An integrated system to assess marine extinctions.

More than 20 global marine extinctions and over 700 local extinctions have reportedly occurred during the past 500 years. However, available methods to determine how many of these species can be confidently declared true disappearances tend to be data-demanding, time-consuming, and not applicable to all taxonomic groups or scales of marine extinctions (global [G] and local [L]). We developed an integrated system to assess marine extinctions (ISAME) that can be applied to any taxonomic group at any geographic scale. We applied the ISAME method to 10 case studies to illustrate the possible ways in which the extinction status of marine species can be categorized as unverified, possibly extinct, or extinct. Of the 10 case studies we assessed, the ISAME method concludes that 6 should be categorized as unverified extinctions due to problems with species' identity and lack of reliable evidence supporting their disappearance (periwinkle-Littoraria flammea [G], houting-Coregonus oxyrinchus [G], long-spined urchin-Diadema antillarum [L], smalltooth sawfish-Pristis pectinata [L], and largetooth sawfish-P. pristis [L]). In contrast, ISAME classified the Guadalupe storm-petrel (Oceanodroma macrodactyla [G]) and the lost shark (Carcharhinus obsolerus [G]) as possibly extinct because the available evidence indicates that their extinction is plausible-while the largetooth sawfish [L] and Steller's sea cow (Hydrodamalis gigas [G]) were confirmed to be extinct. Determining whether a marine population or species is actually extinct or still extant is needed to guide conservation efforts and prevent further biodiversity losses.

Teaching laboratory for large cohorts of undergraduates: Private and social information in fish.

A challenge in the Bachelor's studies in Biology is to strike a balance between reducing the teaching of practical scientific experiments to what is feasible in a short time, and teaching "real" science in undergraduate laboratories for high numbers of participants. We describe a laboratory in behavioral biology, with the primary focus on the student learning. However, also the underlying scientific question and the results of the experiment, namely the behavior of the three-spined stickleback (Gasterosteus aculeatus) in a trade-off situation during foraging, is without a doubt timely and sufficient for scientific studies on this subject, and this through the experiments conducted and data collected by the students. The students rated this laboratory well and learned at the end that social information is certainly important, but that self-learning can be more important, and this not only in small fish, but also for the students themselves.

A new mechanistic approach for the further development of a population with established size bimodality.

Usually, the origin of a within-cohort bimodal size distribution is assumed to be caused by initial size differences or by one discrete period of accelerated growth for one part of the population. The aim of this study was to determine if more continuous pathways exist allowing shifts from the small to the large fraction within a bimodal age-cohort. Therefore, a Eurasian perch population, which had already developed a bimodal size-distribution and had differential resource use of the two size-cohorts, was examined. Results revealed that formation of a bimodal size-distribution can be a continuous process. Perch from the small size-cohort were able to grow into the large size-cohort by feeding on macroinvertebrates not used by their conspecifics. The diet shifts were accompanied by morphological shape changes. Intra-specific competition seemed to trigger the development towards an increasing number of large individuals. A stage-structured matrix model confirmed these assumptions. The fact that bimodality can be a continuous process is important to consider for the understanding of ecological processes and links within ecosystems.

Facing different predators: adaptiveness of behavioral and morphological traits under predation.

Predation is thought to be one of the main structuring forces in animal communities. However, selective predation is often measured on isolated traits in response to a single predatory species, but only rarely are selective forces on several traits quantified or even compared between different predators naturally occurring in the same system. In the present study, we therefore measured behavioral and morphological traits in young-of-the-year Eurasian perch Perca fluviatilis and compared their selective values in response to the 2 most common predators, adult perch and pike Esox lucius. Using mixed effects models and model averaging to analyze our data, we quantified and compared the selectivity of the 2 predators on the different morphological and behavioral traits. We found that selection on the behavioral traits was higher than on morphological traits and perch predators preyed overall more selectively than pike predators. Pike tended to positively select shallow bodied and nonvigilant individuals (i.e. individuals not performing predator inspection). In contrast, perch predators selected mainly for bolder juvenile perch (i.e. individuals spending more time in the open, more active), which was most important. Our results are to the best of our knowledge the first that analyzed behavioral and morphological adaptations of juvenile perch facing 2 different predation strategies. We found that relative specific predation intensity for the divergent traits differed between the predators, providing some additional ideas why juvenile perch display such a high degree of phenotypic plasticity.

Does social context affect boldness in juveniles?

Differences in boldness are common between populations or between related species and are discussed as part of individual coping style, personality, or behavioral syndrome. Boldness has been found to be dependent on experience, social, and environmental contexts. The major aim of the present study was to establish an experimental environment that would allow analyzing the risk-taking behavior of 2 competing invasive goby species. Neogobius melanostomus was more active in the absence of a predator Sander lucioperca than N. fluviatilis and clearly spent more time "swimming" and "feeding" than N. fluviatilis. In addition, N. melanostomus was always faster than N. fluviatilis both when leaving the shelter and reaching offered food. Based on the different behaviors recorded, species-specific boldness scores were established using a principal component analysis. Although there was no overall difference in boldness scores between the 2 species, both competitive conditions and the effect of the predator played significant roles as factors influencing boldness. Neogobius melanostomus was more affected by the presence/absence of the predator than the social circumstances. Neogobius fluviatilis, on the other hand, was more active and bolder in competitive situations. However, when alone, N. fluviatilis was rather inactive and displayed altogether shy behavior, independent of the presence/absence of the predator. Thus, the study confirms the prediction that there are differences in behavior and behavioral plasticity, and therein predator-avoidance strategies, between ecologically similar species of goby living in sympatry. We argue that these differences may be related to differential habitat use of both invasive species that presently dominate the fish community in the Lower Rhine.

Individual consistency and context dependence in group-size preference of Eurasian perch.

Many fish spend a large part of their life in groups. The size of the group influences potential costs and benefits of group living, and depending on context a fish may prefer different group sizes. Group-size preference may also depend on personality, with social individuals expected to prefer larger groups than asocial fish. This study investigates context-dependent group size preference in two populations of a highly social fish, young of the year Eurasian perch. The perch were given a choice between a group of two and a group of eight conspecifics under three different situations: the small group was feeding, the small group had access to shelter, and a control treatment with no extra stimuli. In general, the perch associated more with the large group, but significantly less so during the food treatment. Perceived access to shelter did not affect group size preference compared to the control treatment. Consistent individual differences in social attraction were found within each context, but not among all contexts. Also, an individual's sociability did not correlate with its degree of boldness, indicating a lack of a behavioural syndrome between the two personality traits in the studied populations. The results highlight the importance of considering environmental context when studying social behaviour in obligate social fish, and show the complexity of the concept of sociability as a personality trait by demonstrating context dependence in individual consistency in social behaviour.

Anthropogenic hybridization between endangered migratory and commercially harvested stationary whitefish taxa (Coregonus spp.).

Natural hybridization plays a key role in the process of speciation. However, anthropogenic (human induced) hybridization of historically isolated taxa raises conservation issues. Due to weak barriers to gene flow and the presence of endangered taxa, the whitefish species complex is an excellent study system to investigate the consequences of hybridization in conservation. We focused on three naturally reproductively isolated whitefish taxa in Germany: the endangered, anadromous North Sea houting (NSH) and Baltic houting (BH), which were reintroduced after local extinction, and the commercially stocked European whitefish (EW). To evaluate the genetic integrity of each taxon, source and reintroduced populations of NSH and BH, and EW populations were characterized based on two mitochondrial and 17 microsatellite loci. Additionally, we investigated gill raker counts as an adaptive phenotypic trait. Even though clear genetic and phenotypic differentiation confirmed the houtings as separate evolutionarily significant units, admixture analyses revealed an extensive hybrid zone. Hybridizations were introgressive, positively correlated with genetic diversity, and were reflected in the gill raker counts. The BH distribution range showed higher heterogeneity and stronger admixture than the NSH range. Erroneous stocking with non-native genotypes best explained these patterns, which pose challenges for the conservation of the endangered NSH and BH.

Boldness in two perch populations - long-term differences and the effect of predation pressure.

1. Populations of the same species often display different behaviours, for example, in their response to predators. The question is whether this difference is developed as part of a divergent selection caused by differences in predation pressure, or as a result of phenotypic responses to current environmental conditions. 2. Two populations of Eurasian perch were investigated over a time span of 6 years to see whether risk-taking behaviour in young-of-the-year perch were consistent across cohorts, or if behaviour varied over time with changes in predation regime. 3. Boldness was estimated in aquarium studies by looking at how the fish made trade-offs between foraging in a risky area and staying in shelter. Predation risk of each year and lake was estimated from fishing surveys, using an individual-based model calculating attack rates for cannibalistic perch. 4. The average boldness scores were consistently lower in perch from Fisksjön compared with those in Ängersjön, although the magnitude of the difference varied among years. Variance component analyses showed that differences between lakes in boldness scores only explained 12 per cent of the total variation. Differences between years were contributing at least similarly or more to the total variance, and the variation was higher in Fisksjön than in Ängersjön. 5. The observed risk-taking behaviour of young-of-the-year perch, compared across cohorts, was significantly correlated with the year-specific estimates of cannibalistic attack rates, with lower boldness scores in years with higher predation pressure. In Fisksjön, with significant changes over the years in population structure, the range of both predation risk and boldness scores was wider than in Ängersjön. 6. By following the two perch populations over several years, we have been able to show that the differences in risk-taking behaviour mainly are due to direct phenotypic responses to recent experience of predation risk. Long-term differences in behaviour among perch populations thus reflect consistent differences in predation regime rather than diverging inherent traits.

Match or mismatch: the influence of phenology on size-dependent life history and divergence in population structure.

1. In gape-limited predators, body size asymmetries determine the outcome of predator-prey interactions. Due to ontogenetic changes in body size, the intensity of intra- and interspecific interactions may change rapidly between the match situation of a predator-prey system and the mismatch situation in which competition, including competition with the prey, dominates. 2. Based on a physiologically structured population model using the European perch (Perca fluviatilis), analysis was performed on how prey density (bream, Abramis brama), initial size differences in the young-of-the-year (YOY) age cohort of the predator, and phenology (time-gap in hatching of predator and prey) influence the size structure of the predator cohort. 3. In relation to the seasonality of reproduction, the match situation of the predator-prey system occurred when perch hatched earlier than bream and when no gape-size limitations existed, leading to decreased size divergence in the predator age cohort. Decreased size divergence was also found when bream hatched much earlier than perch, preventing perch predation on bream occurring, which, in turn, increased the competitive interaction of the perch with bream for the common prey, zooplankton; i.e. the mismatch situation in which also the mean size of the age cohort of the predator decreased. 4. In between the total match and the mismatch, however, only the largest individuals of the perch age cohort were able to prey on the bream, while smaller conspecifics got trapped in competition with each other and with bream for zooplankton, leading to enlarged differences in growth that increased size divergence. 5. The modelling results were combined with 7 years of field data in a lake, where large differences in the length-frequency distribution of YOY perch were observed after their first summer. These field data corroborate that phenology and prey density per predator are important mechanisms in determining size differences within the YOY age cohort of the predator. 6. The results demonstrate that the switch between competitive interactions and a predator-prey relationship depended on phenology. This resulted in pronounced size differences in the YOY age cohort, which had far-reaching consequences for the entire predator population.

The annual reproductive cycle of the freshwater mussel Dreissena polymorpha Pallas in lakes.

The annual development of the gonads of Dreissena polymorpha was studied at three sampling sites in two lakes over 3 and 1 1/2 years, respectively. A resting stage occurred after the last spawning in summer/autumn. Oogenesis (accompanied by multiplying segmentation of the oogonia and early growth processes of its oocytes) restarted in specimens at least 1 year old at low temperatures (below 10° C) during winter and early spring. At one location (Fühlinger See) the onset of the spawning season was correlated with an increase of water temperatures above 12° C. At 2 m depth, two main spawning periods in May and August were normally recognized, the first at temperatures of 12-16° C, the second at 16-21° C. It was clearly demonstrated for the first time in Dreissena polymorpha that the oocytes became mature in successive cohorts within one gonad. A female mussel may spawn several times during the reproductive season. At 9 m depth, the onset of spawning also started at about 12° C; this occurred in late summer, with two spawning periods within 1 month at a temperature range of 12-16° C. At another location (Heider Bergsee) the size of the gonads and the oocytes was reduced during April of both years studied, when food supply was low simultaneously with rapidly rising water temperatures in this shallow lake. There was no spawning period during spring. The major spawning period was delayed until July (temperatures 19-22°C). This shows (1) the synchronizing influence of low winter temperatures on the annual reproductive cycle and (2) a temperature threshold of at least 12° C for the start of the spawning processes. The results are discussed with regard to the geographical limits of further spread of Dreissena polymorpha.