Search

Runoff containing road salt (sodium chloride, NaCl) causes the salinization of inland freshwaters, with potentially severe impacts on aquatic species. We performed a mesocosm experiment to test the effects of salinization on plankton community structure in an oligotrophic mountain lake with a limited history of elevated salt concentrations. We exposed plankton communities to a gradient of 30 salt concentrations ranging from 1 to 2900 Cl− mg L−1 for 6 weeks. Adding salt increased zooplankton biomass at concentrations < 500 mg Cl− L−1 while reducing phytoplankton biomass. Zooplankton biomass declined precipitously at higher concentrations, with phytoplankton biomass showing a mirror image increase. The initial increase in zooplankton biomass with salt addition suggests that zooplankton are salt-limited at low ionic concentrations. Additionally, the inverse response of zooplankton and phytoplankton suggests that salinity mainly affects phytoplankton biomass via changes in top-down regulation by grazers.

Knowledge of processes underlying recruitment is critical for understanding marine population dynamics and their response to ocean climate. We investigated the relationship between coastal upwelling and early life history of black rockfish (Sebastes melanops), a midwater aggregating species, and CQB rockfishes (a solitary benthic species complex including Sebastes caurinus, Sebastes maliger, and Sebastes auriculatus), between two oceanographically distinct years on the west coast of Vancouver Island, Canada. We analysed otolith microstructure to determine parturition and settlement dates, pelagic durations, and pre- and postsettlement growth rates. High CQB rockfish recruitment in 2005 was associated with prolonged downwelling and warm ocean temperatures, late parturition dates, fast presettlement growth, short pelagic durations, and small size-at-settlement. In contrast, high black rockfish recruitment in 2006 was associated with strong upwelling and cool ocean temperatures, slow presettlement growth, and protracted pelagic durations. Presettlement growth of both rockfish complexes increased with high sea surface temperature, but was unrelated to chlorophyll a concentration. Our results indicate that the same oceanographic conditions give rise to fast presettlement growth and short pelagic durations for both groups, but that different factors lead to strong recruitment in each.

Daphnia, an ecologically important zooplankton species in lakes, shows both genetic adaptation and phenotypic plasticity in response to temperature and fish predation, but little is known about the molecular basis of these responses and their potential interactions. We performed a factorial experiment exposing laboratory-propagated Daphnia pulicaria clones from two lakes in the Sierra Nevada mountains of California to normal or high temperature (15°C or 25°C) in the presence or absence of fish kairomones, then measured changes in life history and gene expression. Exposure to kairomones increased upper thermal tolerance limits for physiological activity in both clones. Cloned individuals matured at a younger age in response to higher temperature and kairomones, while size at maturity, fecundity and population intrinsic growth were only affected by temperature. At the molecular level, both clones expressed more genes differently in response to temperature than predation, but specific genes involved in metabolic, cellular, and genetic processes responded differently between the two clones. Although gene expression differed more between clones from different lakes than experimental treatments, similar phenotypic responses to predation risk and warming arose from these clone-specific patterns. Our results suggest that phenotypic plasticity responses to temperature and kairomones interact synergistically, with exposure to fish predators increasing the tolerance of Daphnia pulicaria to stressful temperatures, and that similar phenotypic responses to temperature and predator cues can be produced by divergent patterns of gene regulation.

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 (Cl−) 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. This article contains supporting information online at https://www.pnas.org/lookup/suppl/doi:10.1073/pnas.2115033119/-/DCSupplemental

Introgressive hybridization may erode phenotypic divergence along environmental gradients, collapsing locally adapted populations into a hybrid swarm. Alternatively, introgression may promote phenotypic divergence by providing variation on which natural selection can act. In freshwater fishes, water flow often selects for divergent morphological traits in lake versus stream habitats. We tested the effects of introgression on lake–stream morphological divergence in the minnow Owens Tui Chub (Siphateles bicolor snyderi), which has been rendered endangered by introgession from the introduced Lahontan Tui Chub (Siphateles bicolor obesa). Using geometric morphometric analysis of 457 individual Tui Chub from thirteen populations, we found that both native and introgressing parent taxa exhibited divergent body and caudal fin shapes in lake versus stream habitats, but their trajectories of divergence were distinct. In contrast, introgressed populations exhibited intermediate body and caudal fin shapes that were not differentiated by habitat type, indicating that introgression has eroded phenotypic divergence along the lentic–lotic gradient throughout the historic range of the Owens Tui Chub. Individuals within hybrid populations were less morphologically variable than those within parent populations, suggesting hybrid adaptation to selective agents other than water flow or loss of variance by drift., Introgressive hybridization can either erode or promote adaptive phenotypic divergence along environmental gradients. We tested the effects of widespread introgressive hybridization on morphological divergence between lake and stream habitats in the minnow Tui Chub. We found that hybridization has eroded morphological divergence, but comparisons of morphological variance between parent and hybrid populations suggest this outcome may not be maladaptive.

How communities reorganize during climate change depends on the distribution of diversity within ecosystems and across landscapes. Understanding how environmental and evolutionary history constrain community resilience is critical to predicting shifts in future ecosystem function. The goal of our study was to understand how communities with different histories respond to environmental change with regard to shifts in elevation (temperature, nutrients) and introduced predators. We hypothesized that community responses to the environment would differ in ways consistent with local adaptation and initial trait structure. We transplanted plankton communities from lakes at different elevations with and without fish in the Sierra Nevada Mountains in California to mesocosms at different elevations with and without fish. We examined the relative importance of the historical and experimental environment on functional (size structure, effects on lower trophic levels), community (zooplankton composition, abundance and biomass) and population (individual species abundance and biomass) responses. Communities originating from different elevations produced similar biomass at each elevation despite differences in species composition; that is, the experimental elevation, but not the elevation of origin, had a strong effect on biomass. Conversely, we detected a legacy effect of predators on plankton in the fishless environment. Daphnia pulicaria that historically coexisted with fish reached greater biomass under fishless conditions than those from fishless lakes, resulting in greater zooplankton community biomass and larger average size. Therefore, trait variation among lake populations determined the top-down effects of fish predators. In contrast, phenotypic plasticity and local diversity were sufficient to maintain food web structure in response to changing environmental conditions associated with elevation.

The keystone roles of mega-fauna in many terrestrial ecosystems have been lost to defaunation. Large predators and herbivores often play keystone roles in their native ranges, and some have established invasive populations in new biogeographic regions. However, few empirical examples are available to guide expectations about how mega-fauna affect ecosystems in novel environmental and evolutionary contexts. We examined the impacts on aquatic ecosystems of an emerging population of hippopotamus (Hippopotamus amphibus) that has been growing in Colombia over the last 25 yr. Hippos in Africa fertilize lakes and rivers by grazing on land and excreting wastes in the water. Stable isotopes indicate that terrestrial sources contribute more carbon in Colombian lakes containing hippo populations, and daily dissolved oxygen cycles suggest that their presence stimulates ecosystem metabolism. Phytoplankton communities were more dominated by cyanobacteria in lakes with hippos, and bacteria, zooplankton, and benthic invertebrate communities were similar regardless of hippo presence. Our results suggest that hippos recapitulate their role as ecosystem engineers in Colombia, importing terrestrial organic matter and nutrients with detectable impacts on ecosystem metabolism and community structure in the early stages of invasion. Ongoing range expansion may pose a threat to water resources.

Populations at risk of extinction due to climate change may be rescued by adaptive evolution or plasticity. Selective agents, such as introduced predators, may enhance or constrain plastic or adaptive responses to temperature. We tested responses of Daphnia to temperature by collecting populations from lakes across an elevational gradient in the presence and absence of fish predators (long-term selection). We subsequently grew these populations at two elevations in field mesocosms over two years (short-term selection), followed by a common-garden experiment at two temperatures in the lab to measure life-history traits. Both long-term and short-term selection affected traits, suggesting that genetic variation of plasticity within populations enabled individuals to rapidly evolve plasticity in response to high temperature. We found that short-term selection by high temperature increased plasticity for growth rate in all populations. Fecundity was higher in populations from fishless lakes and body size showed greater plasticity in populations from warm lakes (long-term selection). Neither body size nor fecundity were affected by short-term thermal selection. These results demonstrate that plasticity is an important component of the life-history response of Daphnia, and that genetic variation within populations enabled rapid evolution of plasticity in response to selection by temperature.

Variation in resource use among species determines their potential for competition and co-existence, as well as their impact on ecosystem processes. Planktonic crustaceans consume a range of micro-organisms that vary among habitats and species, but these differences in resource consumption are difficult to characterize due to the small size of the organisms. Consumers acquire amino acids from their diet, and the composition of tissues reflects both the use of different resources and their assimilation in proteins. We examined the amino acid composition of common crustacean zooplankton from 14 tropical lakes in Colombia in three regions (the Amazon floodplain, the eastern range of the Andes, and the Caribbean coast). Amino acid composition varied significantly among taxonomic groups and the three regions. Functional richness in amino acid space was greatest in the Amazon, the most productive region, and tended to be positively related to lake trophic status, suggesting the niche breadth of the community could increase with ecosystem productivity. Functional evenness increased with lake trophic status, indicating that species were more regularly distributed within community-wide niche space in more productive lakes. These results show that zooplankton resource use in tropical lakes varies with both habitat and taxonomy, and that lake productivity may affect community functional diversity and the distribution of species within niche space. © 2018, Springer-Verlag GmbH Germany, part of Springer Nature. Acknowledgements NA was supported by a graduate fellowship from Colciencias 1892–2006. JS was supported by a Fulbright Colombia Fellowship, JS and NTJ were supported by NSF DEB 1457737.

The strength of species interactions often varies geographically and locally with environmental conditions. Competitive interactions are predicted to be stronger in benign environments while facilitation is expected to be stronger in harsh ones. We tested these ideas with an aboveground neighbor removal experiment at six salt marshes along the California coast. We determined the effect of removals of either the dominant species, Salicornia pacifica, or the subordinate species on plant cover, aboveground biomass and community composition, as well as soil salinity and moisture. We found that S. pacifica consistently competed with the subordinate species and that the strength of competition varied among sites. In contrast with other studies showing that dominant species facilitate subordinates by moderating physical stress, here the subordinate species facilitated S. pacifica shortly after removal treatments were imposed, but the effect disappeared over time. Contrary to expectations based on patterns observed in east coast salt marshes, we did not see patterns in species interactions in relation to latitude, climate, or soil edaphic characteristics. Our results suggest that variation in interactions among salt marsh plants may be influenced by local‐scale site differences such as nutrients more than broad latitudinal gradients.

Interactions among microbes determine the prevalence of harmful algal blooms that threaten water quality. These interactions can be indirectly mediated by shared resources or consumers, or through interference by the production of allelochemicals. Allelopathic interactions and resource competition have been shown to occur among algae and associated microbes. However, little work has considered seasonal influences on ecosystem structure and function. Here, we report results of our investigations on seasonal changes in the interactions between benthic microbial assemblies and the bloom forming cyanobacterium Microcystis aeruginosa. We show that phosphorus (P) competition and allelopathy by the microbial assembly vary seasonally and inhibit growth of M. aeruginosa. The interactions per unit biomass of the microbial assembly are stronger under winter than summer conditions and inhibit the recruitment of the cyanobacteria, thereby preventing the reoccurrence of cyanobacterial blooms in the following summer. The seasonality of these interactions correlates with changes in composition, metabolic activity and functional diversity of the microbial assembly. Our findings highlight the importance of competitive and allelopathic interactions in regulating the occurrence of harmful algal blooms. Our results also imply that seasonal variation of competition and allelopathy of the microbial assembly might be beneficial to adjust aquatic ecosystem structure and function.

Between 2004 and 2007, Fisheries and Oceans Canada undertook a management action to conserve overfished populations of Inshore Rockfishes by designating 164 Rockfish Conservation Areas (RCAs) closed to most recreational and commercial fishing. However, no research has yet assessed the effectiveness of the RCA network at promoting groundfish population recoveries. We surveyed the fish communities of 35 RCAs and adjacent unprotected areas in southern British Columbia using a remotely operated vehicle (ROV) between 2009 and 2011. We investigated the effect of protection and habitat on fish densities for six species or species groups (Quillback, Yelloweye, Greenstriped Rockfish, Kelp Greenling, Lingcod and all Inshore Rockfish combined) on transects inside and outside of RCAs. Habitat features such as percent rocky substrates and depth influenced fish density while reserve status did not. Next, we calculated habitat-based average densities and used the mean log response ratio (RR) of the density inside to outside of RCAs to determine if the amount of fishing outside the RCA, previous fishing history, the age, area or perimeter to area ratio influenced population recovery. Few positive reserve effects were apparent for any species/group. No clear patterns of RR with age were found for the RCAs, which ranged from 3 to 7 years old at the time of sampling (mean = 4.6). In addition, the intensity of fishing, size, and perimeter-to-area ratio failed to explain RR for most species. There were also no differences in size structure (length) of fish between RCAs and unprotected areas. The results give little indication that demersal fish populations have recovered inside the RCA system. Ongoing monitoring is essential to assess population recovery over time and evaluate the RCAs in terms of criteria such as habitat quality, habitat isolation and the level of compliance in order to enhance their effectiveness.

Resources and temperature play major roles in determining biological production in lake ecosystems. Lakes have been warming and ‘browning’ over recent decades due to climate change and increased loading of terrestrial organic matter. Conflicting hypotheses and evidence have been presented about whether these changes will increase or decrease fish growth within lakes. Most studies have been conducted in low-elevation lakes where terrestrially derived carbon tends to dominate over carbon produced within lakes. Understanding how fish in high-elevation mountain lakes will respond to warming and browning is particularly needed as warming effects are magnified for mountain lakes and treeline is advancing to higher elevations. We sampled 21 trout populations in the Sierra Nevada Mountains of California to examine how body condition and individual growth rates, measured by otolith analysis, varied across independent elevational gradients in temperature and dissolved organic carbon (DOC). We found that fish grew faster at warmer temperatures and higher nitrogen (TN), but slower in high DOC lakes. Additionally, fish showed better body condition in lakes with higher TN, higher elevation and when they exhibited a more terrestrial δ13C isotopic signature. The future warming and browning of lakes will likely have antagonistic impacts on fish growth, reducing the predicted independent impact of warming and browning alone.

Global change involves shifts in multiple environmental factors that act in concert to shape ecological systems in ways that depend on local biotic and abiotic conditions. Little is known about the effects of combined global change stressors on phytoplankton communities, and particularly how these are mediated by distinct community properties such as productivity, grazing pressure and size distribution. Here, we tested for the effects of warming and eutrophication on phytoplankton net growth rate and C:N:P stoichiometry in two phytoplankton cell size fractions (30 µm) in the presence and absence of grazing in microcosm experiments. Because effects may also depend on lake productivity, we used phytoplankton communities from three Dutch lakes spanning a trophic gradient. We measured the response of each community to multifactorial combinations of temperature, nutrient, and grazing treatments and found that nutrients elevated net growth rates and reduced carbon:nutrient ratios of all three phytoplankton communities. Warming effects on growth and stoichiometry depended on nutrient supply and lake productivity, with enhanced growth in the most productive community dominated by cyanobacteria, and strongest stoichiometric responses in the most oligotrophic community at ambient nutrient levels. Grazing effects were also most evident in the most oligotrophic community, with reduced net growth rates and phytoplankton C:P stoichiometry that suggests consumer‐driven nutrient recycling. Our experiments indicate that stoichiometric responses to warming and interactions with nutrient addition and grazing are not universal but depend on lake productivity and cell size distribution., We measured the response of three phytoplankton communities to multifactorial combinations of temperature, nutrient and grazing treatments. Nutrients elevated net growth rates and reduced carbon:nutrient and nitrogen:phosphorus ratios of all communities. Warming effects on growth and stoichiometry depended on lake productivity: warming enhanced growth in the most productive community and caused strongest stoichiometric responses in the least productive community. Grazing reduced C:P and N:P ratios in the least productive community, suggesting consumer‐driven nutrient recycling. Our experiments indicate that stoichiometric responses to warming, and interactions with nutrient supply and grazing, depend on lake productivity and cell size distribution.

Trade‐offs among traits that shape different components of fitness determine the potential for species to coexist in communities and the regulation of ecosystem productivity by consumers vs. resources. We compared functional and structural traits of 14 species of freshwater phytoplankton to test the influence of life history (exponential growth rate, r, and asymptotic density, K), cell volume, cellular fatty acid (FA) biochemistry, and stoichiometry (cellular C:N and C:P ratios) on grazing and growth rate of Daphnia consumers. Grazers exerted the strongest top‐down effects on algae with the highest C:N and C:P, and lower levels of FA unsaturation. In contrast, the grazer's growth rate was most associated with phytoplankton species’ FA unsaturation and chain length. Different traits were related to grazing rate and trophic transfer efficiency, and as a result, the grazing rate on a particular phytoplankton taxon was unrelated to the grazers’ growth rate. The best nutrient competitors (those with high C:N, C:P, and K, which previous experiments showed had lower minimum requirements for nutrients) were most heavily affected by grazing, but did not contribute the most to grazer growth. Our results indicate that Daphnia increase their food consumption rate in order to compensate for low stoichiometric food quality. Strong competitors with low nutrient demands may therefore be subject to the most intense grazing by individual consumers but not support greater production of higher trophic levels.

Lago de Tota, the largest lake in Colombia, is the primary source of water for 250,000 people and a focus of regional economic activity in agriculture, aquaculture, and tourism. Recently, agencies and stakeholders report a shift from the naturally oligotrophic state toward eutrophy. However, the relative contributions of different inputs, including agricultural runoff, aquaculture and municipal wastewaters are unknown, hampering efforts to mitigate nutrient loading. We examined spatial and temporal variation in the trophic state of the lake over one year, as well as stable C and N isotopic profiles of aquatic producers and consumers in relation to two main potential sources (fertilizer and trout feed). We found that Lago de Tota is moderately eutrophic (average chlorophyll-a: 6.4 µg/L, TN: 1.5 mg/L and TP: 0.06 mg/L) with a 32% reduction of transparency over the last 15 years. δ15N and δ13C of aquatic organisms and surface sediments were enriched relative to prehistoric sediments, indicating that human sources dominate the C and N cycles of the lake. δ15N of macrophytes (15.7‰), particulate organic matter (12.5‰), and invertebrates (20.2‰) were enriched relative to trout food (4.6‰), but similar to chicken manure (13.7‰), suggesting that farming in the watershed may be a more important source of N than aquaculture. Our results indicate that Lago de Tota is on a trajectory toward eutrophication with potentially severe consequences for water resources in a rapidly developing mountain region.

Warming, eutrophication (nutrient fertilization) and brownification (increased loading of allochthonous organic matter) are three global trends impacting lake ecosystems. However, the independent and synergistic effects of resource addition and warming on autotrophic and heterotrophic microorganisms are largely unknown. In this study, we investigate the independent and interactive effects of temperature, dissolved organic carbon (DOC, both allochthonous and autochthonous) and nitrogen (N) supply, in addition to the effect of spatial variables, on the composition, richness, and evenness of prokaryotic and eukaryotic microbial communities in lakes across elevation and N deposition gradients in the Sierra Nevada mountains of California, USA. We found that both prokaryotic and eukaryotic communities are structured by temperature, terrestrial (allochthonous) DOC and latitude. Prokaryotic communities are also influenced by total and aquatic (autochthonous) DOC, while eukaryotic communities are also structured by nitrate. Additionally, increasing N availability was associated with reduced richness of prokaryotic communities, and both lower richness and evenness of eukaryotes. We did not detect any synergistic or antagonistic effects as there were no interactions among temperature and resource variables. Together, our results suggest that (a) organic and inorganic resources, temperature, and geographic location (based on latitude and longitude) independently influence lake microbial communities; and (b) increasing N supply due to atmospheric N deposition may reduce richness of both prokaryotic and eukaryotic microbes, probably by reducing niche dimensionality. Our study provides insight into abiotic processes structuring microbial communities across environmental gradients and their potential roles in material and energy fluxes within and between ecosystems.

The relationship between biodiversity and productivity has emerged as a central theme in ecology. Mechanistic explanations for this relationship suggest that the role organisms play in the ecosystem (i.e., niches or functional traits) is a better predictor of ecosystem stability and productivity than taxonomic richness. Here, we tested the capacity of functional diversity in nitrogen uptake in experimental microalgal communities to predict the complementarity effect (CE) and selection effect (SE) of biodiversity on productivity. We grew five algal species as monocultures and as polycultures in pairwise combinations in homogeneous (ammonium, nitrate, or urea alone) and heterogeneous nitrogen (mixed nitrogen) environments to determine whether complementarity between species may be enhanced in heterogeneous environments. We show that the positive diversity effects on productivity in heterogeneous environments resulted from complementarity effects with no positive contribution by species‐specific SEs. Positive biodiversity effects in homogeneous environments, when present (nitrate and urea treatments but not ammonium), were driven both by CE and SE. Our results suggest that functional diversity increases species complementarity and productivity mainly in heterogeneous resource environments. These results provide evidence that the positive effect of functional diversity on community productivity depends on the diversity of resources present in the environment.

Algae hold much promise as a potential feedstock for biofuels and other products, but scaling up biomass production remains challenging. We hypothesized that multispecies assemblages, or polycultures, could improve crop yield when grown in media with mixed nitrogen sources, as found in wastewater. We grew mono- and poly- cultures of algae in four distinct growth media that differed in the form (i.e., nitrate, ammonium, urea, plus a mixture of all three) but not the concentration of nitrogen. We found that mean biomass productivity was positively correlated with algal species richness, and that this relationship was strongest in mixed nitrogen media (on average 88% greater biomass production in 5-species polycultures than in monocultures in mixed nitrogen treatment). We also found that the relationship between nutrient use efficiency and species richness was positive across nitrogen treatments, but greatest in mixed nitrogen media. While polycultures outperformed the most productive monoculture only 0-14% of the time in this experiment, they outperformed the average monoculture 26-52% of the time. Our results suggest that algal polycultures have the potential to be highly productive, and can be effective in recycling nutrients and treating wastewater, offering a sustainable and cost-effective solution for biofuel production.

© The Ecological Society of America. Global demand for transportation fuels will increase rapidly during the upcoming decades, and concerns about fossil-fuel consumption have stimulated research on renewable biofuels that can be sustainably produced from biological feedstocks. However, if unchecked, pathogens and parasites are likely to infect these cultivated biofuel feedstocks, greatly reducing crop yields and potentially threatening the sustainability of renewable bioenergy production efforts. In particular, clonal biofuel crops grown as monocultures at industrial scales will be confronted both by an accumulation of specialist pathogens over time, and by the rapid evolution of those pathogens. We propose possible solutions to these important sustainability problems, with a focus on managing disease risk through crop rotations and the cultivation of multi-species polycultures.

Sea otters are a classic example of a predator controlling ecosystem productivity through cascading effects on basal, habitat-forming kelp species. However, their indirect effects on other kelp-associated taxa like fishes are poorly understood. We examined the effects of sea otter (Enhydra lutris) reintroduction along the west coast of Vancouver Island, Canada on giant kelp (Macrocystis pyrifera) distributions and the trophic niches and growth of two common kelp forest fishes, black (Sebastes melanops) and copper (S. caurinus) rockfishes. We sampled 47 kelp forests, and found that red sea urchins (Strongylocentrotus franciscanus) were eliminated in the presence of otters, and that kelp forests were 3.7 times deeper and 18.8 times larger. Despite order-of-magnitude differences in kelp forest size, adult black and copper rockfishes contained less kelp-derived carbon in their tissues (as measured by stable isotopes of C and N) in regions with otters. Adults of both species had higher mean trophic positions in the presence of otters, indicating more frequent consumption of higher trophic level prey such as fishes. Smaller trophic niche space of rockfishes in the presence of otters indicated a higher degree of trophic specialization. Juvenile black rockfishes rapidly shifted to higher kelp-carbon contents, trophic positions, and body condition factors after settling in kelp forests. The relationships of growth to length, percentage of kelp carbon, and trophic position varied between the two regions, indicating that potential effects of kelp forest size on trophic ontogeny may also affect individual performance. Our results provide evidence that the indirect effects of otters on rockfishes arise largely through the creation of habitat for fishes and other prey rather than a direct trophic connection through invertebrates or other consumers of kelp productivity.

© 2015 by the Ecological Society of America. Understanding the evolutionary potential of organisms to adapt to a changing climate, and the fitness consequences of temperature fluctuations, are critical to forecasting the future of biodiversity. Geographic variation among populations in life history response to temperature mean and variability offers one view of the potential for local adaptation to broaden the thermal niche. We used laboratory growth experiments to examine the effects of temperatures between 138C and 308C on five life history traits and the intrinsic rate of increase for 15 Tigriopus californicus populations distributed over 178 of latitude. Different life history stages showed distinct latitudinal shifts in thermal response, while the temperature of peak population growth consistently declined with increasing latitude. In addition, high-latitude populations grew faster at optimal temperatures but showed steeper fitness declines at high temperature. To test geographic population variation in response to the amplitude of daily thermal fluctuations, we grew three northern and three southern populations and manipulated nightly low and daily high temperatures. We found the lowest fitness overall in the treatment with the highest mean temperature, and the treatment with the greatest variability showed high fitness despite an 88C greater daily range. Population responses to daily thermal variability were unrelated to latitude of origin. Our results indicate that trade-offs between adaptation to high vs. low temperature, and between growth and maturation vs. survival and fecundity, govern local adaptation along the latitudinal gradient. They also indicate that T. californicus populations can maintain fitness over a wide range of daily variability but are more sensitive to small changes in the mean temperature.

Trophic cascades are indirect positive effects of predators on resources via control of intermediate consumers. Larger-bodied predators appear to induce stronger trophic cascades (a greater rebound of resource density toward carrying capacity), but how this happens is unknown because we lack a clear depiction of how the strength of trophic cascades is determined. Using consumer resource models, we first show that the strength of a trophic cascade has an upper limit set by the interaction strength between the basal trophic group and its consumer and that this limit is approached as the interaction strength between the consumer and its predator increases. We then express the strength of a trophic cascade explicitly in terms of predator body size and use two independent parameter sets to calculate how the strength of a trophic cascade depends on predator size. Both parameter sets predict a positive effect of predator size on the strength of a trophic cascade, driven mostly by the body size dependence of the interaction strength between the first two trophic levels. Our results support previous empirical findings and suggest that the loss of larger predators will have greater consequences on trophic control and biomass structure in food webs than the loss of smaller predators.

Genetically engineered (GE) algae offer the promise of producing food, fuel, and other valuable products with reduced requirements for land and fresh water. While the gains in productivity measured in GE terrestrial crops are predicted to be mirrored in GE algae, the stability of phenotypes and ecological risks posed by GE algae in large-scale outdoor cultivation remain unknown. Here, we describe the first US Environmental Protection Agency (EPA)-sanctioned experiment aimed at understanding how GE algae perform in outdoor cultivation. Acutodesmus dimorphus was genetically engineered by the addition of two genes, one for enhanced fatty acid biosynthesis, and one for recombinant green fluorescence protein (GFP) expression; both the genes and their associated phenotypes were maintained during fifty days of outdoor cultivation. We also observed that while the GE algae dispersed from the cultivation ponds, colonization of the trap ponds by the GE strain declined rapidly with increasing distance from the source cultivation ponds. In contrast, many species of indigenous algae were found in every trap pond within a few days of starting the experiment. When inoculated in water from five local lakes, the GE algae's effect on biodiversity, species composition, and biomass of native algae was indiscernible from those of the wild-type (wt) progenitor algae, and neither the GE nor wt algae were able to outcompete native strains. We conclude that GE algae can be successfully cultivated outdoors while maintaining GE traits, and that for the specific GE algal strain tested here they did not outcompete or adversely impact native algae populations when grown in water taken from local lakes. This study provides an initial evaluation of GE algae in outdoor cultivation and a framework to evaluate GE algae risks associated with outdoor GE algae production.

Changing temperature can substantially shift ecological communities by altering the strength and stability of trophic interactions. Because many ecological rates are constrained by temperature, new approaches are required to understand how simultaneous changes in multiple rates alter the relative performance of species and their trophic interactions. We develop an energetic approach to identify the relationship between biomass fluxes and standing biomass across trophic levels. Our approach links ecological rates and trophic dynamics to measure temperature-dependent changes to the strength of trophic interactions and determine how these changes alter food web stability. It accomplishes this by using biomass as a common energetic currency and isolating three temperature-dependent processes that are common to all consumer-resource interactions: biomass accumulation of the resource, resource consumption and consumer mortality. Using this framework, we clarify when and how temperature alters consumer to resource biomass ratios, equilibrium resilience, consumer variability, extinction risk and transient vs. equilibrium dynamics. Finally, we characterise key asymmetries in species responses to temperature that produce these distinct dynamic behaviours and identify when they are likely to emerge. Overall, our framework provides a mechanistic and more unified understanding of the temperature dependence of trophic dynamics in terms of ecological rates, biomass ratios and stability. © 2014 John Wiley & Sons Ltd/CNRS.

Understanding the factors that influence larval dispersal and connectivity among marine populations is critical to the conservation and sustainable management of marine resources. We assessed genetic subdivision among ten populations of copper rockfish (Sebastes caurinus) representing paired samples of outer coast and the heads of inlets in five replicate sounds on the west coast of Vancouver Island, British Columbia, using 17 microsatellite DNA loci. Overall, subdivision (F(ST)) was low (F(ST) = 0.031, P0.001), but consistently higher between paired coast and head of inlet sites (mean FST = 0.047, P0.001) compared to among the five coast sites (mean F(ST) = -0.001, P0.5) or among the five head of inlet sites (mean F(ST) = 0.026, P0.001). Heterozygosity, allelic richness and estimates of effective population size were also lower in head of inlet sites than in coast sites. Bayesian analysis identified two genetic groups across all samples, a single genetic group among only coast samples, two genetic groups among head of inlet samples and two genetic groups within each sound analysed separately. Head of inlet copper rockfish tended to be shorter with lower condition factors and grew more slowly than coast sites fish. Reduced physical connectivity and selection against immigrants in contrasting outer coast-head of inlet environments likely contribute to the evolution of population structure of copper rockfish. Based on genetic connectivity, coast sites appear to be better served by existing marine protected areas than are head of inlet sites.

Microalgae represent one of the most promising groups of candidate organisms for replacing fossil fuels with contemporary primary production as a renewable source of energy. Algae can produce many times more biomass per unit area than terrestrial crop plants, easing the competing demands for land with food crops and native ecosystems. However, several aspects of algal biology present unique challenges to the industrial-scale aquaculture of photosynthetic microorganisms. These include high susceptibility to invading aquatic consumers and weeds, as well as prodigious requirements for nutrients that may compete with the fertiliser demands of other crops. Most research on algal biofuel technologies approaches these problems from a cellular or genetic perspective, attempting either to engineer or select algal strains with particular traits. However, inherent functional trade-offs may limit the capacity of genetic selection or synthetic biology to simultaneously optimise multiple functional traits for biofuel productivity and resilience. We argue that a community engineering approach that manages microalgal diversity, species composition and environmental conditions may lead to more robust and productive biofuel ecosystems. We review evidence for trade-offs, challenges and opportunities in algal biofuel cultivation with a goal of guiding research towards intensifying bioenergy production using established principles of community and ecosystem ecology.

Predators can influence the exchange of carbon dioxide between ecosystems and the atmosphere by altering ecosystem processes such as decomposition and primary production, according to food web theory1,2. Empirical knowledge of such an effect in freshwater systems is limited, but it has been suggested that predators in odd-numbered food chains suppress freshwater carbon dioxide emissions, and predators in even-numbered food chains enhance emissions2,3. Here, we report experiments in three-tier food chains in experimental ponds, streams and bromeliads in Canada and Costa Rica in the presence or absence of fish (Gasterosteus aculeatus) and invertebrate (Hesperoperla pacifica and Mecistogaster modesta) predators. We monitored carbon dioxide fluxes along with prey and primary producer biomass. We found substantially reduced carbon dioxide emissions in the presence of predators in all systems, despite differences in predator type, hydrology, climatic region, ecological zone and level of in situ primary production. We also observed lower amounts of prey biomass and higher amounts of algal and detrital biomass in the presence of predators. We conclude that predators have the potential to markedly influence carbon dioxide dynamics in freshwater systems. Copyright © 2013 Macmillan Publishers Limited.

Climate change shuffles species ranges and creates novel interactions that may either buffer communities against climate change or exacerbate its effect. For instance, facilitation can become more prevalent in salt marshes under stressful conditions while competition is stronger in benign environments. Sea-level rise (SLR) is a consequence of climate change that affects the distribution of stress from inundation and salinity. To determine how interactions early in SLR are affected by changes in these two stressors in Mediterranean-climate marshes, we transplanted marsh turfs to lower elevations to simulate SLR and manipulated cover of the dominant plant species, Salicornia pacifica (formerly Salicornia virginica). We found that both S. pacifica and the subordinate species were affected by inundation treatments, and that subordinate species cover and diversity were lower at low elevations in the presence of S. pacifica than when it was removed. These results suggest that the competitive effect of S. pacifica on other plants is stronger at lower tidal elevations where we also found that salinity is reduced. As sea levels rise, stronger competition by the dominant plant will likely reduce diversity and cover of subordinate species, suggesting that stronger species interactions will exacerbate the effects of climate change on the plant community.

Compliance with spatial fishing regulations (e.g., marine protected areas, fishing closures) is one of the most important, yet rarely measured, determinants of ecological recovery. We used aerial observations of recreational fishing events from creel surveys before, during, and after 77 Rockfish Conservation Areas (RCAs) were established in British Columbia, Canada. There was no evidence of a change in fishing effort in 83% of the RCAs, and effort in five RCAs increased after establishment. Fishing effort in open areas adjacent to the RCAs declined with time and was higher than effort in the RCAs in all 3 years. Next, we used compliance data for 105 RCAs around Vancouver Island to model the drivers of compliance. Compliance was related to the level of fishing effort around the RCA, the size and perimeter-to-area ratio of RCAs, proximity to fishing lodges, and the level of enforcement. Noncompliance in RCAs may be hampering their effectiveness and impeding rockfish recovery. Education and enforcement efforts to reduce fishing effort inside protected areas are critical to the recovery of depleted fish stocks.

Human activities have resulted in rising temperatures and the introduction or extirpation of top predators worldwide. Both processes generate cascading impacts throughout food webs and can jeopardize important ecosystem services. We examined the impact of fish stocking on communities and ecosystems in California mountain lakes across an elevation (temperature and dissolved organic carbon) gradient to determine how trophic cascades and ecosystem function vary with climate. Here, we show that the impact of fish on the pelagic consumer-to-producer biomass ratio strengthened at low elevation, while invertebrate community composition and benthic ecosystem rates (periphyton production and litter decomposition) were most influenced by predators at high elevation. A warming climate may therefore alter the stability of lake ecosystems by shifting the strength of top-down control by introduced predators over food web structure and function.

Conceptual models of adaptive radiation predict that competitive interactions among species will result in an early burst of speciation and trait evolution followed by a slowdown in diversification rates. Empirical studies often show early accumulation of lineages in phylogenetic trees, but usually fail to detect early bursts of phenotypic evolution. We use an evolutionary simulation model to assemble food webs through adaptive radiation, and examine patterns in the resulting phylogenetic trees and species’ traits (body size and trophic position). We find that when foraging trade-offs result in food webs where all species occupy integer trophic levels, lineage diversity and trait disparity are concentrated early in the tree, consistent with the early burst model. In contrast, in food webs in which many omnivorous species feed at multiple trophic levels, high levels of turnover of species’ identities and traits tend to eliminate the early burst signal. These results suggest testable predictions about how the niche structure of ecological communities may be reflected by macroevolutionary patterns.

The exchange of organisms and energy among ecosystems has major impacts on food web structure and dynamics, yet little is known about how climate warming combines with other pervasive anthropogenic perturbations to affect such exchanges. We used an outdoor freshwater mesocosm experiment to investigate the interactive effects of warming, eutrophication, and changes in top predators on the flux of biomass between aquatic and terrestrial ecosystems. We demonstrated that predatory fish decoupled aquatic and terrestrial ecosystems by reducing the emergence of aquatic organisms and suppressing the decomposition of terrestrial plant detritus. In contrast, warming and nutrients enhanced cross-ecosystem exchanges by increasing emergence and decomposition, and these effects were strongest in the absence of predators. Furthermore, we found that warming advanced while predators delayed the phenology of insect emergence. Our results demonstrate that anthropogenic perturbations may extend well beyond ecosystem boundaries by influencing cross-ecosystem subsidies. We find that these changes are sufficient to substantially impact recipient communities and potentially alter the carbon balance between aquatic and terrestrial ecosystems and the atmosphere.

1. Climate change and other human-driven environmental perturbations are causing reductions in biodiversity and impacting the functioning of ecosystems on a global scale. Metacommunity theory suggests that ecosystem connectivity may reduce the magnitude of these impacts if the regional species pool contains functionally redundant species that differ in their environmental tolerances. Dispersal may increase the resistance of local ecosystems to environmental stress by providing regional species with traits adapted to novel conditions. 2. We tested this theory by subjecting freshwater zooplankton communities in mesocosms that were either connected to or isolated from the larger regional species pool to a factorial manipulation of experimental warming and increased salinity. 3. Compensation by regional taxa depended on the source of stress. Warming tolerant regional taxa partially compensated for reductions in heat sensitive local taxa but similar compensation did not occur under increased salinity. 4. Dispersal-mediated species invasions dampened the effects of warming on summer net ecosystem productivity. However, this buffering effect did not occur in the fall or for periphyton growth, the only other ecosystem function affected by the stress treatments. 5. The results indicate that regional biodiversity can provide insurance in a dynamic environment but that the buffering capacity is limited to some ecosystem processes and sources of stress. Maintaining regional biodiversity and habitat connectivity may therefore provide some limited insurance for local ecosystems in changing environments, but is unable to impart resistance against all sources of environmental stress.

The introduction and expansion of exotic species have caused dramatic changes in marine macroalgal assemblages globally. Mechanisms underlying the impacts of macroalgal invaders have been identified for only a few species over small portions of their introduced distribution, and over short time scales. Invasive macroalgae can impact native species through competition for different resources such as light, space or nutrients, as well as modification of abiotic stress and disturbances. We used field experiments to assess the impacts of the invasive macroalga Sargassum muticum (Yendo) Fensholt (Phaeophyceae: Fucales) on native macroalgal communities of the west coast of Vancouver Island, Canada. We manipulated S. muticum in the low intertidal zone to test its effects via both light and space competition at different densities over two consecutive growing seasons. We found non-linear, density-dependent effects of S. muticum on native macroalgal richness. At high cover, S. muticum excludes native species and reduces richness through light competition by shading smaller, understory macroalgae. Light competition was more important than space competition. Native richness was greater when S. muticum cover was reduced to 20% than when it was eliminated entirely, suggesting some facilitation of native species at low cover. The morphology of native macroalgae (canopy, understory, basal) did not predict their response to S. muticum removal, suggesting substantial heterogeneity within functional groups in the types of interactions with S. muticum. The biomass produced by S. muticum failed to compensate for the loss of native biomass due to competition with the exotic, suggesting a negative effect of S. muticum on community productivity. Our results suggest the effects of S. muticum on native macroalgae are largely negative when the invader is abundant, but neutral or positive when the invader is rare.

Until recently, large apex consumers were ubiquitous across the globe and had been for millions of years. The loss of these animals may be humankind’s most pervasive influence on nature. Although such losses are widely viewed as an ethical and aesthetic problem, recent research reveals extensive cascading effects of their disappearance in marine, terrestrial, and freshwater ecosystems worldwide. This empirical work supports long-standing theory about the role of top-down forcing in ecosystems but also highlights the unanticipated impacts of trophic cascades on processes as diverse as the dynamics of disease, wildfire, carbon sequestration, invasive species, and biogeochemical cycles. These findings emphasize the urgent need for interdisciplinary research to forecast the effects of trophic downgrading on process, function, and resilience in global ecosystems.

Ecology Letters (2011) 14: 852–862 Abstract Synergistic interactions between multiple limiting resources are common, highlighting the importance of co-limitation as a constraint on primary production. Our concept of resource limitation has shifted over the past two decades from an earlier paradigm of single-resource limitation towards concepts of co-limitation by multiple resources, which are predicted by various theories. Herein, we summarise multiple-resource limitation responses in plant communities using a dataset of 641 studies that applied factorial addition of nitrogen (N) and phosphorus (P) in freshwater, marine and terrestrial systems. We found that more than half of the studies displayed some type of synergistic response to N and P addition. We found support for strict definitions of co-limitation in 28% of the studies: i.e. community biomass responded to only combined N and P addition, or to both N and P when added separately. Our results highlight the importance of interactions between N and P in regulating primary producer community biomass and point to the need for future studies that address the multiple mechanisms that could lead to different types of co-limitation.

Species interactions come in a variety of forms, from weak to strong, and negative or positive, each with unique consequences for local community structure. However, interactions depend on several biotic, abiotic and scale-dependent variables that make their magnitude and direction difficult to predict. Here, we quantify the relative impacts of multiple factors on species interactions for a diverse array of intertidal organisms, using our own experiments across a range of environments in New Zealand and North America. Interaction strengths are related to organism body size and trophic level, but are relatively insensitive to environmental conditions associated with tidal height. Although species at higher trophic levels exert stronger per-capita effects on other taxa, their population-level impacts are equivalent to basal trophic groups. This indicates that interaction intensity is largely based on requirements for resources, such as food or space, that follow allometric scaling rules. These results demonstrate the potential to predict interactivity based on simple criteria without detailed information on particular species or communities.

If natural communities are assembled according to deterministic rules, coexisting species will represent a nonrandom subset of the potential species pool. We tested for signatures of assembly rules in the distribution of species' traits in Pacific rockfish (Sebastes spp.) assemblages. We used morphology, dietary niche (estimated with stable nitrogen isotopes), and distribution data to identify traits that relate to local-scale resource use (the alpha-niche) and to environmental gradients (the beta-niche). We showed that gill raker morphology was related to trophic position (an alpha-niche axis), while relative eye size was associated with depth habitat (a beta-niche axis). We therefore hypothesized that, within assemblages of coexisting rockfish species, the gill raker trait would be overdispersed (evenly spaced) due to limiting similarity, while relative eye size would be clustered due to environmental filtering. We examined the evolutionary relatedness of coexisting species to ask whether phylogenetic community structure and trait distributions gave similar indications about the roles of assembly processes. We tested the trait distributions and phylogenetic structure of 30 published rockfish assemblages against a null model of random community assembly. As predicted, the gill raker trait tended to be more evenly spaced than expected by chance, as did overall body size, while relative eye size was more clustered than expected. Phylogenetic community structure appeared to reflect historical dispersal and speciation and did not provide consistent support for assembly rules. Our results indicate that rockfish community assembly is nonrandom with regard to species' traits and show how distinguishing traits related to the alpha- and beta-niches and incorporating functional morphology can provide for powerful tests of assembly rules.