Historical and dispersal processes drive community assembly of multiple aquatic taxa in glacierized catchments in the Qinghai-Tibet plateau.

Understanding the relative role of dispersal dynamics and niche constraints is not only a core task in community ecology, but also becomes an important prerequisite for bioassessment. Despite the recent progress in our knowledge of community assembly in space and time, patterns and processes underlying biotic communities in alpine glacierized catchments remain mostly ignored. To fill this knowledge gap, we combined the recently proposed dispersal-niche continuum index (DNCI) with traditional constrained ordinations and idealized patterns of species distributions to unravel community assembly mechanisms of different key groups of primary producers and consumers (i.e., phytoplankton, epiphytic algae, zooplankton, macroinvertebrates, and fishes) in rivers in the Qinghai-Tibet Plateau, the World's Third Pole. We tested whether organismal groups with contrasting body sizes differed in their assembly processes, and discussed their applicability in bioassessment in alpine zones. We found that community structure of alpine river biotas was always predominantly explained in terms of dispersal dynamics and historical biogeography. These patterns are most likely the result of differences in species-specific functional attributes, the stochastic colonization-extinction dynamics driven by multi-year glacier disturbances and the repeated hydrodynamic separation among alpine catchments after the rising of the Qilian mountains. Additionally, we found that the strength of dispersal dynamics and niche constraints was partially mediated by organismal body sizes, with dispersal processes being more influential for microscopic primary producers. Finding that zooplankton and macroinvertebrate communities followed clumped species replacement structures (i.e., Clementsian gradients) supports the notion that environmental filtering also contributes to the structure of high-altitude animal communities in glacierized catchments. In terms of the applied fields, we argue that freshwater bioassessment in glacierized catchments can benefit from incorporating the metacommunity perspective and applying novel approaches to (i) detect the optimal spatial scale for species sorting and (ii) identify and eliminate the species that are sensitive to dispersal-related processes.

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.

Anthropogenic impacts on multiple facets of macroinvertebrate α and ß diversity in a large river-floodplain ecosystem.

Anthropogenic disturbances have become one of the primary causes of biodiversity decline in freshwater ecosystems. Beyond the well-documented loss of taxon richness in increasingly impacted ecosystems, our knowledge on how different facets of α and ß diversity respond to human disturbances is still limited. Here, we examined the responses of taxonomic (TD), functional (FD) and phylogenetic (PD) α and ß diversity of macroinvertebrate communities to human impact across 33 floodplain lakes surrounding the Yangtze River. We found that most pairwise correlations between TD and FD/PD were low and non-significant, whereas FD and PD metrics were instead positively and significantly correlated. All facets of α diversity decreased from weakly to strongly impacted lakes owing to the removal of sensitive species harboring unique evolutionary legacies and phenotypes. By contrast, the three facets of ß diversity responded inconsistently to anthropogenic disturbance: while FDß and PDß showed significant impairment in moderately and strongly impacted lakes as a result of spatial homogenization, TDß was lowest in weakly impacted lakes. The multiple facets of diversity also responded differently to the underlying environmental gradients, re-emphasizing that taxonomic, functional and phylogenetic diversities provide complementary information on community dynamics. However, the explanatory power of our machine learning and constrained ordination models was relatively low and suggests that unmeasured environmental features and stochastic processes may strongly contribute to macroinvertebrate communities in floodplain lakes suffering from variable levels of anthropogenic degradation. We finally suggested guidelines for effective conservation and restoration targets aimed at achieving healthier aquatic biotas in a context of increasing human impact across the 'lakescape' surrounding the Yangtze River, the most important being the control of nutrient inputs and increased spatial spillover effects to promote natural metasystem dynamics.

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.

Comparative transcriptome analysis of scaled and scaleless skins in Gymnocypris eckloni provides insights into the molecular mechanism of scale degeneration.

BACKGROUND: The scale degeneration is thought to be related to the adaptation to the extreme environment with cold climate and high-altitude in schizothoracine fishes. Gymnocypris eckloni, a schizothoracine fish living in plateau waters with the elevation above 2500 m, is nearly esquamate and only covered with shoulder scales and anal scales, making it a good model species to study the molecular mechanism of scale degeneration. RESULTS: The transcriptomes of shoulder scaled skins (SSS), anal scaled skins (ASS) and scaleless skins (NSS) were sequenced and analyzed in G. eckloni at the age of 1 year. Histological examination showed that shoulder scale had completed its differentiation and anal scale just initiated the differentiation. A total of 578,046 unigenes were obtained from the transcriptomes, with 407,799 unigenes annotated in public databases. A total of 428 and 142 differentially expressed unigenes (DEUs) were identified between SSS and NSS, and between ASS and NSS, respectively, with 45 DEUs that were overlapped. Annotation analysis indicated that these DEUs were mainly enriched in Gene Ontology (GO) terms and KEGG pathways associated with bone and muscle formation, such as myofibril, contractile fiber, cytoskeletal protein binding, muscle structure development, cardiac muscle contraction, hypertrophic cardiomyopathy (HCM) and calcium signaling pathway. CONCLUSIONS: Our results would provide insights into the molecular mechanisms of scale degeneration in G. eckloni and other congeneric fishes. In addition, the transcriptome data provides candidate genes and markers for future studies.

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.

Characterization and analysis of the transcriptome in Gymnocypris selincuoensis on the Qinghai-Tibetan Plateau using single-molecule long-read sequencing and RNA-seq.

The lakes on the Qinghai-Tibet Plateau (QTP) are the largest and highest lake group in the world. Gymnocypris selincuoensis is the only cyprinid fish living in lake Selincuo, the largest lake on QTP. However, its genetic resource is still blank, limiting studies on molecular and genetic analysis. In this study, the transcriptome of G. selincuoensis was first generated by using PacBio Iso-Seq and Illumina RNA-seq. A full-length (FL) transcriptome with 75,435 transcripts was obtained by Iso-Seq with N50 length of 3,870 bp. Among all transcripts, 75,016 were annotated to public databases, 64,710 contain complete open reading frames and 2,811 were long non-coding RNAs. Based on all- vs.-all BLAST, 2,069 alternative splicing events were detected, and 80% of them were validated by reverse transcription polymerase chain reaction (RT-PCR). Tissue gene expression atlas showed that the number of detected expressed transcripts ranged from 37,397 in brain to 19,914 in muscle, with 10,488 transcripts detected in all seven tissues. Comparative genomic analysis with other cyprinid fishes identified 77 orthologous genes with potential positive selection (Ka/Ks > 0.3). A total of 56,696 perfect simple sequence repeats were identified from FL transcripts. Our results provide valuable genetic resources for further studies on adaptive evolution, gene expression and population genetics in G. selincuoensis and other congeneric fishes.

Climate-driven synchrony in growth-increment chronologies of fish from the world's largest high-elevation river.

Understanding how sensitive aquatic ecosystems respond to climate change is essential for effective biodiversity conservation and management. The Tibetan Plateau (TP) is one of the most globally sensitive areas to climate change with potentially serious implications for resident fish populations and aquatic food webs. However, how the growth of TP fish responds to climate change, and how this response varies with the trophic level of different species remain unknown. We established growth-increment chronologies of two important Schizothoracinae fishes that are endemic to the TP (e.g., the omnivorous Schizopygopsis younghusbandi and the carnivorous Oxygymnocypris stewartii) from the Yarlung Tsangpo River, using otolith increment width measurements and dendrochronological methods. These growth chronologies were correlated with key indicators of environmental variation (temperature, precipitation, and river discharge) to examine the potential effects of climate change. The two chronologies displayed synchronous responses to recent climate change. In this glacial-fed river, the growth of both fish species was significantly and negatively correlated with the mean annual air temperature, while it was positively but not significantly correlated with precipitation and discharge. The higher trophic level species O. stewartii was more sensitive to climate than was the lower trophic level species S. younghusbandi, with temperature variables explaining a higher proportion of growth variability in O. stewartii (64.6%) than in S. younghusbandi (46.4%). The results collectively indicate that both species are highly sensitive to climate change, which may affect fish growth by altering water environment, fish physiological fitness and food availability. This study provides further empirical evidence of the utility of growth-increment chronologies for investigating the effects of climate change on aquatic ecosystems across different basins and water body types of the TP. These findings can inform conservation and management actions related to addressing climate change on the TP and other high-elevation temperate systems found worldwide.

Strong evidence for changing fish reproductive phenology under climate warming on the Tibetan Plateau.

Phenological responses to climate change have been widely observed and have profound and lasting effects on ecosystems and biodiversity. However, compared to terrestrial ecosystems, the long-term effects of climate change on species' phenology are poorly understood in aquatic ecosystems. Understanding the long-term changes in fish reproductive phenology is essential for predicting population dynamics and for informing management strategies, but is currently hampered by the requirement for intensive field observations and larval identification. In this study, a very low-frequency sampling of juveniles and adults combined with otolith measurements (long axis length of the first annulus; LAFA) of an endemic Tibetan Plateau fish (Gymnocypris selincuoensis) was used to examine changes in reproductive phenology associated with climate changes from the 1970s to 2000s. Assigning individual fish to their appropriate calendar year class was assisted by dendrochronological methods (crossdating). The results demonstrated that LAFA was significantly and positively associated with temperature and growing season length. To separate the effects of temperature and the growing season length on LAFA growth, measurements of larval otoliths from different sites were conducted and revealed that daily increment additions were the main contributor (46.3%), while temperature contributed less (12.0%). Using constructed water-air temperature relationships and historical air temperature records, we found that the reproductive phenology of G. selincuoensis was strongly advanced in the spring during the 1970s and 1990s, while the increased growing season length in the 2000s was mainly due to a delayed onset of winter. The reproductive phenology of G. selincuoensis advanced 2.9 days per decade on average from the 1970s to 2000s, and may have effects on recruitment success and population dynamics of this species and other biota in the ecosystem via the food web. The methods used in this study are applicable for studying reproductive phenological changes across a wide range of species and ecosystems.

Phylogeographic studies of schizothoracine fishes on the central Qinghai-Tibet Plateau reveal the highest known glacial microrefugia.

Pleistocene climatic oscillations have greatly influenced the evolutionary history and distribution pattern of most extant species. However, their effects on species on the Qinghai-Tibet Plateau (QTP) are not well understood. To investigate the effects of past climatic shifts, particularly the Last Glacial Maximum (LGM), on plateau fish, we analysed the phylogeographic structure and demographic history of five closely related taxa of the subfamily Schizothoracinae, a representative endemic taxon of the QTP, from nine endorheic lakes on the central QTP and three peripheral exorheic rivers using the mitochondrial control region (D-loop) sequence and 12 microsatellite (SSR) markers. Phylogram from D-loop haplotypes revealed two well-supported lineages (North and South) separated by the Tanggula Mountains. The results from the D-loop and SSR revealed that endorheic populations possess high genetic diversity and a unique genetic structure. The most recent demographic expansion occurred post-LGM for most endorheic populations and in the last interglacial period for Siling Co and all exorheic populations. Phylogeographic structure, together with species distribution modelling, supports the scenario of multiple glacial refugia on the QTP during the LGM and suggests that Siling Co (4540 m asl) is a cryptic glacial microrefugia for plateau fish, which would be the highest glacial microrefugia known.

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.