Characterising functional trait diversity and trait–environment relationships in fish assemblages of boreal lakes

1. Characterisation of biodiversity is typically based on taxonomic approaches, while much less is known about other related aspects. Functional trait diversity is one such component of biodiversity that has not been addressed rigorously in ecological research until recently. We tested the congruence between taxonomic‐ and trait‐based approaches, and examined how spatial configuration, local abiotic environmental factors and biotic effects interact to influence taxonomic‐ and trait‐based characterisation of freshwater fish assemblages. 2. Fish assemblage data were compiled for 124 lakes in southern Finland. Variance partitioning in both linear regression analyses and redundancy analysis was used to quantify the relative contribution of spatial and environmental variables to taxonomic and functional trait diversity and structure. Additionally, a null model analysis was used to test for the potential effects of interspecific segregation and biotic interactions on the co‐occurrence of species. 3. The species pool was relatively poor. However, trait‐based classification of species indicated that most species belonged to unique functional entities, which suggested low redundancy in species composition. Correlation analysis indicated a very strong relationship between species richness (SR) and the number of unique trait combinations (UTC). Ecoregion‐level heterogeneity in SR and UTC were well represented in a relatively small group of randomly selected lakes (c. 30 lakes). Multiple regressions indicated moderate roles for abiotic environmental variables (i.e. lake surface area, depth, total phosphorous, colour and pH) in determining SR, UTC and the distribution of single trait categories, whereas geographical location was not generally influential. 4. Redundancy analysis revealed similar patterns to those of diversity analyses for taxonomic and associated trait‐based structure, emphasising the effect of abiotic environmental variables and the negligible effect of geographical position. 5. Co‐occurrence analysis indicated significant checkerboard distribution at the whole assemblage level, but interspecific segregation proved to be of relatively minor importance in the constrained analyses, where species pair combinations within trait category groups were evaluated. 6. Our results suggest that taxonomic‐ and trait‐based patterns of boreal lake fish assemblages are strongly interrelated. Environmental filtering through the effects of local abiotic variables seems to have the most prominent role in determining trait‐based assemblage patterns among lakes, which may also be secondarily shaped by biotic interactions. 7. From the applied perspective, it may not necessarily matter whether traditional taxonomic or more novel trait‐based approaches are used in characterising spatial patterns in boreal fish assemblages. However, trait‐based approaches may provide complementary information which cannot be directly revealed by taxonomic data.     

Size dependency in colour patterns of Western Palearctic carabids

Body colouration is of high evolutionary relevance for most animals. Several competing hypotheses exist regarding the evolutionary reasons for animal colouration ranging from predator avoidance and sexual advertisement to neutral selection. Among these hypotheses, biophysical principles suggest the thermoregulatory importance of dark colouration which in turn strongly depends on species body size. This body size – darkness trade‐off is based on sound theoretical background conceptualized in the thermal melanism hypothesis and is confirmed by numerous case studies for individual species. However, evidence for the general relevance of this trade‐off on large spatial and taxonomic scale is still missing. Here we specifically focus on this body size – colouration trade‐off for a hyper‐diverse and cosmopolitan group of insects, namely ground beetles. We combined colour information with trait data and distributional as well as bioclimatic attributes for more than 1000 carabid species from the entire Western Palearctic. We quantified species‐specific body colouration from high‐quality, standardised digital photographs using the Munsell colour system. We detect a strong increase of colour darkness with body size from small to medium‐sized carabids up to a body size threshold of 15 mm which is consistent with the thermal melanism hypothesis. However, body size showed no effect above this threshold and colour darkness remained constantly high which is in accordance with previous ideas about the size‐dependency of thermoregulative control mechanisms (size dependence hypothesis). By demonstrating a strong tendency towards darkness with increasing body size, we illustrate the inter‐specific relevance of body colouration for this cosmopolitan group of ectotherms on a continental scale. The putative thermoregulative trade‐off between body size and melanism seems to be of significant importance for carabids on a broad spatial scale and may be a general but still underestimated phenomenon for ectotherms in general, although other mechanistic drivers cannot be completely neglected.     

Assessing lake typologies and indicator fish species for Italian natural lakes using past fish richness and assemblages

In order to establish a fish-based typology of Italian lakes and identify possible reference and indicator fish species for each lake type, we analysed historical data on fish assemblages of all Italian natural lakes >0.5 km² from the period prior to the major decline in water quality in the 1950s. General linear regression models showed the ecoregion and lake altitude being the best predictors of fish species richness. The number of species was significantly higher in the Alpine than in the Mediterranean ecoregion. Among Alpine lakes, the number of fish species increased significantly with lake volume whilst decreased with altitude. In the Mediterranean lakes, none of the selected parameters was significant. Cluster analysis of fish assemblages (presence/absence) divided the lakes of the Alpine and Mediterranean ecoregions into four and two types, respectively. Pike (Esox lucius), rudd (Scardinius erythrophthalmus) and tench (Tinca tinca) were the main indicator species for the small and mostly shallow lakes in both the Alpine (Type 1) and Mediterranean (Type 6) ecoregions, minnow (Phoxinus phoxinus) for the alpine high altitude lakes (Type 2) and landlocked shad (Alosa fallax lacustris), European whitefish (Coregonus lavaretus) and burbot (Lota lota) for the large and very deep alpine lakes (Type 4). The European whitefish was the only indicator species for the deep Mediterranean lakes (Type 5). These species and associated fish assemblages may be useful indicators in future assessments of the ecological status of Italian lakes according to the European Directives (2000/60/EC and 2008/105/EC).     

Regulation and stability in fish assemblages of neotropical floodplain lakes

Neotropical floodplain lakes provide an excellent opportunity to examine the regulation and stability of fish assemblages. At low water, when lakes are separated, fish are concentrated in the lakes and are presumably subject to strong interspecific interactions that can shape assemblage structure. At high water, when the lakes and river channels become broadly interconnected, ample potential exists for alteration of assemblage structure because eggs, larvae, and older fish may undergo spatial reshuffling among waterbodies. Twenty lakes of the Orinoco River floodplain in Venezuela were surveyed in the early and late dry seasons of two consecutive years. Marked differences in assemblage structure among lakes were established in the early dry season. Changes in assemblage structure during the dry season were large and detectable even at the ordinal level; they reflected a strong reduction of visually oriented fish relative to fish with adaptations to low light. Changes were similar in the two years and were apparently due to species-specific differences in mortality, which was high over the dry season. The annual flood is a strong natural fluctuation that greatly modifies assemblage properties during the wet season. Nevertheless, the potential for alteration of assemblage structure by reshuffling was not realized: assemblage properties early and late in the dry season were similar in the two years, indicating a regular and predictable annual cycle of change in assemblage structure. This regularity can be explained by mechanisms of regulation which appear to be linked to piscivory and the optical environment. In contrast with the prevailing views on neotropical fish assemblages, there seems to be a strong deterministic component to assemblage structure and dynamics in Orinoco floodplain lakes.     

Non-native fishes and native species diversity in freshwater fish assemblages across the United States

The proliferation of non-native species in North American freshwater ecosystems is considered a primary threat to the integrity of native community structure. However, a general understanding of consistent and predictable impacts of non-native species on native freshwater diversity is limited, in part, because of a lack of broad-scale studies including data from numerous localities across multiple drainages. This study uses data from 751 localities collected during the United States Geological Survey (USGS) National Water Quality Assessment (NAWQA) program to examine the influence of non-native fish species on native freshwater fish assemblages across the United States. In general, no significant differences in native fish richness and diversity measures were detected between sites with only native species and sites containing non-native species. However, at sites with non-native species, the number of non-native species present was negatively correlated with native species richness and Shannon diversity and positively correlated with native evenness. Non-native piscivores were negatively correlated with native species richness and Shannon diversity and positively correlated with native evenness. Native piscivores were positively correlated with native richness and diversity and negatively correlated with native evenness at sites with only native species. Our results suggest that from a superficial perspective, native species richness and diversity are not different among sites with and without non-native species. However, when patterns of native species richness and diversity are examined at sites containing non-native species, correlations between non-native and native species richness and diversity imply the expected negative effect of invasive taxa. Additionally, non-native piscivores appear to have a significant negative effect on native taxa and possibly represent a novel selective force on naive native prey.     

Functional diversity and trait–environment relationships of stream fish assemblages in a large tropical catchment

1. The species composition of stream fish assemblages changes across the longitudinal fluvial gradient of large river basins. These changes may reflect both zonation in species distributions and environmental filtering of fish traits as stream environments change from the uplands to the lowlands of large catchments. Previous research has shown that taxonomic diversity generally increases in larger, lowland streams, and the River Continuum Concept, the River Habitat Template and other frameworks have provided expectations for what functional groups of fishes should predominate in certain stream types. However, studies addressing the functional trait composition of fish assemblages across large regions are lacking, particularly in tropical river basins. 2. We examined functional trait–environment relationships and functional diversity of stream fish assemblages in the Río Grijalva Basin in southern Mexico. Traits linked to feeding, locomotion and life history strategy were measured in fishes from streams throughout the catchment, from highland headwaters to broad, lowland streams. Relationships between functional traits and environmental variables at local and landscape scales were examined using multivariate ordination, and the convex hull volume of trait space occupied by fish assemblages was calculated as a measure of functional diversity. 3. Although there were a few exceptions, functional diversity of assemblages increased with species richness along the gradient from uplands to lowlands within the Grijalva Basin. Traits related to swimming, habitat preference and food resource use were associated with both local (e.g. substratum type, pool availability) and landscape‐scale (e.g. forest cover) environmental variables. 4. Along with taxonomic structure and diversity, the functional composition of fish assemblages changed across the longitudinal fluvial gradient of the basin. Trait–environment relationships documented in this study partially confirmed theoretical expectations and revealed patterns that may help in developing a better understanding of general functional responses of fish assemblages to environmental change.     

Length–weight relationships of fish species in the freshwater rift valley lakes of Kenya

This study reports length–weight relationships for 10 species of fish encountered in the only two freshwater lakes wholly in the Rift Valley of Kenya, Naivasha and Baringo. In Lake Naivasha, none of the species analysed was native to the lake. Two were introduced species non‐native to East Africa (Micropterus salmoides, Cyprinus carpio), with three native to East Africa but not to the lake (Tilapia zillii, Oreochromis leucostictus, Barbus paludinosus). In Lake Baringo, all species were native (Oreochromis niloctictus baringoensis, Labeo cylindricus, Barbus intermedius autralis, Barbus lineomaculatus, Clarias gariepinus). Specimens were collected by gill netting in September 2004 and October 2005. With the exception of B. lineomaculatus, the b values in the relationship W = aL^b were between 2.90 and 3.22.     

Implications of species loss in freshwater fish assemblages

Freshwater systems are vulnerable to pollution and species loss often ensues. Are there additional implications for assemblage structure? Here we use Berger‐Parker d. Simpson's I/D and Simpson's F to measure the ecological diversity of pristine and perturbed freshwater fish assemblages in Trinidad. West Indies, and Oklahoma. USA. Although the impacted sites typically had fewer species than expected, they could not be distinguished from unperturbed ones of equivalent richness. Changes in the evenness of these assemblages are thus driven by changes in richness. One practical outcome is that diversity indices may not provide independent verification of the detrimental consequences of pollution. The similarity in structure of naturally and anthropogenically impoverished assemblages provides no grounds for complacency, however, since it ignores the evolutionary history of the species concerned. On the basis of our results we suggest that species provenance may be important in tests of ecological function. Moreover, these investigations should replicate natural patterns of evenness as well as richness.     

Length–weight relationships of 38 fish species of the Magdalena River floodplain lakes

This study reports length–weight relationships for 38 fish species belonging to 18 families inhabiting the Magdalena River floodplain lakes of Colombia. Samples were taken during the high water season of 2008, 2010 and 2011. Two species are presented for the first time. Maximum total length records for 24 species exceed those in FishBase.     

Patterns of fish species richness in China's lakes

Aim To document the patterns of fish species richness and their possible causes in China's lakes at regional and national scales. Location Lakes across China. Methods We compiled data of fish species richness, limnological characteristics and climatic variables for 109 lakes across five regions of China: East region, Northeast region, Southwest region, North‐Northwest region, and the Tibetan Plateau. Correlation analyses, regression models and a general linear model were used to explore the patterns of fish species richness. Results At the national scale, lake altitude, energy availability (potential evapotranspiration, PET) and lake area explained 79.6% of the total variation of the lake fish species richness. The determinants of the fish richness pattern varied among physiographic regions. Lake area was the strongest predictor of fish species richness in the East and Southwest lakes, accounting for 22.2% and 82.9% of the variation, respectively. Annual PET explained 68.7% of the variation of fish richness in the Northeast lakes. Maximum depth, mineralization degree, and lake area explained 45.5% of the fish variation in the lakes of the North‐Northwest region. On the Tibetan Plateau, lake altitude was the first predictor variable, interpreting 32.2% of the variation. Main conclusions Lake altitude was the most important factor explaining the variation of fish species richness across China's lakes, and accounted for 74.5% of the variation. This may stem in part from the fact that the lakes investigated in our study span the largest altitudinal range anywhere in the world. The effects of the lake altitude on fish species richness can be separated into direct and indirect aspects due to its collinearity with PET. We also found that the fish diversity and its determinants were scale‐dependent. Fish species richness was probably energy‐determined in the cold region, while it was best predicted by the lake area in the relatively geologically old region. The independent variables we used only explained a small fraction of the variations in the lake fish species richness in East China and the Tibetan Plateau, which may be due to the effects of human activity and historical events, respectively.     

Multilocus phylogeny and species delimitation within the Natterer's bat species complex in the Western Palearctic

Delimiting species is a crucial issue for many biological disciplines and is of primary importance for designing effective conservation plans. Traditional taxonomy based on morphological characters can be misled by the presence of phenotypic plesiomorphism or adaptative convergence. The use of multiple locus genetic data appears thus as a powerful tool for recognizing species boundaries. In this study, we used six nuclear introns and two mitochondrial markers to conduct a phylogenetic study of the Myotis nattereri species complex in the Western Palearctic. We combined tree-based and non-tree-based analyses, and also used concatenated phylogenetic methods of the separated nuclear and mitochondrial dataset as well as a recent coalescence-based multilocus approach. The strong concordance between the results of the analyses conducted confirms that M. nattereri is a paraphyletic group that is composed of four well-differentiated lineages in the study area. In the framework of the unified species concept, these four clades can be confidently considered as four valid species. This recognition of new cryptic species in the Western Mediterranean region shows that the biodiversity of this well-studied area is still not fully understood.     

山东近海鱼类群落分类多样性

根据相关文献整理了山东近海鱼类名录,并根据1998—2009年山东近海鱼类调查名录,应用平均分类差异指数(Δ+)和分类差异变异指数(Λ+)研究了鱼类分类学多样性特征。结果表明,山东近海鱼类名录包括2纲28目91科169属225种,1998年调查仅2纲11目41科58属62种,2006年调查为1纲13目41科71属78种,2009年调查为1纲9目32科55属62种。1998年—2009年调查鱼类种类远远低于鱼类名录记录的种数,分类阶元包含指数较低,平均每属包含1.1种。根据山东近海鱼类名录计算鱼类平均分类差异指数为66.1,分类差异变异指数为141.7;1998—2009年历次调查鱼类平均分类差异指数在60.9—62.7之间,分类差异变异指数在65.4—92.3之间。将1998—2009年历次调查鱼类群落分类多样性指数计算值叠加到山东近海鱼类总名录的95%置信漏斗曲线图,结果表明大部分调查值在置信漏斗曲线之外,目前山东近海鱼类分类多样性已大幅下降。     

Length‐weight relationships of five fish species from lakes of the Central Amazonian floodplains

Parameters of the length‐weight relationship (LWR) were calculated for five fish species from Amazon Basin. Samplings were carried out in ten lakes of the Solimões River floodplains during the four seasons of the hydrological cycle: rising water, high water, receding water, and low water, during two periods. The first period of eight years included samplings performed from 2001 to 2008 and the second sampling period of two years was from 2012 to 2013. Specimens were captured using gillnets (mesh sizes ranging from 30 to 120 mm between opposite knots) and standardized dimension of 20 m in length × 2 m in height. This study provides the LWR parameters for Acarichthys heckelii, Leporinus trifasciatus, Brycon amazonicus, Curimatella meyeri and Osteoglossum bicirrhosum.     

Gravel dredging alters diversity and structure of riverine fish assemblages

1. Human activities affect fish assemblages in a variety of ways. Large‐scale and long‐term disturbances such as in‐stream dredging and mining alter habitat and hydrodynamic characteristics within rivers which can, in turn, alter fish distribution. Habitat heterogeneity is decreased as the natural riffle–pool–run sequences are lost to continuous pools and, as a consequence, lotic species are displaced by lentic species, while generalist and invasive species displace native habitat specialists. Sediment and organic detritus accumulate in deep, dredged reaches and behind dams, disrupting nutrient flow and destroying critical habitat for habitat specialist species. 2. We used standard ecological metrics such as species richness and diversity, as well as stable isotope analysis of δ¹³C and δ¹⁵N, to quantify the differences in fish assemblages sampled by benthic trawls among dredged and undredged sites in the Allegheny River, Pennsylvania, U.S.A. 3. Using mixed‐effects models, we found that total catch, species richness and diversity were negatively correlated with depth (P < 0.05), while species richness, diversity and proportion of species in lithophilic (‘rock‐loving’) reproductive guilds were lower at dredged than at undredged sites (P < 0.05). 4. Principal components analysis and manova revealed that taxa such as darters in brood hider and substratum chooser reproductive guilds were predominantly associated with undredged sites along principal component axis 1 (PC1 and manova P < 0.05), while nest spawners such as catfish and open substratum spawners including suckers were more associated with dredged sites along PC2 (P < 0.05). 5. Stable isotope analysis of δ¹³C and δ¹⁵N revealed shifts from reliance on shallow water and benthic‐derived nutrients at undredged sites to reliance on phytoplankton and terrestrial detritus at deep‐water dredged sites. Relative trophic positions were also lower at dredged sites for many species; loss of benthic nutrient pathways associated with depth and dredging history is hypothesised. 6. The combination of ecological metrics and stable isotope analysis thus shows how anthropogenic habitat loss caused by gravel dredging can decrease benthic fish abundance and diversity, and that species in substratum‐specific reproductive guilds are at particular risk. The effects of dredging also manifest by altering resource use and nutrient pathways within food webs. Management and conservation decisions should therefore consider the protection of relatively shallow areas with suitable substratum for spawning for the protection of native fishes.     

Predicting the structure and diversity of young-of-the-year fish assemblages in large rivers

1. Interactions between environmental variables and 0+ fish assemblages in the upper River Garonne (France) were quantified during late August 1995.
2. The abundance and diversity of the fish assemblages in floodplain channels were modelled using Artificial Neural Network (ANN) analysis and nine variables: the abundance of the six dominant species, fish specific richness, overall abundance of 0+ fish and the Shannon index of diversity. Multiple regression analysis was also used to assess ANN performance.
3. Using 596 samples, correlation coefficients ( r adjusted) between observed and estimated values of the nine dependent parameters were all highly significant ( P < 0.01). Expected values from the tested data were significantly related to the observed values. The correlation coefficient between observed and estimated values ( r ) varied from 0.70 to 0.85.
4. The ANN provided a high quality prediction, despite the complex nature of the relationship between microhabitat composition and fish abundance.
5. Garson's algorithm was used to provide the explanatory power needed in ecology when using black-box models. Parameters contained in the models (i.e. weighting) were used to determine the relative contributions of explanatory variables and thus to ascertain the structure of fish communities.     

Spatial predictors of fish species composition in European lowland lakes

Composition of animal communities can be shaped by both local and regional processes. Among others, dispersal of organisms links local and regional patterns and determines the similarity of communities at increasing spatial distances. Unique and shared spatial and environmental contributions to fish community composition were calculated for watercourse distances between 49 hydrologically connected lakes in the German lowland area. Variation partitioning indicated a dominant unique effect of spatial predictors on fish community composition, whereas the effects of lake morphometry and productivity were weaker. The spatial effect was attributable to an uneven occurrence of small, littoral fish species found even at the small spatial extension covered here (maximum spatial distance ?550 km). Distance decay of community similarity was moderate, but significant, if all 31 fish species were considered, but the slope of the decay function became steeper if only 11 small‐sized, primarily littoral species were included. These results suggest that fish in European lowland lakes can be considered a metacommunity with limited dispersal along watercourse connections in particular for small‐sized species. The analysis supports that for an appropriate evaluation of spatial effects on fish community similarity, reliable estimates of local richness are required which include in particular also rare, small‐sized species occurring primarily in littoral areas. Furthermore, watercourse distance is a more reliable approximation than Euclidean distance to the real spatial dimension of fish dispersal.     

滇中高原湖泊鱼类多样性的研究

依据历年馆藏的鱼类标本和资料,采用多元逐步回归与多项式逐步回归的分析方法,系统分析了滇中6个高原湖泊鱼类的物种丰度、特有种以及属的数目与7个主要湖泊环境因子之间的关系。分析结果显示,控制湖泊鱼类物种丰度、特有种数和属数的关键湖泊环境因子是湖泊面积和湖岸线长度;其中与湖泊面积呈显著正相关,而与湖岸线长度呈负相关。分析不停留于这一常规的横向比较,而是从历史生物地理学的角度,更深入一步揭示了滇中6个湖泊鱼类多样性的演化实质上是受湖泊的发育阶段所控制的。