Habitat patch dynamics can scale up to influence population demography and diversity with implications for resilience to environmental stochasticity. But how the spatial arrangement and size of habitat patches interact with other components of habitat heterogeneity to shape population diversity at larger spatial scales is not well understood. For riverine fishes, there is increasing evidence that tributary streams provide critical demographic support to main stem rivers. However, the extent to which main stem rivers rely on demographic contributions from tributaries, and the factors underlying this dependence, have not been assessed. Here, we used genetic stock identification to evaluate the effect of tributaries on population diversity of Yellowstone cutthroat trout (Oncorhynchus virginalis bouvieri) occupying the main stem Snake River, Wyoming, USA. We found that the main stem relied almost entirely on tributaries for demographic support, but main stem composition varied spatially among river sections. Distance between habitat patches, catchment area, and groundwater availability acted in concert to determine the contribution of specific tributaries to the main stem, but contributions were ultimately modulated by habitat connectivity. We also found evidence for multi-scale spatial structure in tributary contributions, providing insight into untested drivers of main stem river population diversity. Our results demonstrate how spatially discrete and distributed riverscape attributes influence population diversity at broader spatial scales, illustrating how ecosystem resilience emerges from the dynamic, two-way exchange of individuals and energy across habitat networks. Management plans for large rivers that address the ecological contributions of tributaries may be needed to achieve optimal outcomes. Similarly, conservation strategies that exclusively focus on headwater streams may fail to capture the broader habitat requirements necessary to maintain robust cold-water fish populations and associated recreational fisheries, particularly under global environmental change.

Abstract All organisms contain carbon, nitrogen, and phosphorus in widely ranging amounts and proportions. Integrating existing datasets enables quantification of this variation at global scales. Such efforts could leverage ecological stoichiometry theory, the study of elemental supply and imbalances in ecological interactions, to connect ecological drivers and taxonomic constraints to ecosystem structure and function. Towards this goal, we developed the Limnology Stoichiometric Traits of Organisms In their Chemical Habitats (Limno‐STOICH) database. The Limno‐STOICH database includes 51,576 observations of organismal elemental stoichiometry from >3100 rivers, lakes, wetlands, and other aquatic ecosystem sites on seven continents, derived from 190+ sources. It also includes extensive spatial and temporal metadata to link elemental stoichiometry with ecosystem type, trophic status, etc., and information on organismal data (body size, taxonomic classifications, stable isotope composition) and water physicochemical parameters. The Limno‐STOICH database sets the stage for significant applications across food web ecology, evolutionary ecology, biogeochemistry, and other disciplines.

Abstract Why do some lineages diversify while others do not? This remains a central question in evolutionary ecology. A long-standing assumption, dating to Darwin and embedded in the Unified Neutral Theory of Biodiversity, holds that abundant species should speciate at higher rates. Conversely, theoretical and empirical work highlights the possibility that rare and dispersal-limited clades might be more prone to speciation. Using a birth-death-immigration model with protracted speciation in a multi-population landscape connected by limited dispersal, we show that abundance has a hump-shaped effect on probability of speciation. Our model reveals that intermediate abundance maximizes speciation probability because larger populations disperse more, swamping regional differentiation and inhibiting speciation completion, while smaller populations lack the persistence and incipient speciation needed to diversify. We find empirical support for this prediction with an analysis of data from arthropods endemic to Hawai’i, where genus-level species richness shows a significant hump-shaped relationship with mean genus abundance. These findings provide a mechanistic explanation for a nuanced relationship between abundance and diversification.

Freshwater environments are threatened by multiple anthropogenic stressors. High-elevation mountain lakes are particularly vulnerable to introduced nonnative fish and nutrient deposition because they were historically fishless and typically oligotrophic. To understand the potential effects of fish introduction and nutrient levels on high-elevation lake ecosystems, we assessed differences in zooplankton size, biomass, and density in 76 alpine and subalpine lakes in the Wind River Range, Wyoming, USA, and related those differences to the presence of introduced trout and to food quality (seston nutrient content) and quantity (chlorophyll a concentration). Trout presence, and to a lesser extent trout species, were the strongest predictors of zooplankton composition. Fishless lakes were dominated by low densities of copepods and other large-bodied taxa, and lakes with introduced trout were dominated by high densities of cladocerans, rotifers, and other small-bodied taxa. These assemblage differences are likely because trout reduce or eliminate all large zooplankton taxa by size-selective predation, including predation on Hesperodiaptomus shoshone (S. A. Forbes, 1893), a keystone species that effectively controls populations of rotifers and small crustacean zooplankton taxa. In contrast, the quantity and quality of seston was not associated with zooplankton assemblages. Zooplankton composition in lakes with Rocky Mountain Cutthroat Trout Oncorhynchus virginalis (Girard, 1856) or Golden Trout Oncorhynchus aguabonita (Jordan, 1892) was highly variable, but in lakes with primarily Brook Trout Salvelinus fontinalis (Mitchill, 1814), zooplankton composition was consistently distinct from that in fishless lakes, suggesting that Brook Trout introductions altered the zooplankton assemblage to a greater extent than Cutthroat or Golden trout did. These results contribute to global evidence that predatory fish introductions fundamentally restructure alpine lake food webs. The slow or incomplete recovery of native zooplankton assemblages following fish removal suggests long-term ecological legacies of fish introductions and highlights the importance of understanding factors that promote resilience in high-elevation lake ecosystems.

New species typically evolve over several million years. However, rates of speciation and ecological diversification vary by orders of magnitude across the tree of life, with the fastest shown by some adaptive radiations. Eight hundred endemic species of cichlid fishes emerged and formed entire food webs in Lake Victoria and nearby lakes in East Africa. According to Victorias paleolimnological history, five hundred may have arisen within the past 16,700 years, but molecular phylogenies estimated a much older origin. We reconstruct the age and demography of all Lake Victoria region radiations from whole genomes. We show that indeed, in Lake Victoria all trophic guilds diverged <16,700 years ago, corresponding to between 537 and nearly 30000 speciation events per species per million years, the fastest speciation rate in metazoans. Cichlid radiations in lakes Edward, Albert and Kivu too began <20,000 years ago, an order of magnitude faster than previously thought. Evolutionary transitions between trophic levels led to divergence in effective population sizes as predicted by the trophic pyramid of numbers concept and replicated across three parallel food web radiations. Our results demonstrate that classical theory of trophic interactions in ecologically assembled food webs applies equally to food webs that assembled through rapid adaptive radiation. ### Competing Interest Statement The authors have declared no competing interest.

Top predators have oversized influence on food webs and ecosystem dynamics, and introducing a novel predator to a naive environment can have dramatic consequences for endemic biodiversity. Using genomic data, we find that the colonization of Lake Tanganyika by Lates fishes-the top predators in this ancient lake-occurred more recently than other diverse clades within the lake. Diversification into four endemic Lates species occurred within the lake during a time of dramatic changes in lake levels driven by glacial-interglacial cycles, supporting the hypothesis that these fluctuations were a "species pump" for lacustrine taxa. These lake level fluctuations also likely contributed to multiple admixture events among Lates species during the Pleistocene (~90-500 Kya). Together, our findings suggest a dynamic and environmentally-linked evolutionary history of this predator radiation, and that their colonization of the lake and subsequent diversification likely had dramatic ecosystem consequences for taxa already present in Lake Tanganyika.

Nonlinear dynamics govern ecological processes; thus, understanding thresholds is important for measuring and forecasting the effects of climate change and management of natural resources. However, identifying whether and how such thresholds transfer across ecological levels of organization remains challenging. We argue for a broadening of a foundational organismal concept from ecological stoichiometry theory, the threshold elemental ratio (TER), to study how nonlinear dynamics driven by shifts in limitation operate in evolutionary and ecological processes from organisms to ecosystems. Traditionally, TERs are used to describe the elemental ratio at which the limitation of organismal growth shifts from one element to another. Building on this definition, we make a case for broadening the TER beyond organisms to include populations, clades, communities, and ecosystems. We discuss how TERs may be detected and translated across different ecological levels and evolutionary processes through simulation modeling, literature review, and synthesis of empirical examples from diverse systems and scales including: cyanotoxin production in lakes, alder–salmon dynamics, and the Cambrian explosion. Collectively, we argue that TERs are likely widespread and consequential across levels of ecological organization and that such thresholds manifest from a diversity of mechanisms. Thus, applying the TER concept across ecological levels of organization holds promise for advancing our understanding of nonlinear dynamics from the micro-evolutionary to the macro-ecological.

Nonlinear dynamics govern ecological processes, thus understanding thresholds is important for measuring and forecasting effects of climate change and management of natural resources. However, identifying whether and how such thresholds scale across biological levels of organization remains challenging. Ecological stoichiometry, the study of the balance of multiple elements and energy in ecological systems, provides a framework for scaling thresholds. We broaden a key organismal concept from ecological stoichiometry theory, the Threshold Elemental Ratio (TER), to study how nonlinear dynamics operate in evolutionary and ecological processes across the organizational hierarchy. Traditionally, TERs are used to describe the elemental ratio at which the limitation of organismal growth shifts from one element to another. Following this definition, we make a case for broadening the ecological scale of the TER beyond organisms to include populations, clades, communities, and ecosystems. We show how TERs can be detected and translated across different scales of biological and evolutionary organization through simulation modeling, literature review, and synthesis of empirical examples from diverse systems and ecological scales including: cyanotoxin production in lakes, alder-salmon dynamics, and the Cambrian explosion. Collectively, we demonstrate that TERs are widespread and consequential across levels of biological organization and that such thresholds manifest from a diversity of mechanisms. Thus, scaling of the TER concept holds promise for advancing our understanding of nonlinear dynamics from the micro-evolutionary to macro-ecological. ### Competing Interest Statement The authors have declared no competing interest.

Abstract Dams have negatively affected freshwater biodiversity throughout the world. These negative effects tend to be exacerbated for aquatic taxa with migratory life histories, and for taxa whose habitat is fundamentally altered by the formation of large reservoirs. Sauger (Sander candadensis; Percidae), large‐bodied migratory fishes native to North America, have seen population declines over much of the species' range, and dams are often implicated for their role in blocking access to spawning habitat and otherwise negatively affecting river habitat. Furthermore, hybridization appears to be more frequent between sauger and walleye in the reservoirs formed by large dams. In this study, we examine the role of dams in altering sauger population connectivity and facilitating hybridization with introduced walleye in Wyoming's Wind River and Bighorn River systems. We collected genomic data from individuals sampled over a large spatial scale and replicated sampling throughout the spawning season, with the intent to capture potential variation in hybridization prevalence or genomic divergence between sauger with different life histories. The timing of sampling was not related to hybridization prevalence or population divergence, suggesting limited genetic differences between sauger spawning in different time and places. Overall, there was limited hybridization detected, however, hybridization was most prevalent in Boysen Reservoir (a large impounded section of the Wind River). Dams in the lower Wind River and upper Bighorn River were associated with population divergence between sauger upstream and downstream of the dams, and demographic models suggest that this divergence has occurred in concordance with the construction of the dam. Sauger upstream of the dams exhibited substantially lower estimates of genetic diversity, which implies that disrupted connectivity between Wind River and Bighorn River sauger populations may already be causing negative demographic effects. This research points towards the importance of considering the evolutionary consequences of dams on fish populations in addition to the threats they pose to population persistence.

Molecular phylogenies are a cornerstone of modern comparative biology and are commonly employed to investigate a range of biological phenomena, such as diversification rates, patterns in trait evolution, biogeography, and community assembly. Recent work has demonstrated that significant biases may be introduced into downstream phylogenetic analyses from processing genomic data; however, it remains unclear whether there are interactions among bioinformatic parameters or biases introduced through the choice of reference genome for sequence alignment and variant calling. We address these knowledge gaps by employing a combination of simulated and empirical data sets to investigate the extent to which the choice of reference genome in upstream bioinformatic processing of genomic data influences phylogenetic inference, as well as the way that reference genome choice interacts with bioinformatic filtering choices and phylogenetic inference method. We demonstrate that more stringent minor allele filters bias inferred trees away from the true species tree topology, and that these biased trees tend to be more imbalanced and have a higher center of gravity than the true trees. We find the greatest topological accuracy when filtering sites for minor allele count (MAC) >3-4 in our 51-taxa data sets, while tree center of gravity was closest to the true value when filtering for sites with MAC >1-2. In contrast, filtering for missing data increased accuracy in the inferred topologies; however, this effect was small in comparison to the effect of minor allele filters and may be undesirable due to a subsequent mutation spectrum distortion. The bias introduced by these filters differs based on the reference genome used in short read alignment, providing further support that choosing a reference genome for alignment is an important bioinformatic decision with implications for downstream analyses. These results demonstrate that attributes of the study system and dataset (and their interaction) add important nuance for how best to assemble and filter short-read genomic data for phylogenetic inference.

Adaptive radiations are rich laboratories for exploring, testing, and understanding key theories in evolution and ecology because they offer spectacular displays of speciation and ecological adaptation. Particular challenges to the study of adaptive radiation include high levels of species richness, rapid speciation, and gene flow between species. Over the last decade, high-throughput sequencing technologies and access to population genomic data have lessened these challenges by enabling the analysis of samples from many individual organisms at whole-genome scales. Here we review how population genomic data have facilitated our knowledge of adaptive radiation in five key areas: (1) phylogenetics, (2) hybridization, (3) timing and rates of diversification, (4) the genomic basis of trait evolution, and (5) the role of genome structure in divergence. We review current knowledge in each area, highlight outstanding questions, and focus on methods that facilitate detection of complex patterns in the divergence and demography of populations through time. It is clear that population genomic data are revolutionising the ability to reconstruct evolutionary history in rapidly diversifying clades. Additionally, studies are increasingly emphasising the central role of gene flow, re-use of standing genetic variation during adaptation, and structural genomic elements as facilitators of the speciation process in adaptive radiations. We highlight hybridization—and the hypothesized processes by which it shapes diversification—and questions seeking to bridge the divide between microevolutionary and macroevolutionary processes as rich areas for future study. Overall, access to population genomic data has facilitated an exciting era in adaptive radiation research, with implications for deeper understanding of fundamental evolutionary processes across the tree of life.

Eco-evolutionary interactions following ecosystem change provide critical insight into the ability of organisms to adapt to shifting resource landscapes. Here we explore evidence for the rapid parallel evolution of trout feeding morphology following eco-evolutionary interactions with zooplankton in alpine lakes stocked at different points in time in the Wind River Range (Wyoming, USA). In this system, trout predation has altered the zooplankton species community and driven a decrease in average zooplankton size. In some lakes that were stocked decades ago, we find shifts in gill raker traits consistent with the hypothesis that trout have rapidly adapted to exploit available smaller-bodied zooplankton more effectively. We explore this morphological response in multiple lake populations across two species of trout (cutthroat trout, Oncorhynchus clarkii, and golden trout Oncorhynchus aguabonita) and examine the impact of resource availability on morphological variation in gill raker number among lakes. Furthermore, we present genetic data to provide evidence that historically stocked cutthroat trout populations likely derive from multiple population sources, and incorporate variation from genomic relatedness in our exploration of environmental predictors of feeding morphology. These findings describe rapid adaptation and eco-evolutionary interactions in trout and document an evolutionary response to novel, contemporary ecosystem change.

Although some lineages of animals and plants have made impressive adaptive radiations when provided with ecological opportunity, the propensities to radiate vary profoundly among lineages for unknown reasons. In Africa's Lake Victoria region, one cichlid lineage radiated in every lake, with the largest radiation taking place in a lake less than 16,000 years old. We show that all of its ecological guilds evolved in situ. Cycles of lineage fusion through admixture and lineage fission through speciation characterize the history of the radiation. It was jump-started when several swamp-dwelling refugial populations, each of which were of older hybrid descent, met in the newly forming lake, where they fused into a single population, resuspending old admixture variation. Each population contributed a different set of ancient alleles from which a new adaptive radiation assembled in record time, involving additional fusion-fission cycles. We argue that repeated fusion-fission cycles in the history of a lineage make adaptive radiation fast and predictable.

Explaining broad molecular, phenotypic and species biodiversity patterns necessitates a unifying framework spanning multiple evolutionary scales. Here we argue that although substantial effort has been made to reconcile microevolution and macroevolution, much work remains to identify the links between biological processes at play. We highlight four major questions of evolutionary biology whose solutions require conceptual bridges between micro and macroevolution. We review potential avenues for future research to establish how mechanisms at one scale (drift, mutation, migration, selection) translate to processes at the other scale (speciation, extinction, biogeographic dispersal) and vice versa. We propose ways in which current comparative methods to infer molecular evolution, phenotypic evolution and species diversification could be improved to specifically address these questions. We conclude that researchers are in a better position than ever before to build a synthesis to understand how microevolutionary dynamics unfold over millions of years.

Hybridisation with introduced taxa poses a threat to native fish populations. Mechanisms of reproductive isolation can limit or prevent hybridisation between closely related species. Understanding how these mechanisms interact between the same species across geographically distinct occurrences of secondary contact, and how regional factors influence them, can inform our understanding of hybridisation as a threat and management actions to mitigate this threat. We used data collected on adult fish migration timing and approximate emergence timing of subsequent juvenile fish paired with genomic data to assess whether temporal isolation in the timing of spawning exists between Yellowstone cutthroat trout, rainbow trout and hybrids in the North Fork Shoshone River drainage in northwest Wyoming. We found evidence that Yellowstone cutthroat trout spawn, on average, two to four weeks later than rainbow trout and hybrids and two environmental covariates related to water temperature and discharge were associated with differences in spawning migration timing. Despite statistical support for Yellowstone cutthroat trout spawning later, disproportionately high numbers of rainbow trout and hybrids, paired with extended spawning seasons, lead to substantial overlap between all genotypes. Our results provide further evidence of temporal segregation in the timing of spawning as a mechanism of reproductive isolation between closely related species, but substantial spawning overlap suggests temporal segregation alone will not be enough to curtail hybridisation in conservation populations.

Lake Tanganyika’s pelagic fish sustain the second largest inland fishery in Africa and are under pressure from heavy fishing and global warming related increases in stratification. The strength of water column stratification varies regionally, with a more stratified north and an upwelling-driven, biologically more productive south. Only little is known about whether such regional hydrodynamic regimes induce ecological or genetic differences among populations of highly mobile, pelagic fish inhabiting these different areas. Here, we examine whether the regional contrasts leave distinct isotopic imprints in the pelagic fish of Lake Tanganyika, which may reveal differences in diet or lipid content. We conducted two lake-wide campaigns during different seasons and collected physical, nutrient, chlorophyll, phytoplankton and zooplankton data. Additionally, we analyzed the pelagic fish–the clupeids Stolothrissa tanganicae , Limnothrissa miodon and four Lates species–for their isotopic and elemental carbon (C) and nitrogen (N) compositions. The δ 13 C values were significantly higher in the productive south after the upwelling/mixing period across all trophic levels, implying that the fish have regional foraging grounds, and thus record these latitudinal isotope gradients. By combining our isotope data with previous genetic results showing little geographic structure, we demonstrate that the fish reside in a region for a season or longer. Between specimens from the north and south we found no strong evidence for varying trophic levels or lipid contents, based on their bulk δ 15 N and C:N ratios. We suggest that the development of regional trophic or physiological differences may be inhibited by the lake-wide gene flow on the long term. Overall, our findings show that the pelagic fish species, despite not showing evidence for genetic structure at the basin scale, form regional stocks at the seasonal timescales. This implies that sustainable management strategies may consider adopting regional fishing quotas.

Understanding genetic connectivity plays a crucial role in species conservation decisions, and genetic connectivity is an important component of modern fisheries management. In this study, we investigated the population genetics of four endemic Lates species of Lake Tanganyika (Lates stappersii, L. microlepis, L. mariae and L. angustifrons) using reduced-representation genomic sequencing methods. We find the four species to be strongly differentiated from one another (mean interspecific FST = 0.665), with no evidence for contemporary admixture. We also find evidence for strong genetic structure within L. mariae, with the majority of individuals from the most southern sampling site forming a genetic group that is distinct from the individuals at other sampling sites. We find evidence for much weaker structure within the other three species (L. stappersii, L. microlepis, and L. angustifrons). Our ability to detect this weak structure despite small and unbalanced sample sizes and imprecise geographic sampling locations suggests the possibility for further structure undetected in our study. We call for further research into the origins of the genetic differentiation in these four species—particularly that of L. mariae—which may be important for conservation and management of this culturally and economically important clade of fish.

Additional file 1. Taxonomic information, accession numbers and references of mitochondrial genomes used for phylogenetic analyses.

Background The hydrogeological history of Lake Tanganyika paints a complex image of several colonization and adaptive radiation events. The initial basin was formed around 9-12 million years ago (MYA) from the predecessor of the Malagarasi-Congo River and only 5-6 MYA, its sub-basins fused to produce the clear, deep waters of today. Next to the well-known radiations of cichlid fishes, the lake also harbours a modest clade of only two clupeid species, Stolothrissatanganicae and Limnothrissamiodon. They are members of Pellonulini, a tribe of clupeid fishes that mostly occur in freshwater and that colonized West and Central-Africa during a period of high sea levels during the Cenozoic. There is no consensus on the phylogenetic relationships between members of Pellonulini and the timing of the colonization of Lake Tanganyika by clupeids. Results We use short-read next generation sequencing of 10X Chromium libraries to sequence and assemble the full mitochondrial genomes of S.tanganicae and L.miodon. We then use Maximum likelihood and Bayesian inference to place them into the phylogeny of Pellonulini and other clupeiforms, taking advantage of all available full mitochondrial clupeiform genomes. We identify Potamothrissaobtusirostris as the closest living relative of the Tanganyika sardines and confirm paraphyly for Microthrissa. We estimate the divergence of the Tanganyika sardines around 3.64 MYA [95% CI: 0.99, 6.29], and from P.obtusirostris around 10.92 MYA [95% CI: 6.37-15.48]. Conclusions These estimates imply that the ancestor of the Tanganyika sardines diverged from a riverine ancestor and entered the proto-lake Tanganyika around the time of its formation from the Malagarasi-Congo River, and diverged into the two extant species at the onset of deep clearwater conditions. Our results prompt a more thorough examination of the relationships within Pellonulini, and the new mitochondrial genomes provide an important resource for the future study of this tribe, e.g. as a reference for species identification, genetic diversity, and macroevolutionary studies.