Mercury biomagnification at higher rates than the global average in aquatic ecosystems of the Qinghai-Tibet Plateau.

Mercury biomagnification in aquatic ecosystems is a global issue. Biomagnification patterns and drivers in alpine regions remain poorly understood. Hg biomagnification in the aquatic food web of the Qinghai-Tibetan Plateau (Q-T Plateau) was investigated. A total of 302 fish and macroinvertebrate tissue samples were analysed for total mercury (THg) and nitrogen (δ15N) stable isotope ratios. Overall, 26.75% of fish individuals exceeded the USFWS consumption guidelines. A total of 52.17% of the sampling sites covering different habitats exhibited a significantly positive THg-δ15N relationship, which confirmed the Hg biomagnification potential of Q-T Plateau aquatic ecosystems. The Q-T Plateau Hg biomagnification rates were generally far higher than global averages regardless of the habitat type. Hg in sediments, elevation and population density were positively related to the Hg biomagnification magnitude on the Q-T Plateau, which could be attributed to the disproportionate response of Hg concentrations in macroinvertebrates and fishes along environmental gradients. Our findings offer empirical evidence that fish consumption on the Q-T Plateau poses a substantial Hg exposure risk to people living along river and lake shores. Higher biomagnification rates could further disproportionately accelerate Hg pollution in Q-T Plateau aquatic ecosystems under future anthropogenic activities and climate warming trajectories.

Genetic responses in sexual diploid and unisexual triploid goldfish (Carassius auratus) introduced into a high-altitude environment.

Anthropogenic biological invasions represent major concerns but enable us to investigate rapid evolutionary changes and adaptation to novel environments. The goldfish Carassius auratus with sexual diploids and unisexual triploids coexisting in natural waters is one of the most widespread invasive fishes in Tibet, providing an ideal model to study evolutionary processes during invasion in different reproductive forms from the same vertebrate. Here, using whole-genome resequencing data of 151 C. auratus individuals from invasive and native ranges, we found different patterns of genomic responses between diploid and triploid populations during their invasion of Tibet. For diploids, although invasive individuals derived from two different genetically distinct sources had a relative higher diversity (π) at the population level, their individual genetic diversity (genome-wide observed heterozygosity) was significantly lower (21.4%) than that of source individuals. Population structure analysis revealed that the invasive individuals formed a specific genetic cluster distinct from the source populations. Runs of homozygosity analysis showed low inbreeding only in invasive individuals, and only the invasive population experienced a recent decline in effective population size reflecting founder events. For triploids, however, invasive populations showed no loss of individual genetic diversity and no genetic differentiation relative to source populations. Regions of putative selective sweeps between invasive and source populations of diploids mainly involved genes associated with mannosidase activity and embryo development. Our results suggest that invasive diploids deriving from distinct sources still lost individual genetic diversity resulting from recent inbreeding and founder events and selective sweeps, and invasive triploids experienced no change in genetic diversity owing to their reproduction mode of gynogenesis that precludes inbreeding and founder effects and may make them more powerful invaders.

Across the highest mountain on earth: discordant phylogeographic patterns and recent dispersal of Tibetan stone loaches (Triplophysa) in the Himalayas.

Phylogeographic congruence among co-distributed taxa is regarded as an inherent inference to vicariance events. Nonetheless, incongruent patterns of contemporary lineage divergence among taxa indicated that species differ in their response to common past events. To investigate the role of past events, ecological traits and lineage diversification time in shaping the contemporary phylogeographic patterns, comparative analyses were conducted for Tibetan stone loaches in the Himalayas using three gene markers and two ecological traits (depth of caudal peduncle in their length and presence/absence of posterior chamber of the air bladder). By a thorough sampling in two flanks of the Himalayas, the authors detected that phylogenetic breaks were spatially discordant and divergences of populations were also temporally asynchronous in co-distributed loaches. Estimated divergence time using fossil-calibrated node dating indicated that the Tibetan stone loaches colonised into the south flank of the Himalayas until the Pleistocene. The demographic expansions were also disconcerted between populations in north and south flanks, or east and west Himalayas. Ongoing gene flows between populations in north and south sides implied that the Himalayas do not strictly impede dispersal of cold-adapted species. The results highlight that the quaternary climatic oscillation, in conjunction with ecological traits and lineage diversification time, shaped contemporary phylogenetic patterns of stone loaches in the Himalayas and provide new insights into the biodiversity and composition of species in the Himalayas and surrounding region.

Changes in the Suitable Habitats of Three Endemic Fishes to Climate Change in Tibet.

As one of the most sensitive regions to global climate change, Tibet is subject to remarkable changes in biota over the past decades, including endemic fish species. However, no study has attempted to predict the changes in the distribution of Tibetan fishes, leaving a great blank for aquatic conservation in Tibet. Based on the Maximum Entropy model (MaxEnt), this study predicted the changes in the suitable habitats of three endemic fish species, including two species mainly inhabiting the rivers (Glyptosternon maculatum, Oxygymnocypris stewartii) and one species mainly inhabiting lakes (Gymnocypris selincuoensis) in Tibet under two representative concentration pathways (RCP2.6 and RCP8.5) under two future scenarios (2050 and 2090), and explored the impact of the barrier effects of hydropower projects on the suitable habitats of fish. The results showed that under the four scenarios, the net change in the suitable habitats of the G. maculatum was negative (-2.0--18.8%), while the suitable habitats of the O. stewartii and G. selincuoensis would be expanded, with the net change of 60.0-238.3% and 46.4-56.9%, respectively. Under different scenarios, the suitable habitats of the three species had a tendency to migrate to a higher elevation, and the largest expansion in the range of migration was projected to occur under the 2090-RCP8.5 scenario. In addition, due to the impact of the hydropower projects, the ability of G. maculatum to obtain new suitable habitats from climate change would be reduced by 2.0-8.1%, which was less than the loss induced by climate change (5.5-25.1%), while the suitable habitats of O. stewartii would be reduced by 3.0-9.7%, which was more than the impact of climate change (about 1%). The results of this study have guiding significance for the conservation and management of fish resources diversity in the Yarlung Tsangpo River basin and Siling Co basin of Tibet, and also provide a reference for the coordination and scientific planning of hydropower projects in Tibet.

Predicting impacts of future climate change and hydropower development towards habitats of native and non-native fishes.

Climate change and hydropower development are two primary stressors affecting riverine ecosystems and both stressors facilitate invasions by non-native species. However, little study has focused on how habitats of native and non-native fishes may be affected by independent or combined impacts of such stressors. Here we used the Jinsha River as an example to predict habitat change and distributional shift of native and non-native fishes with species distribution models. The Jinsha River Basin has nearly 40 cascade dams constructed or planned and located in the Tibetan Plateau, which is sensitive to future climate change. Two climate change scenarios and future hydropower development were combined to produce five scenarios of future changes. Under the impacts of independent extreme climate change or hydropower development, non-native fishes showed greater habitat gain in total, while native fishes shifted their distribution into tributaries and higher elevations, and impacts were stronger in combined scenarios. Habitat overlap between the two groups also increased in future scenarios. Certain fish traits correlated with stressors in habitat change prediction. River basins with hydropower development were shown to face higher risk of non-native fishes invasion under future climate change. As the most biodiverse river basins globally are threatened by hydropower development, our results emphasize the importance of regulating non-native fish introduction in reservoirs. Our approaches are also applicable to other systems globally to better understand how hydropower development and climate change may increase invasion risk, and therefore help conserve native species effectively.

Understanding invasion success of Pseudorasbora parva in the Qinghai-Tibetan Plateau: Insights from life-history and environmental filters.

Understanding mechanisms of fish invasion success is crucial to controlling existing invasions and preventing potential future spread. Despite considerable advances in explaining successful fish invasions, little is known about how non-native fish successfully invade alpine freshwater ecosystems. Here, we explore the role of fish life history and environmental factors in contributing to invasion success of Pseudorasbora parva on the Qinghai-Tibet Plateau. We compared life history trait differences between native populations in lowland China with introduced populations in lowland Europe and the high elevation Qinghai-Tibet Plateau. Linear mixed-effects models were used to analyse life-history trait variation across elevation gradients. A random forest model was developed to identify the key environmental filters influencing P. parva invasion success. Life history characteristics differed substantially between native and introduced populations. Compared with native Chinese populations, introduced populations in lowland Europe had smaller body size, higher fecundity, smaller oocytes and earlier maturation. Introduced populations in the Qinghai-Tibet Plateau had smaller body size, lower fecundity, smaller oocytes and later maturation compared with native populations. 1-Year-Length and fecundity in all age classes of females significantly increased with increasing elevation. 2-Year-Length and 3-Year-Length of male significantly increased while maximal longevity and length at first maturity were significantly decreased with the elevation gradient. Habitat type, annual mean temperature, elevation, annual precipitation and precipitation seasonality, were the 5 most important predictors for the occurrence of the P. parva. Our study indicates that invasive P. parva adopt different life history strategies on the plateau compared with invasive populations at low elevations, highlighting that more studies are required for a better understanding of biological invasion under extreme conditions. Considering the ongoing hydrologic alteration and climate change, our study also highlighted that P. parva may expand their distribution range in the future on the Qinghai-Tibet Plateau.

Morphological and genetic divergence between lake and river populations of Triplophysa in Ngangtse Co, Tibet.

Ngangtse Co (4535 m a.s.l., 840 km2) is a typical endorheic lake in Tibet. In 2015, we investigated the fish resource in this lake and its rivers and collected two Triplophysa species, including T. brevicauda and T. stewarti. To understand the evolutionary dynamics of Triplophysa in Ngangtse Co, the adaptive and genetic divergence between river and lake populations were investigated by analysing their morphological characteristics and mitochondrial COI and Cytb sequences. Among all of 277 samples, the ratios of the number of T. brevicauda to T. stewarti were 27:8, 34:11, 14:24, 37:0, 16:23 and 0:83 in river populations RA, RB, RC, RD, lakeside population LSE and lake population L, respectively. For T. brevicauda, a small but significant genetic divergence (mean FST = 0.0890) was detected between population RD and other three river populations, and significant morphological differences of body length/body height and body length/caudal peduncle length were observed between population RD and RA, RB, suggesting an association between differentiation and geographical distance. For T. stewarti, very high levels of genetic differentiation was observed between population L and other populations with the FST values ranging from 0.4737 (L-RA) to 0.7074 (L-RC). In addition, significant differences of most of morphological characteristics were observed between the lake (L) and river (RC) populations, indicating that lake fish of T. stewarti was a population that showed a long-term adaptation to the saltwater lake. These results provide insights on the evolutionary dynamics of Triplophysa in Ngangtse Co and help us to understand the fish speciation in endorheic lakes on the Qinghai-Tibetan Plateau.

The complete mitochondrial genome of Rhynchocypris oxycephalus (Cypriniformes: Cyprinidae).

The Rhynchocypris oxycephalus (Cypriniformes: Cyprinidae) is an ecological important fish in East Asia. In this study, we firstly sequenced the complete mitochondrial genome of R. oxycephalus. The mitogenome is 16,606 bp in length, including 13 protein-coding genes, 22 transfer RNA genes, two ribosomal RNA genes, and two non-coding regions (origin of light-strand replication and control region). The gene content and order is in accord with the common vertebrate form. The nucleotide base composition of H-strand is 28.4% A, 27.2% T, 26.4% C and 18.0% G. The complete mitochondrial genome data would be useful for further studies of genetics, conservation biology and adaptive evolution of R. oxycephalus.

Influence of a large dam on the longitudinal patterns of fish assemblages in Qingyi Stream.

Using seasonally collected data (2009-2010) from 15 sampling sites that represent first- to fifth-order streams within the Qingyi watershed, we examined the spatio-temporal patterns of fish assemblages along two longitudinal gradients to explore the effects of a large dam on fish assemblages at the watershed scale. No significant variation was observed in either species richness or assemblage structure across seasons. Species richness significantly varied according to stream order and gradient. Dam construction appeared to decrease species richness upstream substantially, while a significant decrease between gradients only occurred within fourth-order streams. Along the gradient without the large dam, fish assemblage structures presented distinct separation between two neighboring stream orders, with the exception of fourth-order versus fifth-order streams. However, the gradient disrupted by a large dam displayed the opposite pattern in the spatial variation of fish assemblages related with stream orders. Significant between-gradient differences in fish assemblage structures were only observed within fourth-order streams. Species distributions were determined by local habitat environmental factors, including elevation, substrate, water depth, current discharge, wetted width, and conductivity. Our results suggested that dam construction might alter the longitudinal pattern in fish species richness and assemblage structure in Qingyi Stream, despite the localized nature of the ecological effect of dams.

Development of a fish-based index of biotic integrity for wadeable streams in southern China.

With economic development in China, human-induced pressures on aquatic environments have grown and created an urgent need for tools measuring the ecological condition of aquatic systems. However, biological indicators for wadeable streams in China were poorly developed. This study developed and validated a multi-metric index of fish assemblages for wadeable streams of southern China to meet the requirement of the water project which has been carried on in China in recent years. Fifty-seven stream sites were sampled in April-May and November-December 2010 to develop an index of biotic integrity. A set of 45 candidate metrics were evaluated for range, responsiveness and redundancy, resulting in the selection of six metrics for the index: number of native species, number of rheophilic species, proportion of benthic riffle individuals, number of lithophilic species, number of omnivore species, and number of fish per hour sampling. The publicly available census data were used as independent data set to validate our method. Twenty-three sites were assessed as subject to significant (SP) or non-significant pressures (NSP) based on anthropogenic pressure evaluation. Our index performed well in discriminating NSP and SP sites, which suggested that our method could provide an accurate measure for wadeable streams ecosystem condition. We believe this integrated approach would meet the requirements for the water projects of China, and the process of developing the method could be used as reference for managing the subtropical streams in other areas of China or other states.