The hypoxia avoidance behaviour of juvenile Atlantic cod (<Emphasis Type="Italic">Gadus morhua</Emphasis> L.) depends on the provision and pressure level of an O<Subscript>2</Subscript> refuge

The frequency of low O₂ (hypoxia) has increased in coastal marine areas but how fish avoid deleterious water masses is not yet clear. To assess whether the presence and oxygen pressure (PO₂) level of an O₂ refuge affects the hypoxia avoidance behaviour of fish, individual Atlantic cod (Gadus morhua L.) were exposed to a range of O₂ choices in a 2-way choice chamber at 11.4°C over two different experiments. Cod in the first experiment were allowed access to a fixed O₂ refuge (fully air-saturated seawater) whilst oxygen pressure (PO₂) on the other side was reduced in steps to a critically low level, i.e. 4.3 kPa—a point where cod can no longer regulate O₂ consumption. Under these conditions, cod did not avoid any level of hypoxia and fish swimming speed also remained unchanged. In contrast, strong avoidance reactions were exhibited in a second experiment when fish were again exposed to 4.3 kPa but the safety, i.e. PO₂, of the refuge was reduced. Fish not only spent less time at 4.3 kPa as a result of fewer sampling visits but they also swam at considerably slower speeds. The presence of an avoidance response was thus strongly related to refuge PO₂ and it is unlikely that cod, and possibly other fish species, would enter low O₂ to feed in the wild if a sufficiently safe O₂ refuge was not available. It is therefore hypothesized that the feeding range of fish may be heavily compressed if hypoxia expands and intensifies in future years.     

Carbon conversion and metabolic rate in two marine sponges

The carbon metabolism of two marine sponges, Haliclona oculata and Dysidea avara, has been studied using a ¹³C isotope pulse-chase approach. The sponges were fed ¹³C-labeled diatoms (Skeletonema costatum) for 8 h and they took up between 75 and 85%. At different times, sponges were sampled for total ¹³C enrichment, and fatty acid (FA) composition and ¹³C enrichment. Algal biomarkers present in the sponges were highly labeled after feeding but their labeling levels decreased until none was left 10 days after enrichment. The sponge-specific FAs incorporated ¹³C label already during the first day and the amount of ¹³C label inside these FAs kept increasing until 3 weeks after labeling. The algal-derived carbon captured by the sponges during the 8-h feeding period was thus partly respired and partly metabolized during the weeks following. Apparently, sponges are able to capture enough food during short periods to sustain longer-term metabolism. The change of carbon metabolic rate of fatty acid synthesis due to mechanical damage of sponge tissue was studied by feeding sponges with ¹³C isotope–labeled diatom (Pheaodactylum tricornutum) either after or before damaging and tracing back the ¹³C content in the damaged and healthy tissue. The filtration and respiration in both sponges responded quickly to damage. The rate of respiration in H. oculata reduced immediately after damage, but returned to its initial level after 6 h. The ¹³C data revealed that H. oculata has a higher metabolic rate in the tips where growth occurs compared to the rest of the tissue and that the metabolic rate is increased after damage of the tissue. For D. avara, no differences were found between damaged and non-damaged tissue. However, the filtration rate decreased directly after damage.     

Feeding and non-feeding aggression can be induced in invasive shore crabs by altering food distribution

Human activity can have a large impact on surrounding ecosystems. For example, humans alter resource distributions for other species, potentially modifying these species competitive dynamics. These changes in local competitive processes are frequently associated with species invasions. Here, we investigate how differences in resource distribution affect competitive behaviour using the highly invasive European shore crab (Carcinus maenas). Using a controlled laboratory experiment in combination with behaviour assays and social network analysis, we show that individuals feeding in habitats with clumped food distributions are more aggressive than individuals feeding in habitats where food is evenly dispersed, and this aggression is present even on days where crabs are not feeding. Additionally, this persistent aggression can be induced, suggesting that individuals of this invasive species possess the flexibility to modify their competitive behaviours in response to differences in food distributions. Furthermore, we show how these individual responses can lead to changes in overall organisation of aggressive interactions within a population. We discuss these results in relation to how human impacts can have long-term effects on competitive behavioural strategies, and how behavioural flexibility can allow invasive species to colonise and persist in highly impacted sites such as urban ecosystems.     

Biliverdin-based egg coloration is enhanced by carotenoid supplementation

Avian eggshell color is remarkably variable among and within species and its possible function has long been controversial. Female birds use biliverdin pigment to color their eggs blue and green. Although evidence is accumulating that blue-green eggshell coloration is costly to produce, the exact mechanisms underlying its expression are little studied. Biliverdin pigment is involved in important maintenance functions such as immune and antioxidant defenses and shows similar properties to carotenoid pigments. Carotenoids play a role as immunoenhancers and have the potential to ameliorate the impact of oxidative processes, although their antioxidant function could be minor for some bird species. Important life-history components can be limited by carotenoid availability, including sexual displays and fecundity. Here we explored if biliverdin-based eggshell pigmentation was affected by carotenoid availability in a seabird, by performing a carotenoid supplementation experiment in female blue-footed boobies (Sula nebouxii) after laying their first egg. In this species, blue eggshell color is associated with incubation patterns, and carotenoid availability affects the crucial trade-off between female foot ornamentation and fecundity. We found that under natural conditions, there was a decline in eggshell color with laying order, suggesting pigment limitation for females. However, carotenoid-supplemented females had enhanced second egg coloration compared to controls, and the color increase was independent of their previous pigment allocation to the first egg. Our results suggest that biliverdin-based eggshell coloration is costly to produce and can be alleviated by current carotenoid availability.     

Darwin’s “tug-of-war” vs. starlings’ “horse-racing”: how adaptations for sequential encounters drive simultaneous choice

Charles Darwin aided his private decision making by an explicit deliberation, famously deciding whether or not to marry by creating a list of points in a table with two columns: “Marry” and “Not Marry”. One hundred seventy-two years after Darwin’s wedding, we reconsider whether this process of choice, under which individuals assign values to their options and compare their relative merits at the time of choosing (the tug-of-war model), applies to our experimental animal, the European Starling, Sturnus vulgaris. We contrast this with the sequential choice model that postulates that decision-makers make no comparison between options at the time of choice. According to the latter, behaviour in simultaneous choices reflects adaptations to contexts with sequential encounters, in which the choice is whether to take an opportunity or let it pass. We postulate that, in sequential encounters, the decision-maker assigns (by learning) a subjective value to each option, reflecting its payoff relative to background opportunities. This value is expressed as latency and/or probability to accept each opportunity as opposed to keep searching. In simultaneous encounters, choice occurs through each option being processed independently, by a race between the mechanisms that generate option-specific latencies. We describe these alternative models and review data supporting the predictions of the sequential choice model.     

Ultimate causes and the evolution of altruism

Reconciling the evolution of altruism with Darwinian natural selection is frequently presented as a fundamental problem in biology. In addition to an exponentially increasing literature on specific mechanisms that can permit altruism to evolve, there has been a recent trend to establish general principles to explain altruism in populations undergoing natural selection. This paper reviews and extends one approach to understanding the ultimate causes underlying the evolution of altruism and mechanisms that can realise them, based on the Price equation. From the Price equation, we can see that such ultimate causes equate to the different ways in which the frequency of an altruistic allele in a population can increase. Under this approach, the ultimate causes underlying the evolution of altruism, given some positive fitness costs and benefits, are positive assortment of altruistic alleles with the altruistic behaviour of others, positive deviations from additive fitness effects when multiple altruists interact or bias in the inheritance of altruistic traits. In some cases, one cause can be interpreted in terms of another. The ultimate causes thus identified can be realised by a number of different mechanisms, and to demonstrate its general applicability, I use the Price equation approach to analyse a number of classical mechanisms known to support the evolution of altruism (or cooperation): repeated interaction, ‘greenbeard’ traits, games played on graphs and payoff synergism. I also briefly comment on other important points for the evolution of altruism, such as the ongoing debate over the predominant status of inclusive fitness as the best way to understand its evolution. I conclude by arguing that analysing the evolution of altruism in terms of its ultimate causes is the logical way to approach the problem and that, despite some of its technical limitations, the Price equation approach is a particularly powerful way of doing so.     

AIC model selection and multimodel inference in behavioral ecology: some background,observations, and comparisons

We briefly outline the information-theoretic (I-T) approaches to valid inference including a review of some simple methods for making formal inference from all the hypotheses in the model set (multimodel inference). The I-T approaches can replace the usual t tests and ANOVA tables that are so inferentially limited, but still commonly used. The I-T methods are easy to compute and understand and provide formal measures of the strength of evidence for both the null and alternative hypotheses, given the data. We give an example to highlight the importance of deriving alternative hypotheses and representing these as probability models. Fifteen technical issues are addressed to clarify various points that have appeared incorrectly in the recent literature. We offer several remarks regarding the future of empirical science and data analysis under an I-T framework.     

Stereotypy and variation of the mating call in the Lusitanian toadfish, <Emphasis Type="Italic">Halobatrachus didactylus</Emphasis>

Signal attributes should show different degrees of variability depending on the information to be conveyed. Species identity is usually associated with stereotyped features of a signal, whereas other types of information such as individual quality and motivation are associated with signal plasticity. Lusitanian toadfish males form aggregations during the breeding season and emit a tonal advertisement call (the boatwhistle) to attract mates to their nests. We test the hypothesis that the boatwhistle can convey information both on individual identity and motivation by checking how signal parameters vary with time. We study how the physical (tide level) and social (calling alone or in a chorus) environments and male calling rate affect this advertisement signal and how all these external and internal factors (environment, social and male motivation) blend to modulate the Lusitanian toadfish’s advertisement call. Boatwhistles of each male were very stereotyped in short periods of time (minutes), but intra-male signal variability greatly increased in a longer time scale (days). Nevertheless, significant differences among males could still be found even in a long time scale. Pulse period was the acoustic feature that most contributed to discriminate among males. Tide level and male calling rate modulated boatwhistle characteristics, and there was a differential effect of tide on call attributes depending on male calling rate. Social acoustic environment only affected calling rate. These results suggest that inter-individual differences in call characteristics and call plasticity may mediate both male–male assessment and mate choice.     

Application of a coastal model to simulate present and future inundation and aid coastal management

This paper describes the application of coastal hydro-informatic modelling (using the TELEMAC Modelling System) to address management issues arising from projected hydrodynamical and morphological changes within a shallow, sandy estuarine environment. The model incorporates the complex interaction of ocean, terrestrial and atmospheric processes. The case study of the Dyfi Estuary, on the west coast of Wales, is highlighted here. As sea levels have risen locally and are predicted to rise further, a National Nature Reserve (Borth Bog), which has been reclaimed from tidal waters by embankments, will be at increasing risk from flooding episodes due to overtopping of these embankments at high tide. Present and predicted future tidal-fluvial scenarios have been modelled in the Dyfi Estuary in order to estimate the potential for flooding. In addition, areas of greatest velocity change and potential for sediment erosion/accretion have been identified. A further process that has been investigated is how salt marsh migration is affected by sea-level rise. This case study exemplifies some fundamental and complex physical processes inherent to estuaries, and shows how different management options can be assessed, before their implementation, through a modelling approach.     

Changes in landscape and vegetation of coastal dunes in northwest Europe: a review

In coastal dunes, landscape changes are a rule, rather than an exception. This paper gives an overview of changes in landscape and vegetation with a focus on the past century. The history of dunes is characterised by phases of sand drift, alternated with geomorphological stability. The historical development of dune woodland during these stable phases has been documented for sites all over Europe. Vegetation reconstructions of historical open dune habitats however is very difficult due to limited preservation of fossil remains. People have drastically altered coastal dune landscapes through centuries of exploitation and more recently development of the coast. Historical land use has generally pushed vegetation back into a semi-natural state. During roughly the past century a tendency of increasing fixation and succession is observed on coastal dunes throughout northwest Europe. Six causes of change are discussed. 1) Changes in land use, mainly abandonment of agricultural practices, have led to the development of late successional stages such as scrub and woodland. 2) Crashing rabbit populations due to myxomatosis in the 1950s caused vigorous grass growth and probably stimulated scrub development. 3) A general tendency of landscape fixation is observed due to both natural and anthropogenic factors. 4) Eutrophication, mainly due to atmospheric nitrogen deposition is clearly linked to grass encroachment on acidic but also on some calcareous dunes. 5) The impact of climate change on vegetation is still unclear but probably lengthening of growing season and maybe enhanced CO₂ concentrations have led to an acceleration of succession. 6) A general anthropogenisation of the landscape occurs with rapid spread of non-native species as an important consequence. The reconstruction of a natural reference landscape is considered largely unattainable because of irreversible changes and the long tradition of human impact, in many cases since the development of the dunes. Two contradictory elements need reconciliation. First, the general acceleration of succession and scrub and woodland development in particular is partly caused by a decreased anthropogenic interference in the landscape and deserves more appreciation. Second, most biodiversity values are largely linked to open, early succession dune habitats and are threatened by the same tendency. Apart from internal nature management, in which grazing plays an important part, re-mobilisation of stable, senescent dunes is an important challenge for dune management.