A multi-gene phylogeny of Cephalopoda supports convergent morphological evolution in association with multiple habitat shifts in the marine environment

ABSTRACT: BACKGROUND: The marine environment is comprised of numerous divergent organisms living under similar selective pressures, often resulting in the evolution of convergent structures such as the fusiform body shape of pelagic squids, fishes, and some marine mammals. However, little is known about the frequency of, and circumstances leading to, convergent evolution in the open ocean. Here, we present a comparative study of the molluscan class Cephalopoda, a marine group known to occupy habitats from the intertidal to the deep sea. Several lineages bear features that may coincide with a benthic or pelagic existence, making this a valuable group for testing hypotheses of correlated evolution. To test for convergence and correlation, we generate the most taxonomically comprehensive multi-gene phylogeny of cephalopods to date. We then create a character matrix of habitat type and morphological characters, which we use to infer ancestral character states and test for correlation between habitat and morphology. RESULTS: Our study utilizes a taxonomically well-sampled phylogeny to show convergent evolution in all six morphological characters we analyzed. Three of these characters also correlate with habitat. The presence of an autogenic photophore is correlated with a pelagic habitat, while the cornea and accessory nidamental gland correlate with a benthic lifestyle. Here, we present the first statistical tests for correlation between convergent traits and habitat in cephalopods to better understand the evolutionary history of characters that are adaptive in benthic or pelagic environments, respectively. DISCUSSION: Our study supports the hypothesis that habitat has influenced convergent evolution in the marine environment: benthic organisms tend to exhibit similar characteristics that confer protection from invasion by other benthic taxa, while pelagic organisms possess features that facilitate crypsis and communication in an environment lacking physical refuges. Features that have originated multiple times in distantly related lineages are likely adaptive for the organisms inhabiting a particular environment: studying the frequency and evolutionary history of such convergent characters can increase understanding of the underlying forces driving ecological and evolutionary transitions in the marine environment.     

Evolution of cannibalism in the larval stage of pelagic fish

Larvae of several ocean pelagic fish species, such as tunas and marlins, have been known to have large jaws, but the ecological significance of this unique morphological character has been hardly analyzed in evolutionary ecology. Pelagic spawners produce small and nutrition-poor ova, and spawning and nursery grounds of the open ocean migratory fishes are oligotrophic. We hypothesize that cannibalism would be a possible life style in the larval period and the large mouth gape would be an adaptive morphological characteristic for a cannibal in the oligotrophic pelagic environment. We showed that mouth gape size of the open ocean pelagic fish is significantly larger than that of offshore/coastal pelagic fish in larval period. A mathematical model demonstrated that cannibalism would tend to evolve in high sea environment. Our findings suggest an evolutionary pattern of cannibalism trait in the larval stage of pelagic fishes. This revised version was published online in July 2006 with corrections to the Cover Date.     

Projections of climate‐driven changes in tuna vertical habitat based on species‐specific differences in blood oxygen affinity

Oxygen concentrations are hypothesized to decrease in many areas of the ocean as a result of anthropogenically driven climate change, resulting in habitat compression for pelagic animals. The oxygen partial pressure, pO₂, at which blood is 50% saturated (P₅₀) is a measure of blood oxygen affinity and a gauge of the tolerance of animals for low ambient oxygen. Tuna species display a wide range of blood oxygen affinities (i.e., P₅₀ values) and therefore may be differentially impacted by habitat compression as they make extensive vertical movements to forage on subdaily time scales. To project the effects of end‐of‐the‐century climate change on tuna habitat, we calculate tuna P₅₀ depths (i.e., the vertical position in the water column at which ambient pO₂ is equal to species‐specific blood P₅₀ values) from 21st century Earth System Model (ESM) projections included in the fifth phase of the Climate Model Intercomparison Project (CMIP5). Overall, we project P₅₀ depths to shoal, indicating likely habitat compression for tuna species due to climate change. Tunas that will be most impacted by shoaling are Pacific and southern bluefin tunas—habitat compression is projected for the entire geographic range of Pacific bluefin tuna and for the spawning region of southern bluefin tuna. Vertical shifts in P₅₀ depths will potentially influence resource partitioning among Pacific bluefin, bigeye, yellowfin, and skipjack tunas in the northern subtropical and eastern tropical Pacific Ocean, the Arabian Sea, and the Bay of Bengal. By establishing linkages between tuna physiology and environmental conditions, we provide a mechanistic basis to project the effects of anthropogenic climate change on tuna habitats.     

Understanding Oceanic Migrations with Intrinsic Biogeochemical Markers

Migratory marine vertebrates move annually across remote oceanic water masses crossing international borders. Many anthropogenic threats such as overfishing, bycatch, pollution or global warming put millions of marine migrants at risk especially during their long-distance movements. Therefore, precise knowledge about these migratory movements to understand where and when these animals are more exposed to human impacts is vital for addressing marine conservation issues. Because electronic tracking devices suffer from several constraints, mainly logistical and financial, there is emerging interest in finding appropriate intrinsic markers, such as the chemical composition of inert tissues, to study long-distance migrations and identify wintering sites. Here, using tracked pelagic seabirds and some of their own feathers which were known to be grown at different places and times within the annual cycle, we proved the value of biogeochemical analyses of inert tissue as tracers of marine movements and habitat use. Analyses of feathers grown in summer showed that both stable isotope signatures and element concentrations can signal the origin of breeding birds feeding in distinct water masses. However, only stable isotopes signalled water masses used during winter because elements mainly accumulated during the long breeding period are incorporated into feathers grown in both summer and winter. Our findings shed new light on the simple and effective assignment of marine organisms to distinct oceanic areas, providing new opportunities to study unknown migration patterns of secretive species, including in relation to human-induced mortality on specific populations in the marine environment.     

Long-Term GPS Tracking of Ocean Sunfish Mola mola Offers a New Direction in Fish Monitoring

Satellite tracking of large pelagic fish provides insights on free-ranging behaviour, distributions and population structuring. Up to now, such fish have been tracked remotely using two principal methods: direct positioning of transmitters by Argos polar-orbiting satellites, and satellite relay of tag-derived light-level data for post hoc track reconstruction. Error fields associated with positions determined by these methods range from hundreds of metres to hundreds of kilometres. However, low spatial accuracy of tracks masks important details, such as foraging patterns. Here we use a fast-acquisition global positioning system (Fastloc GPS) tag with remote data retrieval to track long-term movements, in near real time and position accuracy of <70 m, of the world''s largest bony fish, the ocean sunfish Mola mola. Search-like movements occurred over at least three distinct spatial scales. At fine scales, sunfish spent longer in highly localised areas with faster, straighter excursions between them. These ‘stopovers’ during long-distance movement appear consistent with finding and exploiting food patches. This demonstrates the feasibility of GPS tagging to provide tracks of unparalleled accuracy for monitoring movements of large pelagic fish, and with nearly four times as many locations obtained by the GPS tag than by a conventional Argos transmitter. The results signal the potential of GPS-tagged pelagic fish that surface regularly to be detectors of resource ‘hotspots’ in the blue ocean and provides a new capability for understanding large pelagic fish behaviour and habitat use that is relevant to ocean management and species conservation.     

人工集鱼装置对热带金枪鱼类摄食模式的影响研究进展

热带金枪鱼类具有聚集在漂浮物体周围的行为特性,形成的集群表现稳定,可捕性较高。以此特性,金枪鱼围网渔业研制并投放了大量的人工集鱼装置(Fish Aggregation Device,FAD)用于聚集并捕捞金枪鱼。然而,大规模出现的FAD会使某些海域海面漂浮物的密度迅速增加,从而在一定程度上人为地改变了金枪鱼的表层栖息环境,对金枪鱼种群具有一系列可能的潜在负面影响,摄食模式的改变就是其中之一。归纳并综合了近年来国内外关于FAD对金枪鱼类摄食模式影响的相关研究,从摄食行为、日摄食量、饵料种类与组成以及生态位宽度4个方面对比了随附于FAD的金枪鱼和自由状态下同类的不同,发现大多情况下FAD的存在会使金枪鱼的摄食模式发生一定的改变。最后,归纳了过往实验存在的不足,对今后研究的发展方向进行了展望。     

Early‐life foraging: Behavioral responses of newly fledged albatrosses to environmental conditions

In order to survive and later recruit into a population, juvenile animals need to acquire resources through the use of innate and/or learnt behaviors in an environment new to them. For far‐ranging marine species, such as the wandering albatross Diomedea exulans, this is particularly challenging as individuals need to be able to rapidly adapt and optimize their movement strategies in response to the highly dynamic and heterogeneous nature of their open‐ocean pelagic habitats. Critical to this is the development and flexibility of dispersal and exploratory behaviors. Here, we examine the movements of eight juvenile wandering albatrosses, tracked using GPS/Argos satellite transmitters for eight months following fledging, and compare these to the trajectories of 17 adults to assess differences and similarities in behavioral strategies through time. Behavioral clustering algorithms (Expectation Maximization binary Clustering) were combined with multinomial regression analyses to investigate changes in behavioral mode probabilities over time, and how these may be influenced by variations in day duration and in biophysical oceanographic conditions. We found that juveniles appeared to quickly acquire the same large‐scale behavioral strategies as those employed by adults, although generally more time was spent resting at night. Moreover, individuals were able to detect and exploit specific oceanographic features in a manner similar to that observed in adults. Together, the results of this study suggest that while shortly after fledging juvenile wandering albatrosses are able to employ similar foraging strategies to those observed in adults, additional skills need to be acquired during the immature period before the efficiency of these behaviors matches that of adults.     

Dynamic habitat models: using telemetry data to project fisheries bycatch

Fisheries bycatch is a recognized threat to marine megafauna. Addressing bycatch of pelagic species however is challenging owing to the dynamic nature of marine environments and vagility of these organisms. In order to assess the potential for species to overlap with fisheries, we propose applying dynamic habitat models to determine relative probabilities of species occurrence for specific oceanographic conditions. We demonstrate this approach by modelling habitats for Laysan (Phoebastria immutabilis) and black-footed albatrosses (Phoebastria nigripes) using telemetry data and relating their occurrence probabilities to observations of Hawaii-based longline fisheries in 1997-2000. We found that modelled habitat preference probabilities of black-footed albatrosses were high within some areas of the fishing range of the Hawaiian fleet and such preferences were important in explaining bycatch occurrence. Conversely, modelled habitats of Laysan albatrosses overlapped little with Hawaii-based longline fisheries and did little to explain the bycatch of this species. Estimated patterns of albatross habitat overlap with the Hawaiian fleet corresponded to bycatch observations: black-footed albatrosses were more frequently caught in this fishery despite being 10 times less abundant than Laysan albatrosses. This case study demonstrates that dynamic habitat models based on telemetry data may help to project interactions with pelagic animals relative to environmental features and that such an approach can serve as a tool to guide conservation and management decisions.     

Ocean Heat Content Reveals Secrets of Fish Migrations

For centuries, the mechanisms surrounding spatially complex animal migrations have intrigued scientists and the public. We present a new methodology using ocean heat content (OHC), a habitat metric that is normally a fundamental part of hurricane intensity forecasting, to estimate movements and migration of satellite-tagged marine fishes. Previous satellite-tagging research of fishes using archival depth, temperature and light data for geolocations have been too coarse to resolve detailed ocean habitat utilization. We combined tag data with OHC estimated from ocean circulation and transport models in an optimization framework that substantially improved geolocation accuracy over SST-based tracks. The OHC-based movement track provided the first quantitative evidence that many of the tagged highly migratory fishes displayed affinities for ocean fronts and eddies. The OHC method provides a new quantitative tool for studying dynamic use of ocean habitats, migration processes and responses to environmental changes by fishes, and further, improves ocean animal tracking and extends satellite-based animal tracking data for other potential physical, ecological, and fisheries applications.     

Selective advantages conferred by the high performance physiology of tunas,billfishes, and dolphin fish

Tunas are extensively distributed throughout world's oceans and grow and reproduce fast enough to support one of the world's largest commercial fisheries. Yet they are apex predators living in the energy depauperate pelagic environment. It is often presumed that tunas evolved their specialized anatomy, physiology, and biochemistry to be capable of (a) high maximum swimming speeds, (b) high sustained swimming speeds, and/or (c) very efficient swimming, all of which help account for their wide distribution and reproductive success. However, a growing body of data on the energetics and physiological abilities of tunas do not support these assumptions. The three things demonstratively “high performance” about tunas, and probably other pelagic species such as marlin (Makaira spp. and Tetrapturus spp.) and dolphin fish (Coryphaena spp.), are (a) rates of somatic and gonadal growth, (b) rates of digestion, (c) rates of recovery from exhaustive exercise (i.e., clearance of muscle lactate and the concomitant acid load). All of these are energy consuming processes requiring rates of oxygen and substrate delivery above those needed by the swimming muscles for sustained propulsion and for other routine metabolic activities. I hypothesize that the ability of high performance pelagic species (tunas, billfishes, and dolphin fish) to deliver oxygen and metabolic substrates to the tissues at high rates evolved to permit rapid somatic and gonadal growth, rapid digestion, and rapid recovery from exhaustive exercise (abilities central to success in the pelagic environment), not exceptionally high sustained swimming speeds.     

Estimating Trans-Seasonal Variability in Water Column Biomass for a Highly Migratory,Deep Diving Predator

The deployment of animal-borne electronic tags is revolutionizing our understanding of how pelagic species respond to their environment by providing in situ oceanographic information such as temperature, salinity, and light measurements. These tags, deployed on pelagic animals, provide data that can be used to study the ecological context of their foraging behaviour and surrounding environment. Satellite-derived measures of ocean colour reveal temporal and spatial variability of surface chlorophyll-a (a useful proxy for phytoplankton distribution). However, this information can be patchy in space and time resulting in poor correspondence with marine animal behaviour. Alternatively, light data collected by animal-borne tag sensors can be used to estimate chlorophyll-a distribution. Here, we use light level and depth data to generate a phytoplankton index that matches daily seal movements. Time-depth-light recorders (TDLRs) were deployed on 89 southern elephant seals (Mirounga leonina) over a period of 6 years (1999–2005). TDLR data were used to calculate integrated light attenuation of the top 250 m of the water column (LA₂₅₀), which provided an index of phytoplankton density at the daily scale that was concurrent with the movement and behaviour of seals throughout their entire foraging trip. These index values were consistent with typical seasonal chl-a patterns as measured from 8-daySea-viewing Wide Field-of-view Sensor (SeaWiFs) images. The availability of data recorded by the TDLRs was far greater than concurrent remotely sensed chl-a at higher latitudes and during winter months. Improving the spatial and temporal availability of phytoplankton information concurrent with animal behaviour has ecological implications for understanding the movement of deep diving predators in relation to lower trophic levels in the Southern Ocean. Light attenuation profiles recorded by animal-borne electronic tags can be used more broadly and routinely to estimate lower trophic distribution at sea in relation to deep diving predator foraging behaviour.     

Human-induced salinity changes impact marine organisms and ecosystems

Climate change is fundamentally altering marine and coastal ecosystems on a global scale. While the effects of ocean warming and acidification on ecology and ecosystem functions and services are being comprehensively researched, less attention is directed toward understanding the impacts of human-driven ocean salinity changes. The global water cycle operates through water fluxes expressed as precipitation, evaporation, and freshwater runoff from land. Changes to these in turn modulate ocean salinity and shape the marine and coastal environment by affecting ocean currents, stratification, oxygen saturation, and sea level rise. Besides the direct impact on ocean physical processes, salinity changes impact ocean biological functions with the ecophysiological consequences are being poorly understood. This is surprising as salinity changes may impact diversity, ecosystem and habitat structure loss, and community shifts including trophic cascades. Climate model future projections (of end of the century salinity changes) indicate magnitudes that lead to modification of open ocean plankton community structure and habitat suitability of coral reef communities. Such salinity changes are also capable of affecting the diversity and metabolic capacity of coastal microorganisms and impairing the photosynthetic capacity of (coastal and open ocean) phytoplankton, macroalgae, and seagrass, with downstream ramifications on global biogeochemical cycling. The scarcity of comprehensive salinity data in dynamic coastal regions warrants additional attention. Such datasets are crucial to quantify salinity-based ecosystem function relationships and project such changes that ultimately link into carbon sequestration and freshwater as well as food availability to human populations around the globe. It is critical to integrate vigorous high-quality salinity data with interacting key environmental parameters (e.g., temperature, nutrients, oxygen) for a comprehensive understanding of anthropogenically induced marine changes and its impact on human health and the global economy.     

Habitat and behaviour of yellowfin tuna Thunnus albacares in the Gulf of Mexico determined using pop-up satellite archival tags

This study presents the first data on movement, habitat use and behaviour for yellowfin tuna Thunnus albacares in the Atlantic Basin. Six individuals were tracked in the Gulf of Mexico using pop-up satellite archival tags. Records up to 80 days in length were obtained, providing information on depth and temperature preferences as well as horizontal movements. Thunnus albacares in the Gulf of Mexico showed a strong preference for the mixed layer and thermocline, consistent with findings for this species in other ocean basins. Fish showed a diel pattern in depth distribution, remaining in surface and mixed layer waters at night and diving to deeper waters during the day. The vertical extent of T. albacares habitat appeared to be temperature limited, with fish generally avoiding waters that were >6° C cooler than surface waters. The vertical and thermal habitat usage of T. albacares differs from that of bigeye Thunnus obesus and bluefin Thunnus thynnus , Thunnus orientalis and Thunnus maccoyii tunas. These results are consistent with the results of earlier studies conducted on T. albacares in other oceans.     

Unique adaptations of the metabolic biochemistry of tunas and billfishes for life in the pelagic environment

Synopsis Virtually all characteristics of tunas and billfishes reflect their highly charged lifestyles as apex predators in the oceanic pelagic environment. The adaptations they possess for efficient and rapid swimming, efficient and rapid food processing, turnover of nutrients and storage and mobilization of internal fuel supplies, and for rapid recovery rates, are discussed. Overall, tunas and billfishes are designed for high performance, at both sustainable and burst swimming speeds, but there are several differences between tunas and billfishes. Tunas' aerobic metabolic capacities exceed those of ectothermic fishes, including billfishes and other scombrids, by virtue of their elevated red muscle temperatures, and because heart and white muscle aerobic capacities are significantly greater in tunas. The adaptations for high performance involve some costs, including the need for a constant high energy input to sustain high metabolic rates, high activity levels, and endothermy, Yet, tunas and billfishes have adopted successful lifestyles, as evidenced by their large numbers and biomass within the marine environment. Although our knowledge of these fishes has increased dramatically during the past 15 years, there are major gaps in our understanding of the metabolic biochemistry and physiology of these fishes, and these are highlighted so that additional research can be directed towards filling these gaps.Paper from the International Union of Biological Societies symposium The biology of tunas and billfishes: an examination of life on the knife edge, organized by Richard W. Brill and Kim N. Holland.     

Trait biogeography of marine copepods – an analysis across scales

Functional traits, rather than taxonomic identity, determine the fitness of individuals in their environment: traits of marine organisms are therefore expected to vary across the global ocean as a function of the environment. Here, we quantify such spatial and seasonal variations based on extensive empirical data and present the first global biogeography of key traits (body size, feeding mode, relative offspring size and myelination) for pelagic copepods, the major group of marine zooplankton. We identify strong patterns with latitude, season and between ocean basins that are partially (c. 50%) explained by key environmental drivers. Body size, for example decreases with temperature, confirming the temperature‐size rule, but surprisingly also with productivity, possibly driven by food‐chain length and size‐selective predation. Patterns unrelated to environmental predictors may originate from phylogenetic clustering. Our maps can be used as a test‐bed for trait‐based mechanistic models and to inspire next‐generation biogeochemical models.     

Biodiversity's Big Wet Secret: The Global Distribution of Marine Biological Records Reveals Chronic Under-Exploration of the Deep Pelagic Ocean

Background

Understanding the distribution of marine biodiversity is a crucial first step towards the effective and sustainable management of marine ecosystems. Recent efforts to collate location records from marine surveys enable us to assemble a global picture of recorded marine biodiversity. They also effectively highlight gaps in our knowledge of particular marine regions. In particular, the deep pelagic ocean – the largest biome on Earth – is chronically under-represented in global databases of marine biodiversity.

Methodology/Principal Findings

We use data from the Ocean Biogeographic Information System to plot the position in the water column of ca 7 million records of marine species occurrences. Records from relatively shallow waters dominate this global picture of recorded marine biodiversity. In addition, standardising the number of records from regions of the ocean differing in depth reveals that regardless of ocean depth, most records come either from surface waters or the sea bed. Midwater biodiversity is drastically under-represented.

Conclusions/Significance

The deep pelagic ocean is the largest habitat by volume on Earth, yet it remains biodiversity''s big wet secret, as it is hugely under-represented in global databases of marine biological records. Given both its value in the provision of a range of ecosystem services, and its vulnerability to threats including overfishing and climate change, there is a pressing need to increase our knowledge of Earth''s largest ecosystem.     

Climate impacts and oceanic top predators: moving from impacts to adaptation in oceanic systems

Climate impacts are now widely reported from coastal marine systems, but less is known for the open ocean. Here we review progress in understanding impacts on large pelagic species presented at an international workshop for the Climate Impacts on Oceanic Top Predators programme, and discuss the future with regard to the next phase of adaptation-focused research. Recent highlights include a plan to map the distribution of key species in the foodweb using both acoustics and biochemical techniques, and development of a new data sharing and access tool for fisheries and associated data, including socio-economic information. A common research focus in pelagic ecosystems is on understanding climate variability and climate change impacts on marine species, but a greater emphasis on developing future scenarios and adaptation options is needed. Workshop participants also concluded that engagement with and provision of science support to regional fisheries management organisations are critical elements for ensuring successful uptake of research. This uptake will be required for future management of fisheries as global warming continues such that some open ocean top predators can be sustainably harvested, impacts on conservation-dependent species can be avoided, and ecosystem function is not compromised.     

Climate change alters stability and species potential interactions in a large marine ecosystem

We have little empirical evidence of how large‐scale overlaps between large numbers of marine species may have altered in response to human impacts. Here, we synthesized all available distribution data (>1 million records) since 1992 for 61 species of the East Australian marine ecosystem, a global hot spot of ocean warming and continuing fisheries exploitation. Using a novel approach, we constructed networks of the annual changes in geographical overlaps between species. Using indices of changes in species overlap, we quantified changes in the ecosystem stability, species robustness, species sensitivity and structural keystone species. We then compared the species overlap indices with environmental and fisheries data to identify potential factors leading to the changes in distributional overlaps between species. We found that the structure of the ecosystem has changed with a decrease in asymmetrical geographical overlaps between species. This suggests that the ecosystem has become less stable and potentially more susceptible to environmental perturbations. Most species have shown a decrease in overlaps with other species. The greatest decrease in species overlap robustness and sensitivity to the loss of other species has occurred in the pelagic community. Some demersal species have become more robust and less sensitive. Pelagic structural keystone species, predominately the tunas and billfish, have been replaced by demersal fish species. The changes in species overlap were strongly correlated with regional oceanographic changes, in particular increasing ocean warming and the southward transport of warmer and saltier water with the East Australian Current, but less correlated with fisheries catch. Our study illustrates how large‐scale multispecies distribution changes can help identify structural changes in marine ecosystems associated with climate change.     

A comparative study of asymmetric migration events across a marine biogeographic boundary

In many nonclonal, benthic marine species, geographic distribution is mediated by the dispersal of their larvae. The dispersal and recruitment of marine larvae may be limited by temperature gradients that can affect mortality or by ocean currents that can directly affect the movements of pelagic larvae. We focus on Point Conception, a well-known biogeographic boundary between the Californian and Oregonian biogeographic provinces, to investigate whether ocean currents affect patterns of gene flow in intertidal marine invertebrates. The predominance of pelagically dispersing species with northern range limits at Point Conception suggests that ocean currents can affect species distributions by erecting barriers to the dispersal of planktonic larvae. In this paper, we investigate whether the predominantly southward currents have left a recognizable genetic signature in species with pelagically dispersing larvae whose ranges span Point Conception. We use patterns of genetic diversity and a new method for inferring cladistic migration events to test the hypothesis that southward currents increase southward gene flow for species with pelagically dispersing larvae. We collected mitochondrial DNA (mtDNA) sequence data for the barnacles Balanus glandula and Chthamalus fissus and also reanalyzed a previously published mtDNA dataset (Strongylocentrotus purpuratus, Edmands et al. 1996). For all three species, our cladistic approach identified an excess of southward migration events across Point Conception. In data from a fourth species with nondispersing larvae (Nucella emarginata, Marko 1998), our method suggests that ocean currents have not played a role in generating genetic structure.     

Living krill,zooplankton and experimental investigations: a discourse on the role of krill and their experimental study in marine ecology

Krill occupy critical positions in a many marine ecosystems and have been the subject of a number of concerted studies yet there are large areas of their biology that still remain a mystery. Most species of krill are open ocean animals, which makes direct observation and sampling difficult. Krill also exhibit a number of physiological and behavioural attributes which frustrate attempts to understand their life history. Krill are conceptually difficult to come to terms with; they are obviously different from larger marine organisms such as squid, fish, whales and fish yet they are also quite distinct from those animals classed as zooplankton such as copepods. Despite these differences they have most often been grouped with zooplankton and have been studied using techniques developed for animals which are orders of magnitude smaller than they. This mismatch has affected our view of their interactions with the physical world and also affects their perceived trophic interactions. Their size and mobility also interferes with our ability to sample them effectively and thus to develop our appreciation of their true role in the marine ecosystem. Understanding how intermediate-sized animals, such as krill, function in aquatic ecosystem is critical to better management of the marine environment.