner Gulf of Naples (Tyrrhenian Sea, Western Mediterranean). The sampling site, located between the littoral and the open sea systems, has very active hydrography that affects plankton communities. The present work was aimed at establishing whether, in such a dynamic and variable environment, species associations and homogeneous periods could be identified as characteristic and stable features of the mesozooplankton over the period 1984–2006. Hierarchical clustering was applied to assess species associations based on a matrix of similarities between species (R-mode), and homogeneous periods based on a matrix of similarities between observations (Q-mode). The Indicator Value index [IndVal, Dufrene and Legendre (1997) Species assemblages and indicator species: the need for a flexible asymmetrical approach. Ecol. Monogr., 67, 345–366] was calculated to identify species characterizing each period. Five taxonomic groups with well-defined composition and abundance were identified as robust associations that likely reflect different modes of community functioning. The temporal course of these associations was largely shaped by strong seasonal forcing comprising both physical and biological (e.g. trophic) signals. These associations persisted over the long term, thus indicating some stable characters in the Naples zooplankton time-series, providing evidence of resilience in communities in highly variable coastal conditions.

Variation among northeast Atlantic regions in the responses of zooplankton to climate change: Not all areas follow the same path

Broad scale climate forcing can interact with local environmental processes to affect the observed ecological phenomena. This causes potential problems of over-extrapolation for results from a limited number of sites or the averaging out of region-specific responses if data from too wide an area are combined. In this study, an area similar in extent to the Celtic Biscay Large Marine Ecosystem, but including off-shelf areas, was partitioned using clustering of satellite chlorophyll (chl-a) measurements. The resulting clusters were used to define areas over which to combine copepod data from the Continuous Plankton Recorder. Following filtering due to data limitations, nine regions were defined with sufficient records for analysis. These regions were consistent with known oceanographic structure in the study area. Off-shelf regions showed a progressively later timing in the seasonal peak of chl-a measurements moving northwards. Generalised additive models were used to estimate seasonal and multiannual signals in the adult and juvenile stages of Calanus finmarchicus, C. helgolandicus and the Paracalanus–Pseudocalanus group. Associations between variables (sea surface temperature (SST), phenology and annual abundance) differed among taxonomic groups, but even within taxonomic groups, relationships were not consistent across regions. For example, in the deep waters off Spain and Portugal the annual abundance of Calanus finmarchicus has a weak positive association with SST, in contrast to the pattern in most other regions. The regions defined in this study provide an objective basis for investigations into the long term dynamics of plankton populations and suggest suitable sub regions for deriving pelagic system indicators.

Optimal foraging strategies: Lévy walks balance searching and patch exploitation under a very broad range of conditions

While evidence for optimal random search patterns, known as Lévy walks, in empirical movement data is mounting for a growing list of taxa spanning motile cells to humans, there is still much debate concerning the theoretical generality of Lévy walk optimisation. Here, using a new and robust simulation environment, we investigate in the most detailed study to date (24×10(6) simulations) the foraging and search efficiencies of 2-D Lévy walks with a range of exponents, target resource distributions and several competing models. We find strong and comprehensive support for the predictions of the Lévy flight foraging hypothesis and in particular for the optimality of inverse square distributions of move step-lengths across a much broader range of resource densities and distributions than previously realised. Further support for the evolutionary advantage of Lévy walk movement patterns is provided by an investigation into the 'feast and famine' effect, with Lévy foragers in heterogeneous environments experiencing fewer long 'famines' than other types of searchers. Therefore overall, optimal Lévy foraging results in more predictable resources in unpredictable environments.

Long-term annual primary production in the Ulleung Basin as a biological hot spot in the East/Japan Sea

Although the Ulleung Basin is an important biological hot spot in East/Japan Sea (hereafter the East Sea), very limited knowledge for seasonal and annual variations in the primary productivity exists. In this study, a recent decadal trend of primary production in the Ulleung Basin was analyzed based on MODIS-derived monthly primary production for a better annual production budget. Based on the MODIS-derived primary production, the mean daily primary productivity was 766.8 mg C m-2 d-1 (SD=+/- 196.7 mg C m-2 d-1) and the annual primary productivity was 280.2 g C m-2 yr-1 (SD=+/- 14.9 g C m-2 yr-1) in the Ulleung Basin during the study period. The monthly contributions of primary production were not largely variable among different months, and a relatively small interannual production variability was also observed in the Ulleung Basin, which indicates that the Ulleung Basin is a sustaining biologically productive region called as hot spot in the East Sea. However, a significant recent decline in the annual primary production was observed in the Ulleung Basin after 2006. Although no strong possibilities were found in this study, the current warming sea surface temperature and a negative phase PDO index were suggested for the recent declining primary production. For a better understanding of subsequent effects on marine ecosystems, more intensive interdisciplinary field studies will be required in the Ulleung Basin.

Beyond long-term averages: making biological sense of a rapidly changing world

Biological responses to climate change are typically communicated in generalized terms such as poleward and altitudinal range shifts, but adaptation efforts relevant to management decisions often require forecasts that incorporate the interaction of multiple climatic and nonclimatic stressors at far smaller spatiotemporal scales. We argue that the desire for generalizations has, ironically, contributed to the frequent conflation of weather with climate, even within the scientific community. As a result, current predictions of ecological responses to climate change, and the design of experiments to understand underlying mechanisms, are too often based on broad-scale trends and averages that at a proximate level may have very little to do with the vulnerability of organisms and ecosystems. The creation of biologically relevant metrics of environmental change that incorporate the physical mechanisms by which climate trains patterns of weather, coupled with knowledge of how organisms and ecosystems respond to these changes, can offer insight into which aspects of climate change may be most important to monitor and predict. This approach also has the potential to enhance our ability to communicate impacts of climate change to nonscientists and especially to stakeholders attempting to enact climate change adaptation policies.