God playing dice, revisited: determinism and indeterminism in studies of stochastic phenotypic variation.

Empirical studies of phenotypic variation show that genetic and environmental heterogeneity account for only part of it. Usually, the magnitude of the residual variation is comparable with that of the genetic component, while notably exceeding the magnitude of the environmental component. This can be interpreted in two ways. A deterministic interpretation associates it with artifacts such as measurement error and genetic and environmental heterogeneity that is unaccounted for. An indeterministic interpretation argues that it is random or stochastic phenotypic variation (SPV) resulting from developmental instability - a developing organism's inability to produce a consistent phenotype in a given environment. Classical example of debates between determinists and indeterminists took place about a century ago in quantum physics. In discussing Heidelberg's Uncertainty Principle, Einstein metaphorically expressed his deterministic position: 'God does not play dice with universe'. The indeterministic Uncertainty Principle, however, was eventually widely accepted. Currently, most biologists implicitly or explicitly support deterministic interpretations of phenotypic variation patterns. Here, a wide range of data on morphological traits (studied with analysis of fluctuating asymmetry) and non-morphological traits are analyzed to provide evidence that SPV is not an artifact, but a valid phenomenon. This evidence supports conclusions that observed associations between SPV and stress can be analyzed in the framework of dynamic energy budget theory, and are inextricably linked through energy homeostasis.

Aging three-spined sticklebacks Gasterosteus aculeatus: comparison of estimates from three structures.

In order to assess the accuracy and reliability of age estimates from calcified structures in the three-spined stickleback Gasterosteus aculeatus, we evaluated intra and inter-reader repeatability from three structures: otoliths, gill covers and pelvic spines). Average age estimates were also compared between the structures. The overall intra-reader repeatability of age estimates were highest for otoliths (69%), lowest for gill covers (53%) and intermediate for spine cross-sections (63%). Although four of the seven readers had the highest intra-reader repeatability score for spine cross-sections, the inter-reader variance in this structure was much higher than in others. Otoliths were the easiest in terms of their pre-analysis treatment and exchange of materials (as digital images) between readers. In addition, otoliths are more well-studied compared with the other structures with respect to their development through ontogenesis; hence, age estimates based on otoliths should be the most reliable. Therefore, our recommendation is that whenever possible, analysis of otoliths should be the preferred approach for aging G. aculeatus.

Fluctuating asymmetry as a pollution monitor: The Australian estuarine smooth toadfish Tetractenos glaber (Teleostei: Tetraodontidae).

The relationship between pollution level in estuarine sediment and fluctuating asymmetry (FA) of resident smooth toadfish Tetractenos glaber was evaluated. A total of 188 fish from Sydney and Hawkesbury River estuaries (5 locations from each) were analysed for 28 bilateral skull bone characters. Sediment pollution was quantified based on analysis of heavy metals (Cd, Cr, Cu, Ni, Pb and Zn) and organochlorine pesticides (DDT, DDD, DDE, chlordane, dieldrin, lindane). Sediment toxicity was characterized using the mean quotient approach (MERMQ) and ranged from low to moderate level for heavy metals and from low to severe for organochlorides. The mean shape and directional asymmetry of fish bones differed among locations, suggesting a response to local environments. FA was positively correlated with organochlorine pesticides across locations, but not with heavy metals. These results suggest that fish FA could be a useful estimator of stress caused by organic toxicity based on the MERMQ approach.

Cryptic or pseudocryptic: can morphological methods inform copepod taxonomy? An analysis of publications and a case study of the Eurytemora affinis species complex.

Interest in cryptic species has increased significantly with current progress in genetic methods. The large number of cryptic species suggests that the resolution of traditional morphological techniques may be insufficient for taxonomical research. However, some species now considered to be cryptic may, in fact, be designated pseudocryptic after close morphological examination. Thus the "cryptic or pseudocryptic" dilemma speaks to the resolution of morphological analysis and its utility for identifying species. We address this dilemma first by systematically reviewing data published from 1980 to 2013 on cryptic species of Copepoda and then by performing an in-depth morphological study of the former Eurytemora affinis complex of cryptic species. Analyzing the published data showed that, in 5 of 24 revisions eligible for systematic review, cryptic species assignment was based solely on the genetic variation of forms without detailed morphological analysis to confirm the assignment. Therefore, some newly described cryptic species might be designated pseudocryptic under more detailed morphological analysis as happened with Eurytemora affinis complex. Recent genetic analyses of the complex found high levels of heterogeneity without morphological differences; it is argued to be cryptic. However, next detailed morphological analyses allowed to describe a number of valid species. Our study, using deep statistical analyses usually not applied for new species describing, of this species complex confirmed considerable differences between former cryptic species. In particular, fluctuating asymmetry (FA), the random variation of left and right structures, was significantly different between forms and provided independent information about their status. Our work showed that multivariate statistical approaches, such as principal component analysis, can be powerful techniques for the morphological discrimination of cryptic taxons. Despite increasing cryptic species designations, morphological techniques have great potential in determining copepod taxonomy.

Typology of historical sources and the reconstruction of long-term historical changes of riverine fish: a case study of the Austrian Danube and northern Russian rivers.

Historical data are widely used in river ecology to define reference conditions or to investigate the evolution of aquatic systems. Most studies rely on printed documents from the 19th century, thus missing pre-industrial states and human impacts. This article discusses historical sources that can be used to reconstruct the development of riverine fish communities from the Late Middle Ages until the mid-20th century. Based on the studies of the Austrian Danube and northern Russian rivers, we propose a classification scheme of printed and archival sources and describe their fish ecological contents. Five types of sources were identified using the origin of sources as the first criterion: (i) early scientific surveys, (ii) fishery sources, (iii) fish trading sources, (iv) fish consumption sources and (v) cultural representations of fish. Except for early scientific surveys, all these sources were produced within economic and administrative contexts. They did not aim to report about historical fish communities, but do contain information about commercial fish and their exploitation. All historical data need further analysis for a fish ecological interpretation. Three case studies from the investigated Austrian and Russian rivers demonstrate the use of different source types and underline the necessity for a combination of different sources and a methodology combining different disciplinary approaches. Using a large variety of historical sources to reconstruct the development of past fish ecological conditions can support future river management by going beyond the usual approach of static historical reference conditions.

Coastal fisheries in the Eastern Baltic Sea (Gulf of Finland) and its basin from the 15 to the Early 20th centuries.

The paper describes and analyzes original data, extracted from historical documents and scientific surveys, related to Russian fisheries in the southeastern part of the Gulf of Finland and its inflowing rivers during the 15- early 20(th) centuries. The data allow tracing key trends in fisheries development and in the abundance of major commercial species. In particular, results showed that, over time, the main fishing areas moved from the middle part of rivers downstream towards and onto the coastal sea. Changes in fishing patterns were closely interrelated with changes in the abundance of exploited fish. Anadromous species, such as Atlantic sturgeon, Atlantic salmon, brown trout, whitefish, vimba bream, smelt, lamprey, and catadromous eel were the most important commercial fish in the area because they were abundant, had high commercial value and were easily available for fishing in rivers. Due to intensive exploitation and other human-induced factors, populations of most of these species had declined notably by the early 20(th) century and have now lost commercial significance. The last sturgeon was caught in 1996, and today only smelt and lamprey support small commercial fisheries. According to historical sources, catches of freshwater species such as roach, ide, pike, perch, ruffe and burbot regularly occurred, in some areas exceeding half of the total catch, but they were not as important as migrating fish and no clear trends in abundance are apparent. Of documented marine catch, Baltic herring appeared in the 16(th) century, but did not become commercially significant until the 19(th) century. From then until now herring have been the dominant catch.

Phylogeography of amphi-boreal fish: tracing the history of the Pacific herring Clupea pallasii in North-East European seas.

BACKGROUND: The relationships between North Atlantic and North Pacific faunas through times have been controlled by the variation of hydrographic circumstances in the intervening Arctic Ocean and Bering Strait. We address the history of trans-Arctic connections in a clade of amphi-boreal pelagic fishes using genealogical information from mitochondrial DNA sequence data. The Pacific and Atlantic herrings (Clupea pallasii and C. harengus) have basically vicarious distributions in the two oceans since pre-Pleistocene times. However, remote populations of C. pallasii are also present in the border waters of the North-East Atlantic in Europe. These populations show considerable regional and life history differentiation and have been recognized in subspecies classification. The chronology of the inter-oceanic invasions and genetic basis of the phenotypic structuring however remain unclear. RESULTS: The Atlantic and Pacific herrings both feature high mtDNA diversities (large long-term population sizes) in their native basins, but an ocean-wide homogeneity of C. harengus is contrasted by deep east-west Pacific subdivision within Pacific C. pallasii. The outpost populations of C. pallasii in NE Europe are identified as members of the western Pacific C. pallasii clade, with some retained inter-oceanic haplotype sharing. They have lost diversity in colonization bottlenecks, but have also thereafter accumulated abundant new variation. The data delineate three phylogeographic groups within the European C. pallasii: herring from the inner White Sea; herring from the Mezen and Chesha Bays; and a strongly bottlenecked peripheral population in Balsfjord of the Norwegian Sea. CONCLUSIONS: The NE European outposts of C. pallasii are judged to be early post-glacial colonists from the NW Pacific. A strong regional substructure has evolved since that time, in contrast to the apparent broad-scale uniformity maintained by herrings in their native basins. The structure only partly matches the previous biological concepts based on seasonal breeding stocks or geographical subspecies designations. The trans-Arctic herring phylogeography is notably similar to those of the amphi-boreal mollusk taxa Macoma and Mytilus, suggesting similar histories of inter-oceanic connections. We also considered the time dependency of molecular rates, critical for interpreting timing of relatively recent biogeographical events, by comparing the estimates from coding and non-coding mitochondrial regions of presumably different mutation dynamics.

Direct and indirect climate forcing in a multi-species marine system.

Interactions within and between species complicate quantification of climate effects, by causing indirect, often delayed, effects of climate fluctuations and compensation of mortality. Here we identify direct and indirect climate effects by analysing unique Russian time-series data from the Norwegian Sea-Barents Sea ecosystem on the first life stages of cod, capelin, herring and haddock, their predators, competitors and zooplanktonic prey. By analysing growth and survival from one life stage to the next (eggs-larvae-juveniles-recruits), we find evidence for both bottom-up, direct and top-down effects of climate. Ambient zooplankton biomass predicts survival of all species, whereas ambient temperature mainly affects survival through effects on growth. In warm years, all species experienced improved growth and feeding conditions. Cohorts born following a warm year will, however, experience increased predation and competition because of increased densities of subadult cod and herring, leading to delayed climate effects. While climate thus affects early growth and survival through several mechanisms, only some of the identified mechanisms were found to be significant predictors of population growth. In particular, our findings exemplify that climate impacts are barely propagated to later life stages when density dependence is strong.