Ontogeny of behaviour in larvae of marine demersal fishes

The development of behaviours that are relevant to larval dispersal of marine, demersal fishes is poorly understood. This review focuses on recent work that attempts to quantify the development of swimming, orientation, vertical distribution and sensory abilities. These behaviours are developed enough to influence dispersal outcomes during most of the pelagic larval stage. Larvae swim in the ocean at speeds similar to the currents found in many locations and at 3–15 body lengths per second (BL s⁻¹), although, based on laboratory measurements, species from cold environments swim slower than those from warm environments. At least in warm-water species, larvae swim in an inertial hydrodynamic environment for most of their pelagic period. Unfed swimming endurance is >10 km from about 8–10 mm, and reaches more than 50 km before settlement in several species. Larval fishes are efficient swimmers. In most species, a large majority of larvae have orientated swimming in the ocean, but the precision of orientation does not improve with growth. Swimming direction of the larvae frequently changes ontogenetically. Vertical distribution changes ontogenetically in most species, and both ontogenetic ascents and descents are found. Development of schooling is poorly understood, but it may influence speed, orientation and vertical distribution. Sensory abilities (hearing, olfaction, vision) form early, are well developed and are able to detect cues relevant to orientation for most of the pelagic larval stage. All this indicates that the passive portion of the pelagic larval duration will be short, at least in most warm-water species, and that behaviour must be taken into account when considering dispersal, and in particular in dispersal models. Although quantitative information on the ontogeny of some behaviours is available for a relatively small number of species, more research in this field is required, especially on species from colder waters.     

Visual conditions and habitat shape the coloration of the Eurasian perch (Perca fluviatilis L.): a trade‐off between camouflage and communication?

In theory, selection for effective camouflage (i.e. dull coloration) in fish should be strongest when the conditions for visual predation are most favourable, such as in structurally simple pelagic habitats. By contrast, in more sheltered (e.g. littoral) habitats, selection may favour effective intra‐specific communication (i.e. bright coloration) (at the expense of crypsis). Poor transparency, as in highly humic waters, should constrain colour adaptations. We investigated phenotypic variation in body coloration of Eurasian perch (Perca fluviatilis L.) in littoral and pelagic habitats of four humic boreal lakes. Perch from the most transparent lake had the lightest and less coloured belly and perch were more colourful in the littoral habitats than in the pelagic areas, with the pattern being clearest in the most transparent lake. In addition, perch in the most transparent lake exhibited sexual dichromatism, with males having a more colourful belly than the females, whereas no indications of sexual dichromatism were found in more humic lakes. Moreover, in the most transparent lake, the condition of fish correlated with bright belly coloration in the littoral, but with dull belly coloration in the pelagic habitat. The results obtained in the present study suggest that selection on perch coloration may differ between lakes as a result of visual properties of the water, and within lakes as a result of divergent selection for camouflage and communication in pelagic and littoral habitats. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 47–59.     

Comparative visual acuity of coleoid cephalopods

The pelagic realm of the ocean is characterized by extremelyclear water and a lack of surfaces. Adaptations to the visualecology of this environment include transparency, fluorescence,bioluminescence, and deep red or black pigmentation. While thesignals that pelagic organisms send are increasingly well-understood,the optical capabilities of their viewers, especially for predatorswith camera-like vision such as fish and squid, are almost unknown.Aquatic camera-like vision is characterized by a spherical lensfocusing an image on the retina. Here, we measured the resolvingpower of the lenses of eight species of pelagic cephalopodsto obtain an approximation of their visual capabilities. Wedid this by focusing a standard resolution target through dissectedlenses and calculating their modulation transfer functions.The modulation transfer function (MTF) is the single most completeexpression of the resolving capabilities of a lens. Since theoptical and retinal capabilities of an eye are generally well-matched,we considered our measurements of cephalopod lens MTF to bea good proxy for their visual capabilities in vivo. In general,squid have optical capabilities comparable to other organismsgenerally assumed to have good vision, such as fish and birds.Surprisingly, the optical capability of the eye of Vampyroteuthisinfernalis rivals that of humans.     

Water Transparency Drives Intra-Population Divergence in Eurasian Perch (Perca fluviatilis)

Trait combinations that lead to a higher efficiency in resource utilization are important drivers of divergent natural selection and adaptive radiation. However, variation in environmental features might constrain foraging in complex ways and therefore impede the exploitation of critical resources. We tested the effect of water transparency on intra-population divergence in morphology of Eurasian perch (Perca fluviatilis) across seven lakes in central Sweden. Morphological divergence between near-shore littoral and open-water pelagic perch substantially increased with increasing water transparency. Reliance on littoral resources increased strongly with increasing water transparency in littoral populations, whereas littoral reliance was not affected by water transparency in pelagic populations. Despite the similar reliance on pelagic resources in pelagic populations along the water transparency gradient, the utilization of particular pelagic prey items differed with variation in water transparency in pelagic populations. Pelagic perch utilized cladocerans in lakes with high water transparency and copepods in lakes with low water transparency. We suggest that under impaired visual conditions low utilization of littoral resources by littoral perch and utilization of evasive copepods by pelagic perch may lead to changes in morphology. Our findings indicate that visual conditions can affect population divergence in predator populations through their effects on resource utilization.     

Interactions of pelagic cnidarians and ctenophores with fish: a review

Medusae, siphonophores and ctenophores (here grouped as `pelagic coelenterates') interact with fish in several ways. Some interactions are detrimental to fish populations, such as predation by gelatinous species on pelagic eggs and larvae of fish, the potential competition for prey among pelagic coelenterates and fish larvae and zooplanktivorous fish species, and pelagic coelenterates serving as intermediate hosts for fish parasites. Other interactions are positive for fish, such as predation by fish on gelatinous species and commensal associations among fish and pelagic coelenterates. The interactions range from beneficial for the gelatinous species (food, parasite removal), to negative (predation on them). We review existing information and present new data on these topics. Although such interactions have been documented frequently, the significance to either fish or pelagic coelenterate populations is poorly understood. The effects of pelagic coelenterates on fish populations are of particular interest because of the great importance of fisheries to the global economy. As fishing pressures mount, it becomes increasingly important to understand how they may influence the balance between pelagic coelenterates and fish.     

Does new technology inspire new directions? Examples drawn from pelagic visual ecology

Oceanography has seen the recent development of many new toolsand techniques. The subfield of pelagic visual ecology in particularhas benefited from the development of more reliable, portable,and economic tools and techniques that can be taken to sea includingspectrometers, microspectrophotometery, electroretinography,and ultraviolet and polarization imaging systems. These advanceshave led to a relative wealth of data on the visual physiologyof pelagic species and on the optical properties of these speciesand their environment. These data, particularly in combinationwith computational methods, have tested long-standing hypothesesin pelagic ecology and led to new hypotheses and research directions.While the ability to study pelagic species still lags far behindwhat is possible in terrestrial and coastal environments, arenaissance may be developing in the study of the integrativebiology of pelagic species.     

High Optical Transmission in a Hybrid Plasmonic-Optical Structure with a Continuous Metal Film

Metals are naturally opaque for electromagnetic (EM) waves below violet frequency due to the Coulomb screening effect. In this letter, we demonstrate high optical transparency of a seamless continuous metal film by sandwiching it in a hybrid plasmonic-optical structure. The proposed structure consists of a plasmonic array and an optical cavity, which exhibits magnetic plasmon (MP) resonance and optical Fabry-Perot (FP) resonance, respectively. An optical transparency of 84% in the near-IR regime is achieved making use of interaction between the plasmonic and optical modes. Furthermore, spectral tunability of the high transparency is demonstrated and robustness under oblique incidence is examined. This work may give insights into plasmonic-optical interactions and may be a potential candidate for transparent electrodes.     

Computational visual ecology in the pelagic realm

Visual performance and visual interactions in pelagic animals are notoriously hard to investigate because of our restricted access to the habitat. The pelagic visual world is also dramatically different from benthic or terrestrial habitats, and our intuition is less helpful in understanding vision in unfamiliar environments. Here, we develop a computational approach to investigate visual ecology in the pelagic realm. Using information on eye size, key retinal properties, optical properties of the water and radiance, we develop expressions for calculating the visual range for detection of important types of pelagic targets. We also briefly apply the computations to a number of central questions in pelagic visual ecology, such as the relationship between eye size and visual performance, the maximum depth at which daylight is useful for vision, visual range relations between prey and predators, counter-illumination and the importance of various aspects of retinal physiology. We also argue that our present addition to computational visual ecology can be developed further, and that a computational approach offers plenty of unused potential for investigations of visual ecology in both aquatic and terrestrial habitats.     

Yeasts from an oligotrophic lake in Patagonia (Argentina): diversity, distribution and synthesis of photoprotective compounds and extracellular enzymes

Nahuel Huapi (NH) Lake is an oligotrophic temperate lake of glacial origin with high transparency, surrounded by well-developed forests and located at San Carlos de Bariloche, Nahuel Huapi National Park, in Patagonia, Argentina. In this lake, we characterized yeast distribution and diversity along a south-to-north transect and established a relationship between the ability to produce photoprotective compounds (PPCs) (carotenoid pigments and mycosporines) and the occurrence of yeast at different collection points. Subsurface water samples were filtered for yeast isolation. Total yeast counts ranged between 22 and 141?CFU?L(-1) , and the highest values corresponded to the most impacted sites. Littoral sites had a low proportion of yeast-producing PPCs and this group prevailed in pelagic sites. This is probably a result of the high transparency of the water and the increased UV exposure. The yeast community from NH Lake showed a high species richness and a uniform distribution of taxa between pelagic and border collection points. Yeasts were identified as belonging to 14 genera and 34 species. Rhodotorula mucilaginosa and Cryptococcus victoriae were the most frequently found species, representing 14.4% and 13.6% of the total yeast isolates, respectively. Most of the yeast isolates demonstrated at least one extracellular enzymatic activity (mainly cellulase and lipase activities), which suggested that these microorganisms are metabolically active in the lake.     

Polarization sensitivity as a contrast enhancer in pelagic predators: lessons from in situ polarization imaging of transparent zooplankton

Because light in the pelagic environment is partially polarized, it has been suggested that the polarization sensitivity found in certain pelagic species may serve to enhance the contrast of their transparent zooplankton prey. We examined its potential during cruises in the Gulf of Mexico and Atlantic Ocean and at a field station on the Great Barrier Reef. First, we collected various species of transparent zooplankton and micronekton and photographed them between crossed polarizers. Many groups, particularly the cephalopods, pelagic snails, salps and ctenophores, were found to have ciliary, muscular or connective tissues with striking birefringence. In situ polarization imagery of the same species showed that, while the degree of underwater polarization was fairly high (approx. 30% in horizontal lines of sight), tissue birefringence played little to no role in increasing visibility. This is most likely due to the low radiance of the horizontal background light when compared with the downwelling irradiance. In fact, the dominant radiance and polarization contrasts are due to unpolarized downwelling light that has been scattered from the animal viewed against the darker and polarized horizontal background light. We show that relatively simple algorithms can use this negative polarization contrast to increase visibility substantially.     

The Influence of Topographic and Dynamic Cyclic Variables on the Distribution of Small Cetaceans in a Shallow Coastal System

The influence of topographic and temporal variables on cetacean distribution at a fine-scale is still poorly understood. To study the spatial and temporal distribution of harbour porpoise Phocoena phocoena and the poorly known Risso’s dolphin Grampus griseus we carried out land-based observations from Bardsey Island (Wales, UK) in summer (2001–2007). Using Kernel analysis and Generalized Additive Models it was shown that porpoises and Risso’s appeared to be linked to topographic and dynamic cyclic variables with both species using different core areas (dolphins to the West and porpoises to the East off Bardsey). Depth, slope and aspect and a low variation in current speed (for Risso’s) were important in explaining the patchy distributions for both species. The prime temporal conditions in these shallow coastal systems were related to the tidal cycle (Low Water Slack and the flood phase), lunar cycle (a few days following the neap tidal phase), diel cycle (afternoons) and seasonal cycle (peaking in August) but differed between species on a temporary but predictable basis. The measure of tidal stratification was shown to be important. Coastal waters generally show a stronger stratification particularly during neap tides upon which the phytoplankton biomass at the surface rises reaching its maximum about 2–3 days after neap tide. It appeared that porpoises occurred in those areas where stratification is maximised and Risso’s preferred more mixed waters. This fine-scale study provided a temporal insight into spatial distribution of two species that single studies conducted over broader scales (tens or hundreds of kilometers) do not achieve. Understanding which topographic and cyclic variables drive the patchy distribution of porpoises and Risso’s in a Headland/Island system may form the initial basis for identifying potentially critical habitats for these species.     

Ecomorphology of retinal structures in zooplanktivorous haplochromine cichlids (Pisces) from Lake Victoria

Synopsis Before the decline of the species flock of haplochromine cichlids of Lake Victoria due to the Nile perch upsurge, there were many co-existing haploehromine species such as the taxonomically and ecologically well-studied zooplanktivores of the Mwanza Gulf. In spite of the scarcely separated niches of some of these species, no sign of competition for space or food could be demonstrated. As is argued in this paper, optical differentiation could well be an aspect of adaptive radiation of these zooplanktivores, particularly among the highly sympatric species. Our hypothesis is based on the morphological modifications of retinal structures in nine zooplanktivorous species. Interspecific variation was observed in composition, size and density of the photoreceptors and ganglion cells. The analyses included the intraretinal variation and size dependency of some of the structural parameters. The optical functions deduced from retinal structure indicate distinct interspecific differences in sensitivity thresholds and a slight differentiation in visual resolution. These functions correlate poorly with the photic conditions of the species-specific habitats. The optical properties can, on the other hand, be connected with the more subtle differentiation in food items and feeding behaviour among these species. It is our concluding hypothesis, that the optical differentiation among the haplochromine zooplanktivores primarily served resource partitioning by different modes of visual prey detection rather than niche partitioning by habitat.     

The Transparent Goby, Aphia MinutaReview of Biology and Fisheries of a Paedomorphic European Fish

The transparent goby, Aphia minuta, is one of the few pelagic species of the family Gobiidae. Its geographic range covers the eastern Atlantic Ocean, from Gibraltar to Norway and the Mediterranean basin including the Azov and Black seas. However, it does not occur along the north African coasts. The species is characterized by a unique larval morphology and precocious sexual maturation, i.e. a type of heterochrony known as progenesis. The transparent goby is an annual species that lives less than 1 year, with a maximum size of less than 60 mm. The breeding season is quite long and spawning takes place at least twice during its short lifespan. Just after spawning, all breeders quickly die probably as a consequence of the degeneration of the intestinal epithelium by apoptosis, marking the end of the entire cohort. Spending most of its life in the water column, it is a planktotrophic feeder, relying mainly on small copepods. In turn the transparent goby is preyed on by several species of fish and larger cephalopods. During ontogenesis, t hree phases characterize the species: a “pelagic phase”, composed of larval stages hatched from demersal eggs that inhabit shallow coastal waters; an “aggregated phase”, composed of juveniles that gather in schools in shallow waters during winter; finally, a “demersal phase”, composed of adults that in spring migrate offshore with a more dispersed distribution in proximity of the bottom. Despite its small size, this species is largely exploited by local small-scale fisheries developed mainly in the western and central Mediterranean. The fishing fleets, harboured in several sites in Spain and Italy, are generally composed of small vessels. Most of them use very selective gear such as purse-seine nets to catch schools of the transparent goby during winter. For these artisanal fisheries, the transparent goby represents, although seasonally, a very important source of income, yielding locally up to more than 100 tons per fishing season and 75 kg/vessel/day. This review critically examines the published literature on the biology and fisheries of the transparent goby, aiming to provide useful tools for appropriate management and for a sustainable exploitation of this important resource.     

Dissociation of visual associative and motor learning in Drosophila at the flight simulator

Ever since operant conditioning was studied experimentally, the relationship between associative learning and possible motor learning has become controversial. Although motor learning and its underlying neural substrates have been extensively studied in mammals, it is still poorly understood in invertebrates. The visual discriminative avoidance paradigm of Drosophila at the flight simulator has been widely used to study the flies' visual associative learning and related functions, but it has not been used to study the motor learning process. In this study, newly-designed data analysis was employed to examine the flies' solitary behavioural variable that was recorded at the flight simulator-yaw torque. Analysis was conducted to explore torque distributions of both wild-type and mutant flies in conditioning, with the following results: (1) Wild-type Canton-S flies had motor learning performance in conditioning, which was proved by modifications of the animal's behavioural mode in conditioning. (2) Repetition of training improved the motor learning performance of wild-type Canton-S flies. (3) Although mutant dunce(1) flies were defective in visual associative learning, they showed essentially normal motor learning performance in terms of yaw torque distribution in conditioning. Finally, we tentatively proposed that both visual associative learning and motor learning were involved in the visual operant conditioning of Drosophila at the flight simulator, that the two learning forms could be dissociated and they might have different neural bases.     

Cladocera paleocommunity to disentangle the impact of anthropogenic and climatic stressors on a deep subalpine lake ecosystem (Lake Iseo,Italy)

In big lakes with strong anthropogenic pressure, it is usually difficult to disentangle the impacts of climate variability from those driven by eutrophication. The present work aimed at the reconstruction of change in the species distribution and density of subfossil Cladocera in Lake Iseo (Italy) in relation to climate and anthropogenic pressure. We related subfossil Cladocera species composition and density in an 80-cm sediment core collected in the pelagic zone of Lake Iseo to long-term temperature trends and phosphorus concentration inferred by diatoms frustules. The Cladocera remains detected in Lake Iseo sediment reflected the species composition and density of modern pelagic Cladocera assemblages. Cladocera rapidly respond to environmental change, and that climate change combined with eutrophication can induce changes in community composition and species density. At the beginning of twentieth century, when global warming was not yet so accentuated, the nutrient increase in water resulted as the principal driver in determining the long-term development of plankton communities and pelagic food web structure. Moreover, catchment-related processes may decisively affect both species composition and density of the lake planktonic communities due to the decrease of lake water transparency induced by input of inorganic material from the catchment area to the lake. The paleolimnological investigation, through the combined study of biotic and abiotic factor, allowed clarifying the synergic effects of the most important drivers of change in lake ecosystems, suggesting that climatic factors should be considered with nutrient availability as determinant element in controlling the temporal development of plankton communities and pelagic food web structure.

     

Influence of physical environmental factors on the composition and horizontal distribution of summer larval fish assemblages off Mallorca island (Balearic archipelago, western Mediterranean)

Two ichthyoplankton surveys were carried out in two areas offMallorca island (Balearic archipelago, western Mediterranean)in June 1996 and August 1996, respectively. The aim of bothsurveys was to assess the influence of physical environmentalfactors on the horizontal distribution of larval fish assemblages,focusing on larvae of large migratory pelagic fish species.Canonical correspondence analysis (CCA) indicated that in theJune survey, the patterns of horizontal distribution of fishlarvae were mainly conditioned by depth and by the distributionof two surface water masses: the Modified Atlantic Waters (MAW),of recent Atlantic origin, and the older Surface MediterraneanWaters (SMW). The effect of depth gradient was clear in mesopelagicand neritic species, but it was not so evident in the larvaeof large pelagic species which presented a highly patchy distribution.Contrastingly, the August patterns of horizontal fish larvaedistribution were significantly correlated with the surfacesalinity gradient resulting from successive MAW inflows, whereasdepth did not show any significant effect, probably linked tothe bottom topography with a very narrow shelf area. The dataobtained highlight that the Balearic Islands constitute an importantspawning ground for most of the large pelagic fishes inhabitingMediterranean waters, both highly migratory and resident species.     

A REVIEW OF THE PROBLEMATIC OSTEOSTRACAN GENUS AUCHENASPIS AND ITS ROLE IN THYESTIDIAN EVOLUTION

Abstract:  The poorly understood osteostracan genus Auchenaspis , from the Upper Silurian of England, belongs to the thyestidians (Thyestiida), a controversial and important group that is key to the understanding of the evolution of the Osteostraci and paired pectoral fins. The two under-studied species of this genus are comprehensively reviewed using anatomical and novel statistical approaches. No reliable anatomical characters can be used to distinguish between the two species, yet discriminant analysis finds significant separation with respect to size. Potential causes of this conflict are discussed. The findings enable Auchenaspis to be considered in terms of wider thyestidian evolution and contradict previous interpretations of the genus as representing a paraphyletic taxon, whose morphology is intermediate between the general osteostracan condition and the highly derived tremataspids.     

The red and the black: bioluminescence and the color of animals in the deep sea

The colors of deep-sea species are generally assumed to be cryptic,but it is not known how cryptic they are and under what conditions.This study measured the color of approximately 70 deep-sea species,both pelagic and benthic, and compared the results with twosets of predictions: 1) optimal crypsis under ambient light,2) optimal crypsis when viewed by bioluminescent "searchlights."The reflectances of the pelagic species at the blue-green wavelengthsimportant for deep-sea vision were far lower than the predictedreflectances for crypsis under ambient light and closer to thezero reflectance prediction for crypsis under searchlights.This suggests that bioluminescence is more important than ambientlight for the visual detection of pelagic species at mesopelagicdepths. The reflectances of the benthic species were highlyvariable and a relatively poor match to the substrates on whichthey were found. However, estimates of the contrast sensitivityof deep-sea visual systems suggest that even approximate matchesmay be sufficient for crypsis in visually complex benthic habitats.Body coloration was generally uniform, but many crabs had strikingpatterns that may serve to disrupt the outlines of their bodies.     

Longitudinal differentiation among pelagic populations in a planktic foraminifer

Evolutionary processes in marine plankton have been assumed to be dependent on the oceanic circulation system, which transports plankton between populations in marine surface waters. Gene flow facilitated by oceanic currents along longitudinal gradients may efficiently impede genetic differentiation of pelagic populations in the absence of confounding marine environmental effects. However, how responsible oceanic currents are for the geographic distribution and dispersal of plankton is poorly understood. We examined the phylogeography of the planktic foraminifer Pulleniatina obliquiloculata in the Indo-Pacific Warm Pool (IPWP) by using partial small subunit ribosomal DNA (SSU rDNA) sequences. We found longitudinal clines in the frequencies of three distinct genetic types in the IPWP area. These frequencies were correlated with environmental factors that are characteristic of three water masses in the IPWP. Noteworthy, populations inhabiting longitudinally distant water masses at the Pacific and Indian sides of the IPWP were genetically different, despite transportation of individuals via oceanic currents. These results demonstrate that populations of pelagic plankton have diverged genetically among different water masses within a single climate zone. Changes of the oceanic circulation system could have impacted the geographic patterns of dispersal and divergence of pelagic plankton.