Variation in calanoid copepod resting egg abundance among lakes with different acidification histories

Abstract The maintenance of species and genetic diversity within zooplankton egg banks may be crucial to the re-establishment of zooplankton communities following historical disturbance, such as anthropogenic acidification which globally caused widespread damage to ecological communities. Despite this, no other study has described basic characteristics of zooplankton egg banks among lakes with different acidification histories, such as variation in resting egg concentration. Theoretically, habitats with frequent periods of harsh environmental conditions are expected to select for resting egg production or prolonged dormancy in zooplankton, which would increase the size of the resting egg bank in lake sediments. In this study, we compared abundances of viable and inviable calanoid copepod resting eggs among three freshwater lakes with different acidification histories. While Swan Lake underwent major chemical and biological changes from acid and metal deposition, Teardrop and Bat lakes were relatively unaffected by historical acidification and had comparatively constant, but different pH over time. We also tested the effect of age on the viability of resting eggs. As predicted, higher numbers of viable resting eggs were found in recent sediments from acid-recovering Swan Lake compared to study lakes with relatively temporally constant environments (Teardrop and Bat lakes) when the total number of eggs was held as a covariate. We detected this result in spite of similar pelagic abundances of Leptodiaptomus minutus, the dominant species in zooplankton communities of these lakes. This pattern did not necessarily hold for inviable egg concentrations since these eggs were more abundant in both Swan and Bat lakes compared to Teardrop Lake in older sediments (1939-1951, 1800s). Within study lakes, the abundance of viable resting eggs declined with increased egg age. Further study is required to test mechanisms underlying these patterns

Lake salinization drives consistent losses of zooplankton abundance and diversity across coordinated mesocosm experiments

Human-induced salinization increasingly threatens inland waters; yet we know little about the multifaceted response of lake communities to salt contamination. By conducting a coordinated mesocosm experiment of lake salinization across 16 sites in North America and Europe, we quantified the response of zooplankton abundance and (taxonomic and functional) community structure to a broad gradient of environmentally relevant chloride concentrations, ranging from 4 to ca. 1400 mg Cl- L-1. We found that crustaceans were distinctly more sensitive to elevated chloride than rotifers; yet, rotifers did not show compensatory abundance increases in response to crustacean declines. For crustaceans, our among-site comparisons indicate: (1) highly consistent decreases in abundance and taxon richness with salinity; (2) widespread chloride sensitivity across major taxonomic groups (Cladocera, Cyclopoida, and Calanoida); and (3) weaker loss of functional than taxonomic diversity. Overall, our study demonstrates that aggregate properties of zooplankton communities can be adversely affected at chloride concentrations relevant to anthropogenic salinization in lakes.Peer reviewe

Widespread variation in salt tolerance within freshwater zooplankton species reduces the predictability of community-level salt tolerance

The salinization of freshwaters is a global threat to aquatic biodiversity. We quantified variation in chloride (Cl-) tolerance of 19 freshwater zooplankton species in four countries to answer three questions: (1) How much variation in Cl- tolerance is present among populations? (2) What factors predict intraspecific variation in Cl- tolerance? (3) Must we account for intraspecific variation to accurately predict community Cl- tolerance? We conducted field mesocosm experiments at 16 sites and compiled acute LC(50)s from published laboratory studies. We found high variation in LC(50)s for Cl- tolerance in multiple species, which, in the experiment, was only explained by zooplankton community composition. Variation in species-LC50 was high enough that at 45% of lakes, community response was not predictable based on species tolerances measured at other sites. This suggests that water quality guidelines should be based on multiple populations and communities to account for large intraspecific variation in Cl- tolerance.Peer reviewe

Current water quality guidelines across North America and Europe do not protect lakes from salinization

Human-induced salinization caused by the use of road deicing salts, agricultural practices, mining operations, and climate change is a major threat to the biodiversity and functioning of freshwater ecosystems. Yet, it is unclear if freshwater ecosystems are protected from salinization by current water quality guidelines. Leveraging an experimental network of land-based and in-lake mesocosms across North America and Europe, we tested how salinization-indicated as elevated chloride (C-) concentration-will affect lake food webs and if two of the lowest Cl- thresholds found globally are sufficient to protect these food webs. Our results indicated that salinization will cause substantial zooplankton mortality at the lowest Cl- thresholds established in Canada (120 mg Cl-/L) and the United States (230 mg Cl-/L) and throughout Europe where Cl- thresholds are generally higher. For instance, at 73% of our study sites, Cl- concentrations that caused a >= 50% reduction in cladoceran abundance were at or below Cl thresholds in Canada, in the United States, and throughout Europe. Similar trends occurred for copepod and rotifer zooplankton. The loss of zooplankton triggered a cascading effect causing an increase in phytoplankton biomass at 47% of study sites. Such changes in lake food webs could alter nutrient cycling and water clarity and trigger declines in fish production. Current Cl- thresholds across North America and Europe clearly do not adequately protect lake food webs. Water quality guidelines should be developed where they do not exist, and there is an urgent need to reassess existing guidelines to protect lake ecosystems from human-induced salinization.Peer reviewe

Global Patterns and Controls of Nutrient Immobilization On Decomposing Cellulose In Riverine Ecosystems

Microbes play a critical role in plant litter decomposition and influence the fate of carbon in rivers and riparian zones. When decomposing low-nutrient plant litter, microbes acquire nitrogen (N) and phosphorus (P) from the environment (i.e., nutrient immobilization), and this process is potentially sensitive to nutrient loading and changing climate. Nonetheless, environmental controls on immobilization are poorly understood because rates are also influenced by plant litter chemistry, which is coupled to the same environmental factors. Here we used a standardized, low-nutrient organic matter substrate (cotton strips) to quantify nutrient immobilization at 100 paired stream and riparian sites representing 11 biomes worldwide. Immobilization rates varied by three orders of magnitude, were greater in rivers than riparian zones, and were strongly correlated to decomposition rates. In rivers, P immobilization rates were controlled by surface water phosphate concentrations, but N immobilization rates were not related to inorganic N. The N:P of immobilized nutrients was tightly constrained to a molar ratio of 10:1 despite wide variation in surface water N:P. Immobilization rates were temperature-dependent in riparian zones but not related to temperature in rivers. However, in rivers nutrient supply ultimately controlled whether microbes could achieve the maximum expected decomposition rate at a given temperature

Prioritizing taxa for genetic reference database development to advance inland water conservation

Biodiversity loss has accelerated over the past century and freshwater species overall are among those experiencing greatest declines. Genetic resources have the potential to help evaluate the full magnitude of this loss and represent a key tool to effectively allocate conservation resources and monitor the success of restoration efforts. The full power of genetic resources will be realized when the daunting task of referencing all DNA sequences of freshwater organisms is complete. Here, we quantified the availability and distribution of barcode and genome data for freshwater macroscopic organisms in Canada, a country rich in inland water resources and thus particularly vulnerable to aquatic species losses. Impressively, most inland water species (86 %) were represented by barcodes recorded in the BOLD Systems database, while very few had full genomes available (<4 %) in the NCBI database. We identified barcode data deficiencies in northern regions and for taxa assessed as most at risk or without sufficient information for conservation status classification. As expected, the speciose insect group had a lower-than-average number of records per species and a high proportion of data deficient species without adequate barcode coverage. This study highlights where future sequencing resources should be prioritized within initiatives such as the Canada BioGenome Project and BIOSCAN Canada and provides a workflow that could be applied internationally to inform conservation management plans and to mitigate biodiversity loss

Causes of maladaptation

Evolutionary biologists tend to approach the study of the natural world within a framework of adaptation, inspired perhaps by the power of natural selection to produce fitness advantages that drive population persistence and biological diversity. In contrast, evolution has rarely been studied through the lens of adaptation's complement, maladaptation. This contrast is surprising because maladaptation is a prevalent feature of evolution: population trait values are rarely distributed optimally; local populations often have lower fitness than imported ones; populations decline; and local and global extinctions are common. Yet we lack a general framework for understanding maladaptation; for instance in terms of distribution, severity, and dynamics. Similar uncertainties apply to the causes of maladaptation. We suggest that incorporating maladaptation-based perspectives into evolutionary biology would facilitate better understanding of the natural world. Approaches within a maladaptation framework might be especially profitable in applied evolution contexts – where reductions in fitness are common. Toward advancing a more balanced study of evolution, here we present a conceptual framework describing causes of maladaptation. As the introductory article for a Special Feature on maladaptation, we also summarize the studies in this Issue, highlighting the causes of maladaptation in each study. We hope that our framework and the papers in this Special Issue will help catalyze the study of maladaptation in applied evolution, supporting greater understanding of evolutionary dynamics in our rapidly changing world

Evolutionary shifts in copepod acid tolerance in an acid-recovering lake indicated by resurrected resting eggs

Abstract We employed zooplankton resting eggs to track population-level shifts in acid tolerance over the last century in a boreal shield lake recovering from acidification. Shifts in mean and variance of ecological tolerances of species that occupy ecosystems recovering from anthropogenic stress are important to consider because of their potential to influence eco-evolutionary dynamics at community and ecosystem levels. In a laboratory experiment, we compared juvenile survival of Leptodiaptomus minutus copepods hatched from resting eggs from three time periods (80-to 100-year-old: pre-industrial, 20-to 50-year-old: lake acidification, and 8-to 10-year-old: recovery of lake-water back to pH C 6.0) under several pH treatments. Mitochondrial DNA was used to confirm species identity and to test for population bottlenecks as a possible mechanism to explain our results. We expected that nauplii hatched from eggs deposited prior to industrialization (lake-water pH &amp; 6.0) and from the period of pH recovery (lake-water pH C 6.0) would have lower mean and more variable survival at acidic pH compared to nauplii hatched from the period of peak lake acidification (lake-water pH &amp; 4.7). Our results, which are likely a combination of both genetic and environmental effects, suggest support for this hypothesis. Nauplii hatched from eggs deposited during the period of acidification in George Lake had reduced variation in pH tolerance compared to the recovery period. This was likely driven by strong selection rather than genetic drift because we found no evidence of a population bottleneck. However, we could not detect differences in the variance of naupliar survival between pre-industrial and acidic time periods. Trends in mean naupliar survival from different time periods matched findings from other field-based studies that detected a relationship between lake acidification history and acid tolerance in L. minutus

Ecological linkages between community and genetic diversity in zooplankton among boreal shield lakes. Ecology 90:2275–2286

Abstract. Ecological linkages between species diversity in communities and genetic diversity in populations have potential to influence the assembly of communities in habitats recovering from human disturbance, but few studies have attempted to synthesize relationships between these levels of biological organization, especially for locally adapted species. No such studies have been done in freshwater ecosystems despite the plethora of environmental stressors plaguing aquatic communities around the world. We present the first study to test (1) whether diversity and dissimilarity among communities and populations of a locally adapted species are correlated and (2) whether communities and population haplotypes respond differently to environmental selection and spatial structure of habitats. We used a fragment of mitochondrial DNA (mtDNA) belonging to the gene cytochrome oxidase subunit I (COI) as a neutral tag to discriminate among different population haplotype variants. In boreal lakes with different histories of exposure to anthropogenic acidification, diversity and dissimilarity metrics for crustacean zooplankton communities and locally adapted populations of an abundant and broadly distributed calanoid copepod species, Leptodiaptomus minutus, did not correlate. This discord was likely because zooplankton communities responded more strongly to acidity and acidity-related environmental variables than spatial structure of lakes, whereas the distribution of L. minutus haplotypes was more strongly governed by spatial structure of lakes than environmental selection. Although spatial structure was the dominant driver of haplotype structure among L. minutus lake populations, there were similarities in the types of environmental variables that influenced the distributions of species in communities and haplotypes in populations. How haplotype diversity among populations relates to community diversity depends on the relative influence of spatial structure of habitats and selection at each of these scales of biological organization

Appendix C. A 95% statistical-parsimony cladogram of Leptodiaptomus minutus cytochrome oxidase I haplotypes from 38 lakes.

A 95% statistical-parsimony cladogram of Leptodiaptomus minutus cytochrome oxidase I haplotypes from 38 lakes