Parasite distribution on client reef fish determines cleaner fish foraging patterns

Recent evidence suggests that cleaner fish Labroides dimidiatus effectively control parasite densities on client reef fish that actively visit them to have parasites and dead or infected tissue removed. These findings support the hypothesis that clients benefit from cleaning, However, they do not show how cleaners reduce the parasite load of their clients. Cleaners could selectively feed on parasites or parasite removal could be a side product of cleaners foraging indifferently on the client surface, resulting in the removal of healthy mucus and scales also. To investigate cleaner fish foraging behaviour, we infected individuals of the surgeon fish Ctenochaetus striatus, with parasitic monogeneans on one body side, while the other body side was parasite free. We then allowed these clients to interact with L, dimidiatus. We found that the duration of interactions depended on parasite load, and that cleaners spent both more time and took more bites per time unit on the infected than on the uninfected side, Our data thus support the idea that parasite abundance determines food patch quality for cleaners. The overall outcome of cleaning interactions is thus likely to benefit the clients.

Learning of foraging skills by fish

This chapter outlines the relationships between a number of key factors that influence learning and memory, and illustrates them by reference to studies on the foraging behaviour of fish. Learning can lead to significant improvements in foraging performance in only a few exposures, and at least some fish species are capable of adjusting their foraging strategy as patterns of patch profitability change. There is also evidence that the memory window for prey varies between fish species, and that this may be a function of environmental predictability. Convergence between behavioural ecology and comparative psychology offers promise in terms of developing more mechanistically realistic foraging models and explaining apparently 'suboptimal' patterns of behaviour. Foraging decisions involve the interplay between several distinct systems of learning and memory, including those that relate to habitat, food patches, prey types, conspecifics and predators. Fish biologists, therefore, face an interesting challenge in developing integrated accounts of fish foraging that explain how cognitive sophistication can help individual animals to deal with the complexity of the ecological context.

Cleaner fish become hosts: a novel form of parasite transmission

Adult bucephalid trematodes (Digenea) generally only occur in piscivorous fish. Within labrid fishes they are very rare, however, we have found them in labrid cleaner fish that feed on the ectoparasites of fish. We surveyed 969 labrid fishes from the tropical Pacific and found bucephalids only in cleaners (Lahroides dimidiatus, L. bicolor, and Bodianus axillaris) and none in piscivores. The prevalences of bucephalids in L. dimidiatus at Lizard Island, Heron Island, Orpheus Island (all on the Great Barrier Reef), New Caledonia, and Moorea (French Polynesia) were 51, 47, 67, 56, and 67%, respectively. All of the L. bicolor examined from Moorea were infected. Bucephalids were highly prevalent in all size classes of L. dimidiatus from Lizard Island. Bucephalids were found in a 1.6-cm long juvenile L. dimidiatus, in which, piscivory is highly unlikely. We examined the literature on the worldwide bucephalid fauna in labrids and all hosts were found to be cleaners (Symphodus tinca, S. mediterraneus, L. dimidiatus, L. bicolor, and Bodianus axillaris) except Notolabrus parilus, whose ecology is unknown. We suggest that cleaners eat bucephalid metacercariae directly from the exterior surface of client fish during cleaning interactions. This is the first evidence of digeneans in the diet of L. dimidiatus, and the first study to show this novel form of parasite transmission where infective stages are eaten as a result of cleaning behaviour. Cleaning-mediated parasite transmission may result in behavioural modification of second intermediate hosts because clients and parasites both benefit from transmission. If the infection is costly to cleaners and acquired during cheating behaviour, then this parasite might regulate mutualism. Alternatively, if infective stages are targeted, infection by these bucephalids may be a negative consequence of an honest foraging strategy.

Parasitic isopod Anilocra apogonae, a drag for the cardinal fish Cheilodipterus quinquelineatus

Cymothoid isopods Anilocra apogonae are regular ectoparasites of the cardinal fish Cheilodipterus quinquelineatus on the Great Barrier Reef. To determine whether this large isopod, attached to the head of the fish, affects the physiology and behaviour of its host, we conducted morphological measurements to obtain a condition index and several laboratory experiments on fish with and without isopods. The condition index did not vary between parasitised and non-parasitised wild fish. However, we found that parasitised fish lost more weight than unparasitised fish when fed a low food ration. Parasitised fish also had a higher rate of oxygen consumption than non-parasitised fish. When maintaining body posture in calm water, parasitised fish had an elevated pectoral fin beat frequency, probably because the isopod attaches asymmetrically, causing an asymmetrical weight balance for which the fish needs to compensate. Moreover, the sustained aerobic swimming speed as well as the swimming endurance at high water speeds were reduced in parasitised fish, possibly because of the drag from the parasite. The results suggest that parasites can have significant effects on fish even if this is not revealed by their body condition index in the wild. The metabolic effects found imply that parasitised fish may have to spend more time foraging to compensate for their higher metabolism. This could expose them to a higher risk of being eaten, a situation made worse by an impaired swimming ability that may reduce their capacity to escape a predator.

Client fish ectoparasite loads and cleaner shrimp Urocaridella sp c hunger levels affect cleaning behaviour

Cleaning is a classic example of mutualism and determining the factors that maintain the balance between the costs and benefits for mutualist partners can assist our understanding of how cleaning relationships are maintained. Optimal foraging theory suggests two factors that might help to maintain the relationship between cleaners and their clients: client ectoparasite load and cleaner hunger levels. The ecological relevance and importance of foraging by cleaner fish in marine systems has been demonstrated repeatedly, yet there is little information available on this behaviour in cleaner shrimp. To determine whether cleaner shrimp base their choice of client fish on food patch quality (i.e. client fish ectoparasite load) we offered the yellow-beaked cleaner shrimp Urocaridella sp. c a choice of parasitized and unparasitized rock cods, Cephalopholis cyanostigma. To determine whether cleaner shrimp hunger levels influence cleaning time, we manipulated hunger levels in Urocaridella sp. c and examined their behaviour towards parasitized client fish. Cleaner shrimp preferred parasitized to unparasitized client fish and food-deprived cleaner shrimp cleaned parasitized rock cods more frequently than satiated cleaner shrimp did. Therefore, variations in client fish ectoparasite load and cleaner shrimp hunger level are two factors that affect the balance in this mutualism. Finally, our results meet some of the assumptions of biological market theory, a framework used to understand cooperative interactions, and thus this framework is suggested for future studies on this cleaning system.

The Genetic Speciation of Archaeological Fish Bone: A Feasibility Study from Southeast Queensland

Current genetic methods enable highly specific identification of DNA from modern fish bone. The applicability of these methods to the identification of archaeological fish bone was investigated through a study of a sample from late Holocene southeast Queensland sites. The resultant overall success rate of 2% indicates that DNA analysis is, as yet, not feasible for identifying fish bone from any given site. Taphonomic issues influencing the potential of genetic identification methods are raised and discussed in light of this result.

Mummy! Ball! Fish!: Why English-learning children produce nouns earlier than verbs

When English-learning children begin using words the majority of their early utterances (around 80%) are nouns. Compared to nouns, there is a paucity of verbs or non-verb relational words, such as 'up' meaning 'pick me up'. The primary explanations to account for these differences in use either argue in support of a 'cognitive account', which claims that verbs entail more cognitive complexity than nouns, or they provide evidence challenging this account. In this paper I propose an additional explanation for children's noun/verb asymmetry. Presenting a 'multi-modal account' of word-learning based on children's gesture and word combinations, I show that at the one-word stage English-learning children use gestures to express verb-like elements which leaves their words free to express noun-like elements.

Patterns of species diversity in the gastrointestinal helminths of a coral reef fish, Epinephelus merra (Serranidae), from French Polynesia and the south Pacific Ocean

Large-scale patterns of species diversity in the gastrointestinal helminth faunas of the coral reef fish Epinephelus merra (Serranidae) were investigated in French Polynesia and the South Pacific Ocean. The richer barrier reef community in French Polynesia supported richer parasite communities in E. merra than that on the fringing reef. While parasite communities among fish from the same archipelago were similar, differences in potential host species and the distance between archipelagos may have contributed to a qualitative difference in parasite communities between archipelagos. Digenean community diversity in coral reef fishes was greater in the western South Pacific, following similar patterns in free-living species. However, overall species diversity of camallanid nematodes of coral reef fishes does not appear to have been similarly affected.

Catecholamine release in heat-stressed Antarctic fish causes proton extrusion by the red cells

Two species of Antarctic fish were stressed by moving them from seawater at -1 degrees C to seawater at 10 degrees C and holding them for a period of 10 min. The active cryopelagic species Pagothenia borchgrevinki maintained heart rate while in the benthic species Trematomus bernacchii there was an increase in heart rate. Blood pressure did not change in either species. Both species released catecholamines into the circulation as a consequence of the stress. P. borchgrevinki released the greater amounts, having mean plasma concentrations of 177 +/- 54 nmol.l(-1) noradrenaline and 263 +/- 131 nmol.l(-1) adrenaline at 10 min. Pla.sma noradrenaline concentrations rose to 47 +/- 14 nmol.l(-1) and adrenaline to 73 +/- 28 nmol.l(-1) in T. bernacchii. Blood from P. borchgrevinki was tonometered in the presence of isoprenaline. A fall in extracellular pH suggests the presence of a Na+/H+ antiporter on the red cell membrane, the first demonstration of this in an Antarctic fish. Treatment with the beta-adrenergic antagonist drug sotalol inhibited swelling of red blood cells taken from temperature-stressed P. borchgrevinki, suggesting that the antiporter responds to endogenous catecholamines.

Studies of evolutionary temperature adaptation: Muscle function and locomotor performance in Antarctic fish

1, Studies of evolutionary temperature adaptation of muscle and locomotor performance in fish are reviewed with a focus on the Antarctic fauna living at subzero temperatures. 2. Only limited data are available to compare the sustained and burst swimming kinematics and performance of Antarctic, temperate and tropical species. Available data indicate that low temperatures limit maximum swimming performance and this is especially evident in fish larvae. 3, In a recent study, muscle performance in the Antarctic rock cod Notothenia coriiceps at 0 degrees C was found to be sufficient to produce maximum velocities during burst swimming that were similar to those seen in the sculpin Myoxocephalus scorpius at 10 degrees C, indicating temperature compensation of muscle and locomotor performance in the Antarctic fish. However, at 15 degrees C, sculpin produce maximum swimming velocities greater than N, coriiceps at 0 degrees C, 4, It is recommended that strict hypothesis-driven investigations using ecologically relevant measures of performance are undertaken to study temperature adaptation in Antarctic fish, Recent detailed phylogenetic analyses of the Antarctic fish fauna and their temperate relatives will allow a stronger experimental approach by helping to separate what is due to adaptation to the cold and what is due to phylogeny alone.

The eyes of deep-sea fish I: Lens pigmentation, tapeta and visual pigments

Deep-sea fish, defined as those living below 200 m, inhabit a most unusual photic environment, being exposed to two sources of visible radiation: very dim downwelling sunlight and bioluminescence, both of which are, in most cases. maximal at wavelengths around 450-500 nm. This paper summarises the reflective properties of the ocular tapeta often found in these animals the pigmentation of their lenses and the absorption characteristics of their visual pigments. Deepsea tapeta usually appear blue to the human observer. reflecting mainly shortwave radiation. However, reflection in other parts of the spectrum is not uncommon and uneven tapetal distribution across the retina is widespread. Perhaps surprisingly, given the fact that they live in a photon limited environment, the lenses of some deep-sea teleosts are bright yellow, absorbing much of the shortwave part of the spectrum. Such lenses contain a variety of biochemically distinct pigments which most likely serve to enhance the visibility of bioluminescent signals. Of the 195 different visual pigments characterised by either detergent extract or microspectrophotometry in the retinae of deep-sea fishes, cn. 87% have peak absorbances within the range 468-494 nm. Modelling shows that this is most likely an adaptation for the detection of bioluminescence. Around 13% of deep-sea fish have retinae containing more than one visual pigment. Of these, we highlight three genera of stomiid dragonfishes, which uniquely produce far red bioluminescence from suborbital photophores. Using a combination of longwave-shifted visual pigments and in one species (Malacosteus niger) a chlorophyll-related photosensitizer. these fish have evolved extreme red sensitivity enabling them to see their own bioluminescence and giving them a private spectral waveband invisible to other inhabitants of the deep-ocean. (C) 1998 Elsevier Science Ltd. All rights reserved.

The eyes of deep-sea fish II. Functional morphology of the retina

Three different aspects of the morphological organisation of deep-sea fish retinae are reviewed: First, questions of general cell biological relevance are addressed with respect to the development and proliferation patterns of photoreceptors, and problems associated with the growth of multibank retinae, and with outer segment renewal are discussed in situations where there is no direct contact between the retinal pigment epithelium and the tips of rod outer segments. The second part deals with the neural portion of the deep-sea fish retina. Cell densities are greatly reduced, yet neurohistochemistry demonstrates that all major neurotransmitters and neuropeptides found in other vertebrate retinae are also present in deep-sea fish. Quantitatively, convergence rates in unspecialised parts of the retina are similar to those in nocturnal mammals. The differentiation of horizontal cells makes it unlikely that species with more than a single visual pigment are capable of colour vision. In the third part. the diversity of deep-sea fish retinae is highlighted. Based on the topography of ganglion cells, species are identified with areae or foveae located in various parts of the retina, giving them a greatly improved spatial resolving power in specific parts of their visual fields. The highest degree of specialisation is found in tubular eyes. This is demonstrated in a case study of the scopelarchid retina, where as many as seven regions with different degrees of differentiation can be distinguished, ranging from an area giganto cellularis, regions with grouped rods to retinal diverticulum. (C) 1998 Elsevier Science Ltd. All rights reserved.

A systemic secondary vessel system is present in the teleost fish Tandanus tandanus and absent in the elasmobranchs Carcharhinus melanopterus and Rhinobutos typus and in the dipnoan Neoceratodus forsteri

A system of secondary vessels emerging from the primary vessels as numerous coiled capillaries has been described in numerous teleost and holost fishes. The systemic secondary vessels of the teleost Tandanus tandanus are typical of this system and are described in this study. The existence of a secondary vessel system has been postulated in the elasmobranch group. No secondary vessel origins, as seen in the teleosts, are present in the elasmobranchs Rhinobatos typus and Carcharhinus melanopterus. Vessels with a similar distribution to secondary arteries are observed but these are venous rather than arterial in nature and do not connect with the primary arteries. Like the secondary veins in teleosts, the cutaneous veins in R. typus contain blood with a low haematocrit. There is no morphological evidence for a secondary vessel system in the dipnoan Neoceratodus forsteri.

Optics of the developing fish eye: comparisons of Matthiessen's ratio and the focal length of the lens in the black bream Acanthopagrus butcheri (Sparidae, Teleostei)

Matthiessen's ratio (distance from centre of lens to retina: lens radius) was measured in developing black bream, Acanthopagrus butcheri (Sparidae, Teleostei). The value decreased over the first 10 days post-hatch from 3.6 to 2.3 along the nasal and from four to 2.6 along temporal axis. Coincidentally, there was a decrease in the focal ratio of the lens (focal length:lens radius). Morphologically, the accommodatory retractor lentis muscle appeared to become functional between 10-12 days post-hatch. The results suggest that a higher focal ratio compensates for the relatively high Matthiessen's ratio brought about by constraints of small eye size during early development. Combined with differences in axial length, this provides a means for larval fish to focus images from different distances prior to the ability to accommodate. (C) 1999 Elsevier Science Ltd. All rights reserved.