Expanding marine protected areas to include degraded coral reefs.

Marine protected areas (MPAs) are a commonly applied solution to coral reef degradation, yet coral reefs continue to decline worldwide. We argue that expanding the range of MPAs to include degraded reefs (DR-MPA) could help reverse this trend. This approach requires new ecological criteria for MPA design, siting, and management. Rather than focusing solely on preserving healthy reefs, our approach focuses on the potential for biodiversity recovery and renewal of ecosystem services. The new criteria would help identify sites with the highest potential for recovery and the greatest resistance to future threats (e.g., increased temperature and acidification) and sites that contribute to MPA connectivity. The DR-MPA approach is a compliment rather than a substitute for traditional MPA design approaches. We believe that the DR-MPA approach can enhance the natural, or restoration-assisted, recovery of DRs and their ecosystem services; increase total reef area available for protection; promote more resilient and better-connected MPA networks; and improve conditions for human communities dependent on MPA ecosystem services.

Light scattering by selected zooplankton from the Gulf of Aqaba.

Light scattering by zooplankton was investigated as a major factor undermining transparency camouflage in these pelagic animals. Zooplankton of differing transparencies--including the hyperiid amphipod Anchylomera blossevillei, an unknown gammarid amphipod species, the brine shrimp Artemia salina, the euphausiid shrimp Euphausia diomedeae, the isopod Gnathia sp., the copepods Pontella karachiensis, Rhincalanus sp. and Sapphirina sp., the chaetognath Sagitta elegans and an enteropneust tornaria larva--were illuminated dorsally with white light (400-700 nm). Spectral measurements of direct transmittance as well as relative scattered radiances at angles of 30 degrees , 90 degrees , 150 degrees and 180 degrees from the light source were taken. The animals sampled had transparencies between 1.5% and 75%. For all species, the highest recorded relative scattered radiance was at 30 degrees , with radiances reaching 38% of the incident radiance for the amphipod A. blossevillei. Scattering patterns were also found to be species-specific for most animals. Relative scattered radiances were used to estimate sighting distances at different depths. These calculations predict that all of the examined zooplankton are brighter than the background radiance when viewed horizontally, or from diagonally above or below at shallow depths. Thus, in contrast to greater depths, the best strategy for detecting transparent zooplankton in the epipelagic environment may be to search for them from above while looking diagonally downwards, looking horizontally or looking from below diagonally upwards. Looking directly upwards proved to be more beneficial than the other viewing angles only when the viewed animal was at depths greater than 40 m.

Can artificial reefs mimic natural reef communities? The roles of structural features and age.

In light of the deteriorating state of coral reefs worldwide, the need to rehabilitate marine environments has greatly increased. Artificial reefs (ARs) have been suggested as a tool for reef conservation and rehabilitation. Although successions of AR communities have been thoroughly studied, current understanding of the interactions between artificial and natural reefs (NRs) is poor and a fundamental question still to be answered is that of whether AR communities can mimic adjacent NR communities. We suggest three alternative hypotheses: Neighboring ARs and NRs will (1) achieve a similar community structure given sufficient time; (2) be similar only if they possess similar structural features; (3) always differ, regardless of age or structural features. We examined these hypotheses by comparing the community structure on a 119-year old shipwreck to a neighboring NR. Fouling organisms, including stony and soft corals, sponges, tunicates, sea anemones and hydrozoans were recorded and measured along belt transects. The ahermatypic stony coral Tubastrea micrantha dominated vertical AR regions while the soft corals Nephthea sp. and Xenia sp. dominated both artificial and natural horizontal surfaces. Our results support the second hypothesis, indicating that even after a century an AR will mimic its adjacent NR communities only if it possesses structural features similar to those of the natural surroundings. However, if the two differ structurally, their communities will remain distinct.

Fouling reefal communities on artificial reefs: does age matter?

Man-made submerged structures, including shipwrecks, offering substrata for fouling organisms and fish, have been classified secondarily as artificial reefs (ARs). The current approach in AR design is that of low-profile structures placed on the seabed and attempting to mimic natural reef (NR) communities with the aim of mitigating degraded marine ecosystems. To examine the validity of this concept, a long-term comparison of the developing AR fouling communities to those of nearby NRs is required. A survey of the fouling reefal organisms was conducted on seven shipwrecks (Red Sea, Egypt), comprising three young (ca 20 years old) and four old (>100 years old) unplanned ARs, in comparison to nearby NR communities. The hypothesis tested was that the age of the ARs shapes the structure of their fouling coral communities. The results demonstrated distinct differences between ARs and NRs and between young and old ARs. While the species composition on ARs may resemble that of NRs after approximately 20 years, obtaining a similar extent of coral cover may require a full century. Moreover, differences in structural features between ARs and NRs may lead to differences in species composition that persist even after 100 years.

Migrating locusts can detect polarized reflections to avoid flying over the sea.

The desert locust Schistocerca gregaria is a well known migrating insect, travelling long distances in swarms containing millions of individuals. During November 2004, such a locust swarm reached the northern coast of the Gulf of Aqaba, coming from the Sinai desert towards the southeast. Upon reaching the coast, they avoided flying over the water, and instead flew north along the coast. Only after passing the tip of the gulf did they turn east again. Experiments with tethered locusts showed that they avoided flying over a light-reflecting mirror, and when given a choice of a non-polarizing reflecting surface and a surface that reflected linearly polarized light, they preferred to fly over the former. Our results suggest that locusts can detect the polarized reflections of bodies of water and avoid crossing them; at least when flying at low altitudes, they can therefore avoid flying over these dangerous areas.

Modular organization of adaptive colouration in flounder and cuttlefish revealed by independent component analysis.

Flounders and cuttlefish have an impressive ability to change colouration, for camouflage and, in the case of cuttlefish, for communication. We pursue the hypothesis that these diverse patterns are created by combining a small number of distinct pattern modules. Independent component analysis (ICA) is a powerful tool for identifying independent sources of variation in linear mixtures of signals. Two versions of ICA are used, one assuming that sources have independence over time, and the other over space. These reveal the modularity of the skin colouration system, and suggest how the pattern modules are combined in specific behavioural contexts. ICA may therefore be a useful tool for studying animal camouflage and communication.

The linearly polarized light field in clear, tropical marine waters: spatial and temporal variation of light intensity, degree of polarization and e-vector angle.

Sensitivity to polarized light is widespread among marine animals, including crustaceans, cephalopods and some fishes. They use this ability to orient and find prey, and possibly for a number of other visual tasks. Unlike the ultraviolet-sensitive polarization receptors of most insects, the polarization receptors of marine invertebrates tend to be maximally sensitive near 500 nm, suggesting that polarized light in water differs from that in air. The underwater field of partially linearly polarized light has been studied for nearly 50 years, but data are still limited and sparse. We measured the submarine polarized light field from 350 to 600 nm throughout the day on a coral reef in the Florida Keys at a depth of 15m using the underwater laboratory Aquarius as a research platform. Our results show that the angle of polarization as viewed along any given line of sight at this depth is a relatively simple function of solar position and that the degree of polarization is greatest 60-90 degrees from the sun. Both e-vector angle and degree of polarization vary only slightly with wavelength, although light is sometimes less polarized in the ultraviolet. Since light is most intense at medium wavelengths and polarization is nearly maximal at these wavelengths, invertebrate polarization photoreceptors are spectrally well placed. Also, the relative spectral constancy of the angle and degree of polarization supports fish polarization sensitivity, which relies on spectrally diverse photoreceptor sets.

Cuttlefish use polarization sensitivity in predation on silvery fish.

Cephalopods are sensitive to the linear polarization characteristics of light. To examine if this polarization sensitivity plays a role in the predatory behavior of cuttlefish, we examined the preference of Sepia officinalis when presented with fish whose polarization reflection was greatly reduced versus fish whose polarization reflection was not affected. Cuttlefish preyed preferably on fish with normal polarization reflection over fish that did not reflect linearly polarized light (n = 24, chi 2 = 17.3, P < 0.0001), implying that polarization sensitivity is used during predation. We suggest that polarization vision is used to break the countershading camouflage of light-reflecting silvery fish.

An ethogram of body patterning behavior in the biomedically and commercially valuable squid Loligo pealei off Cape Cod, Massachusetts.

Squids have a wide repertoire of body patterns; these patterns contain visual signals assembled from a highly diverse inventory of chromatic, postural, and locomotor components. The chromatic components reflect the activity of dermal chromatophore organs that, like the postural and locomotor muscles, are controlled directly from the central nervous system. Because a thorough knowledge of body patterns is fundamental to an understanding of squid behavior, we have compiled and described an ethogram (a catalog of body patterns and associated behaviors) for Loligo pealei. Observations of this species were made over a period of three years (> or = 440 h) and under a variety of behavioral circumstances. The natural behavior of the squid was filmed on spawning grounds off Cape Cod (northwestern Atlantic), and behavioral trials in the laboratory were run in large tanks. The body pattern components--34 chromatic (including 4 polarization components), 5 postural, and 12 locomotor--are each described in detail. Eleven of the most common body patterns are also described. Four of them are chronic, or long-lasting, patterns for crypsis; an example is Banded Bottom Sitting, which produces disruptive coloration against the substrate. The remaining seven patterns are acute; they are mostly used in intraspecific communication among spawning squids. Two of these acute patterns--Lateral Display and Mate Guarding Pattern--are used during agonistic bouts and mate guarding; they are visually bright and conspicuous, which may subject the squids to predation; but we hypothesize that schooling and diurnal activity may offset the disadvantage presented by increased visibility to predators. The rapid changeability and the diversity of body patterns used for crypsis and communication are discussed in the context of the behavioral ecology of this species.

Behavioural evidence for polarisation vision in stomatopods reveals a potential channel for communication.

Polarisation sensitivity (PS) - the ability to detect the orientation of polarised light - occurs in a wide variety of invertebrates [1] [2] and vertebrates [3] [4] [5], many of which are marine species [1]. Of these, the crustacea are particularly well documented in terms of their structural [6] and neural [7] [8] adaptations for PS. The few behavioural studies conducted on crustaceans demonstrate orientation to, or local navigation with, polarised sky patterns [9]. Aside from this, the function of PS in crustaceans, and indeed in most animals, remains obscure. Where PS can be shown to allow perception of polarised light as a 'special sensory quality' [1], separate from intensity or colour, it has been termed polarisation vision (PV). Here, within the remarkable visual system of the stomatopod crustaceans (mantis shrimps) [10], we provide the first demonstration of PV in the crustacea and the first convincing evidence for learning the orientation of polarised light in any animal. Using new polarimetric [11] and photographic methods to examine stomatopods, we found striking patterns of polarisation on their antennae and telson, suggesting that one function of PV in stomatopods may be communication [12]. PV may also be used for tasks such as navigation [5] [9] [13], location of reflective water surfaces [14] and contrast enhancement [1] [15] [16] [17] [18]. It is possible that the stomatopod PV system also contributes to some of these functions.

Polarization contrast vision in Octopus.

While the ability to analyze polarized light is widespread among animals, its contribution to form vision has not yet been documented. We tested the hypothesis that polarization vision can be used for object discrimination, by training octopuses to distinguish between targets on the basis of the presence or absence of a pattern produced by a 90 degrees polarization contrast within the target. Octopuses recognized a 90 degrees contrast pattern within a single target, when presented either on a horizontal/vertical axis or on a 45 degrees/135 degrees axis. They were able to transfer their learning to new situations and to detect a polarization contrast when the orientations of the e-vector of light passing through the target center and background differed by as little as 20 degrees. Polarization vision may provide information similar to that available from color vision and thus serve to enhance the detection and recognition of objects.

Polarization vision in cuttlefish in a concealed communication channel?

Polarization sensitivity is well documented in marine animals, but its function is not yet well understood. Of the cephalopods, squid and octopus are known to be sensitive to the orientation of polarization of incoming light. This sensitivity arises from the orthogonal orientation of neighboring photoreceptors. Electron microscopical examination of the retina of the cuttlefish Sepia officinalis L. revealed the same orthogonal structure, suggesting that cuttlefish are also sensitive to linearly polarized light. Viewing cuttlefish through an imaging polarized light analyzer revealed a prominent polarization pattern on the arms, around the eyes and on the forehead of the animals. The polarization pattern disappeared when individuals lay camouflaged on the bottom and also during extreme aggression display, attacks on prey, copulation and egg-laying behavior in females. In behavioral experiments, the responses of cuttlefish to their images reflected from a mirror changed when the polarization patterns of the reflected images were distorted. These results suggest that cuttlefish use polarization vision and display for intraspecific recognition and communication.

Specialization of retinal function in the compound eyes of mantis shrimps.

Visual function and its specialization at the level of the retina were studied in 13 species of stomatopod crustaceans, representing three superfamilies: Gonodactyloidea, Lysiosquilloidea, and Squilloidea. We measured attenuation and irradiance spectra in the environment of each species, at the actual depths and times of activity where we observed individuals. We also characterized the intrahabdomal filters of all study species and determined the absolute spectral sensitivity functions and approximate photon capture rates of all photoreceptor classes below the level of the 8th retinular cell in seven of these species. Shallow-water gonodactyloid species have four distinct classes of intrarhabdomal filters, producing photoreceptors that are relatively insensitive but which have the broadest spectral coverage of all. Deep-water gonodactyloids and all lysiosquilloids have filters that are spectrally less diverse. These species often discard the proximal filter classes of one or more receptor types. As a result, their retinas are more sensitive but have reduced spectral range or diversity. The single squilloid species has the most sensitive photoreceptors of any we observed, due to the lack both of intrarhabdomal filters and tiered photoreceptors. Photon absorption rates, at the times of animal activity, were similar in most photoreceptor classes of all species, whether the receptors were tiered or untiered, or filtered or unfiltered. Thus, the retinas of stomatopods are specialized to operate at similar levels of stimulation at the times and depths of actual use, while evidently maintaining the greatest possible potential for spectral coverage and discrimination.

Extreme Diel Fluctuations of Oxygen in Diffusive Boundary Layers Surrounding Stony Corals.

The diffusive boundary layers surrounding sessile marine organisms have been implicated in controlling an organism's metabolism and growth. We studied boundary layers surrounding hermatypic corals by monitoring oxygen concentrations on a submillimetric scale. Oxygen concentration within the boundary layers varied from supersaturation during the day to anoxia at night, although the ambient water composition remained constant. Detailed mapping and oxygen measurements revealed diel oxygen fluctuations from supersaturation (373% air saturation) in the light to complete oxygen depletion at darkness in the massive coral Favia favus. Exposure to a 5-cm/s current reduced the boundary layer thickness from 2.44 mm to 1.90 mm, allowing more rapid oxygen exchange across the diffusive boundary layer. Similar patterns were found in the branching coral Stylophora pistillata. In massive corals, the thickness of the diffusive boundary layer was negatively correlated with the size of the polyp. We suggest that the distribution of corals in areas of differential turbulence is related to the thickness of the diffusive boundary layers surrounding them.