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The effect of aerial drying of biofilmed surfaces to simulate a tidal emersion upon the settlement preferences of spirorbid and bryozoan larvae was investigated using choice experiments with still water conditions carried out in the laboratory. Aerial drying of biofilmed slates and pieces of Fucus serratus for 1 h at 20°C negated their usual settlement-inducing properties to larvae of Spirorbis spirorbis, S. tridentatus and Flustrellidra hispida. Such larval settlement preference may contribute to observed variations in the natural distribution of these species in the intertidal.
Stable carbon isotope measurements (δ13C) were used to assess the sources of carbon assimilated by the fan mussel Pinna nobilis, in sea grass Posidonia oceanica meadows, and an associated shrimp Pontonia pinnophylax which occurs within this bivalve's mantle cavity. The primary carbon sources available to both animals displayed a wide range of δ13C values, from −12·3 to −22·3‰. The δ13C and δ15N of Pinna nobilis and Pontonia pinnophylax suggest that they assimilate carbon from similar sources, occupy comparable trophic levels and that their association is commensal.
The distribution and spatio–temporal variation of Helicolenus dactylopterus (Pisces: Scorpaenidae) population was studied from 816 bottom trawls developed along the Iberian coast. Multifactor analysis of variance was used to test the differences in abundance and biomass and cluster analysis was applied to detect differences in population structure. The results showed a bathymetric and latitudinal gradient in abundance and population structure along three geographic sectors in the surveyed area. In the southern area, the species was more frequent and showed a wider bathymetric distribution range than northwards. The depth range with the maximum frequency of occurrence was also wider in the southern area than northwards. Similar trends were observed in terms of abundance and biomass, with the highest and lowest indices in the southern and northern areas, respectively. The age composition of the catches showed that the population consisted mainly of young-of-the-year and juvenile fish. This fraction of the population is well represented, at a mesoscale, along the whole surveyed area, but adults are well represented only at a local scale, at the deepest strata surveyed in the southern area. In this case, recruits of age 0 and juveniles up to 4-y old were restricted to depths shallower than 500 m, while adult fish older than 6 y of age appeared below this depth. In the southern area, seasonal changes in the population structure were also observed, with modes of small fish (3–6 cm) from March to June, as a consequence of the species recruitment to the bottom. Some direct and indirect factors of biogeographic, environmental and anthropogenic origin affecting the observed gradient are discussed.
Geostatistical analysis was used to investigate the effect of drift kelp on the spatial distribution of the sea urchin Tetrapygus niger. The positions of all sea urchins were mapped in four experimental plots in the rocky intertidal zone of the central Chilean coast. When drift macroalgae were added, the sea urchins left the substratum irregularities, increased in number inside the experimental plots, and tended to form a dense aggregation around the kelp. After the drift macroalgae was removed, the aggregations disappeared and the sea urchins returned to the depressions and/or interstices of the substratum. The results suggest that the influx of drift kelp is the triggering factor in the formation of dense aggregations of this species.
The anatomy of the hook apparatus that attaches egg strings to the fish parasites Haemobaphes cyclopterina, Lernaeocera branchialis, Lernaeocera lusci, Lernaeenicus sprattae, Sarcotretes scopeli and Pennella balaenoptera (Copepoda: Pennellida) is described and illustrated. The hook rises from a cupulate base, extending posteriorly and anteriorly in the body cavity. The suspension of the apparatus in the trunk of the different species differs, but the function is similar. The hook tip enters the genital antrum, nearly penetrates the proximal end of the egg string, and continues into a notch on the antrum wall. The apex of the egg string acquires a concave depression like the finger end of a glove. In this way the string is mechanically attached inside the female genital segment. The mobile ectoparasites Lepeophtheirus salmonis and Hatschekia hippoglossi have hooks which function similarly, but perforate the strings.
Enhydrosoma curvirostre is redescribed from archived and new material with particular attention to the structure of the mouthparts and other details of the fine structure not previously reported. It is shown that the species cannot be maintained in Enhydrosoma but is more closely related to Cletodes, the genus in which the original author placed it. However, it is shown that the species must be assigned to a new genus, Spinapecruris, as a result of possessing autapomorphies on the maxillule (only seven setae on the basis), thoracopods 2–4 (a large spine on the distal margin) and possibly the caudal ramus (proximal insertion of seta VII) and lacking apomorphies which may define Cletodes.
Microphytobenthos are significant primary producers in many coastal systems. It is therefore important to quantify their biomass and productivity. Chlorophyll-a is often used as an index for microphytobenthic biomass. However, complications arise as most studies of sediment properties have been on a millimetre scale, whilst chemical and biological gradients in the surface layers of sediment occur over a microscale. The development of a new technique, the Cryolander (Wiltshire et al., 1997; Wiltshire, 2000), now allows microscale analysis of the sediment surface. Areas of high and low diatom biomass were compared using two coring techniques of different vertical resolution; the Cryolander method, with a vertical resolution of 0·2 mm and plastic core tubes (coarse coring), with a vertical resolution of 5 mm. Results indicated that, except at extreme biomass levels, coarse coring does not detect statistically significant differences in chlorophyll-a between obviously diverse sample sites. This may lead to misinterpretation of seasonal and spatial data when coarse coring is used. Furthermore microscale sectioning allows distinctions to be made between chlorophyll-a measured in the photic zone (photosynthetically active biomass (PAB)) and chlorophyll-a measured below the photic zone (photosynthetically inactive biomass (PIB)), allowing accurate determination of biomass specific primary production.
This is the first report on the presence of basking sharks Cetorhinus maximus in Galician waters (north-west Iberian Peninsula) from sightings, strandings and incidental catches by fishing gear. Morphometric, biological, geographic and temporal data on 19 basking sharks recorded from 1988 to 1998 are presented. Average total length is 401·4 cm. Male:female ratio is 0·6. Seventy per cent of records are from incidental fishing catches in bottom gill nets. Approximately 74% of sharks were recorded during February, March and April, which may suggest that the species occurs seasonally in this area.
There is sometimes a significant bias in the sex ratio of fish caught by longline. Usually, more females than males are caught. The possible reasons for unequal sex ratios in longline catches are listed and discussed. One sex could be more common in the area where the fishery takes place because there really is an unequal sex ratio in the population or because the other sex preferentially occurs in different places. Alternatively, longline fishery might preferentially catch one of the sexes. This could be a result of size difference between the sexes and thus a different response to the given hook size or bait size. Finally, sexes could differ in their feeding behaviour. There is growing evidence that females—not only of fish—are ‘energy maximizers’: they find food faster and spend more time feeding than do males. Thus, fishing methods using bait are likely to catch a higher proportion of females than fishing methods that do not use bait.
The purpose of the present study was to assess the relationship between sediment composition and biological community structure in mixed sands and gravel deposits of the eastern English Channel. Although some species are clearly associated with particular sediment types, the results confirm the lack of correspondence between community composition of the benthos and particle size distribution in unconsolidated sand and gravel deposits. The results also suggest that sample-to-sample variability commonly recorded in the species composition of macrofauna may reflect significant under-sampling by conventional grab sampling techniques. The implications of this for environmental monitoring and impact studies is discussed.
Arcachon Bay was the last important shell culture area of the French Atlantic coast without a stock assessment of the accidentally introduced Crepidula fornicata (Gastropoda). Following a stratified sampling strategy, 205 stations were sampled with a dredge. The total biomass of C. fornicata was estimated at 155 tn (confidence limits: 72 tn) scattered on 2·4 km2, i.e. about 5% of the infralittoral area. This biomass is very low compared to other exploited shellfish sites. Amongst mean environmental parameters (such as bathymetry, temperature, salinity, sediment grain-size, anthropic activity), the major keyfactors differentiating Arcachon Bay from other French sites were searched to explain the differences in C. fornicata colonization pattern. It seems that the low biomass of C. fornicata, 30 y after its occurrence was first recorded in the bay, could be correlated to: (1) the scarcity of subtidal muddy sediments which are suitable for C. fornicata settlement; (2) the presence of large Zostera spp. beds on both intertidal and subtidal areas; and (3) the absence of bottom trawl fishing. Growth, production and productivity (P/B) ratios were compared between five stations characterized by different hydrological and sediment parameters. Growth and productivity (0·3 y−1) were not affected by mean biomass suggesting a lack of population regulation by intraspecific competition.
Parental care in birds and mammals is so familiar to all of us that it seems unlikely that it can hold any fresh surprises or offer any new insights. However, there are important aspects of parental care that are commonly overlooked when its role in evolution is discussed. Parental care is one of the major routes through which information is transferred across generations. It is largely through the effects of parental care that animal traditions become established. The information transmitted through parental care relates to all the aspects of life; some is used everyday, some only rarely. Information is transmitted through several different but usually interacting channels, and is essential for the survival and reproduction of the offspring. A look at some typical parental behaviour, that of the common domestic mouse, will show the remarkable range and importance of the information that is transmitted from parent to offspring.
Dusk is a good feeding time for village mice. The small, four-month-old, greyishbrown female domestic mouse silently scales the outer wall of the village grocer's warehouse. She enters the warehouse through a small crack in the wall, and quickly slides down to the piles of bags containing pinhead oatmeal and canary seed. This urine-marked route leads safely to the best source of solid food around. It was first introduced to her by her mother, three months ago, and has been used by her ever since, at least twice a day, at dawn and dusk. […]
To understand how traditions originate and how they evolve, we must first establish the relationship between learning, memory and social organisation. Not everything that is learnt becomes a habit, not every habit involves social interactions and not every social habit is transmitted across generations. We therefore need to know what learning entails, how patterns of behaviour are memorised and how they lead to the formation of traditions. Our purpose is not to describe the neural mechanisms of learning and memory, but rather to outline the psychological, ecological and social conditions that influence how behaviour patterns are generated, remembered and transmitted. ‘Learning’ and ‘remembering’ are not simple and unitary processes, however: different species rely to varying extents on several types of learning and memory. This affects the nature of the habits they develop, and whether or not and in what manner these habits form cultural traditions. To get a better understanding of the different types of learning and their consequences, we will return again to the Judaean hills and observe the behaviour of some of their inhabitants.
It is late spring, and the dry shrubland of the Judaean hills, with its small oak trees dotted among scattered low bushes and wild herbs, is swarming with life. As the daylight fades away, a female orb-web spider, suspended in mid-air on a thin thread stretched between two flowering bushes, is busy constructing her orb-web. Seed-collecting harvester ants move hurriedly along well-trodden earth roads to and from a nearby underground nest. […]
This book is about the way in which the evolution of birds and mammals is affected by social learning and by the traditions formed by social learning. From observation and experiment, we know that higher animals can acquire information from or through the behaviour of others, and through their own behaviour they can transmit this information to the next generation. Variations in such socially acquired and transmitted behaviour-influencing information cannot be under direct genetic control, since animals with very similar genes can have, and pass on, very different behaviours and traditions. There is clearly another inheritance system, a behavioural system of information transmission, which is superimposed on the genetic system. Some years ago we decided that the evolutionary consequences of this additional tier of variation and inheritance were worth exploring, and set out to see how our view of the evolution of higher animals is altered by incorporating non-genetic behavioural inheritance and the traditions that it produces. This book is the outcome of that endeavour.
We found that adding the behavioural system of information transmission has some radical implications for the current gene-centred view of evolution. For example, the classical distinctions between development and evolution become very blurred. An animal tradition is the product of a historical, evolutionary process, yet it can be formed and transmitted only if it is actively constructed during the behavioural development of individuals and groups.
If you ask a biologist to explain the evolution of the elaborate morning song of a great tit, the subtle food preferences of a domestic mouse, or the efficient hunting techniques of a pack of wolves, what sort of explanation will you get? The chances are you will be told that this type of behaviour can readily be explained by the conventional theory of natural selection acting on genetic differences between individuals. Ever since Darwin, the theory of natural selection has been applied to all sorts of biological problems, from the origin of life to the origin of language, and for most of this century it has been assumed that genetic differences between individuals underlie the variation on which natural selection acts. It is not surprising, therefore, that behavioural evolution is also seen as the outcome of the selection of genetic variations. But is this view correct? In this book we are going to argue that when applied to the behaviour of higher animals, conventional evolutionary theory is rarely adequate and is often misleading. Natural selection acting on genetic differences between individuals is not a sufficient explanation for the evolution of the behaviour of the great tit, the mouse or the wolf.
To understand why we are not satisfied with the current application of Darwin's theory to behaviour, we need to go back to basics. Darwin's theory depends on some fundamental properties of biological entities: on their ability to reproduce, on the differences between individuals and on the heritable nature of some of these differences.
In The Descent of Man and The Expression of the Emotions in Man and Animals, Darwin argues for evolutionary continuity between the minds of man and higher animals, stressing that higher animals share with us many complex mental capacities:
the difference in mind between man and the higher animals, great as it is, is certainly one of degree and not of kind. We have seen that the senses and intuitions, the various emotions and faculties, such as love, memory, attention, curiosity, imitation, reason, &c., of which man boasts, may be found in an incipient, or even sometimes a well-developed condition, in lower animals.
(Darwin, 1871, p. 105)
In one form or another, the continuity thesis is accepted by all evolutionary biologists. Even when a large mental gap between the minds of animals and man is recognised, the interpretation of this gap is based on the assumption that there is an underlying genetic and evolutionary continuity. However, notice how Darwin framed his statement: he did not claim that we are psychologically and cognitively simpler than we believe we are – that we are psychologically more like ‘lower’ animals. On the contrary, Darwin believed that ‘lower’ animals are more complex than is usually thought – that they are more similar to us, possessing more sophisticated capacities than we usually grant them.
In this book, we have followed Darwin's approach, emphasising the learning capacities of higher animals, particularly their ability to learn from others.