Temporal scales and patterns of invertebrate biodiversity dynamics in boreal lakes recovering from acidification

Despite international policy implementation to reduce atmospheric acid deposition and restore natural resources from cultural acidification, evidence of ecological recovery is equivocal. Failure to meet recovery goals means that acidification still threatens biodiversity in many areas of the world. Managers thus need information to manage biodiversity, especially its components that are sensitive to stress (acid-sensitive taxa). We analyzed 20-year time series (1988-2007) of water quality and littoral invertebrates in acidified and circum-neutral lakes across Sweden to evaluate regional biodiversity dynamics and the extent to which changes in water quality affect these dynamics. We used multivariate time series modeling to (1) test how individual species groups within invertebrate communities track changes in the abiotic environment and (2) reveal congruencies of taxon contributions to species group change across lakes. Chemical recovery in the lakes was equivocal, and increases of pH and alkalinity were observed in subsets of acidified and circum-neutral lakes. Time series analyses revealed two different patterns of species groups for invertebrate communities across lakes; the first species group showed monotonic change over time, while the second group showed fluctuating temporal patterns. These independent species groups correlated distinctly with different sets of environmental variables. Recovery of pH and alkalinity status was associated with species group patterns only in a few lakes, highlighting an overall weak recovery of invertebrate species. The sets of species, including acid-sensitive taxa, composing these species groups differed markedly across lakes, highlighting context-specific responses of invertebrates to environmental variation. These results are encouraging because disparate local-scale dynamics maintain the diversity of sensitive invertebrate species on a regional scale, despite persisting acidification problems. Our study can inform and help refine current acidification-related policy focused on sensitive biodiversity elements.     

Trajectories of zooplankton recovery in the Little Rock Lake whole-lake acidification experiment.

Understanding the factors that affect biological recovery from environmental stressors such as acidification is an important challenge in ecology. Here we report on zooplankton community recovery following the experimental acidification of Little Rock Lake, Wisconsin, USA. One decade following cessation of acid additions to the northern basin of Little Rock Lake (LRL), recovery of the zooplankton community was complete. Approximately 40% of zooplankton species in the lake exhibited a recovery lag in which biological recovery to reference basin levels was delayed by 1-6 yr after pH recovered to the level at which the species originally responded. Delays in recovery such as those we observed in LRL may be attributable to "biological resistance" wherein establishment of viable populations of key acid-sensitive species following water quality improvements is prevented by other components of the community that thrived during acidification. Indeed, we observed that the recovery of species that thrived during acidification tended to precede recovery of species that declined during acidification. In addition, correspondence analysis indicated that the zooplankton community followed different pathways during acidification and recovery, suggesting that there is substantial hysteresis in zooplankton recovery from acidification. By providing an example of a relatively rapid recovery from short-term acidification, zooplankton community recovery from experimental acidification in LRL generally reinforces the positive outlook for recovery reported for other acidified lakes.     

The interplay between environmental conditions and allee effects during the recovery of stressed zooplankton communities

Many important ecological phenomena depend on the success or failure of small introduced populations. Several factors are thought to influence the fate of small populations, including resource and habitat availability, dispersal levels, interspecific interactions, mate limitation, and demographic stochasticity. Recent field studies suggest that Allee effects resulting from mate limitation can prevent the reestablishment of sexual zooplankton species following a disturbance. In this study, we explore the interplay between Allee effects and local environmental conditions in determining the population growth and establishment of two acid-sensitive zooplankton species that have been impacted by regional anthropogenic acidification. We conducted a factorial design field experiment to test the impact of pH and initial organism densities on the per capita population growth (r) of the sexual copepod Epischura lacustris and the seasonally parthenogenetic cladoceran Daphnia mendotae. In addition, we conducted computer simulations using r values obtained from our experiments to determine the probability of extinction for small populations of acid-sensitive colonists that are in the process of colonizing recovering lakes. The results of our field experiment demonstrated that local environmental conditions can moderate the impacts of Allee effects for E. lacustris: Populations introduced at low densities had a significantly lower r at pH 6 than at pH 7. In contrast, r did not differ between pH 6 and 7 environments when E. lacustris populations were introduced at high densities. D. mendotae was affected by pH levels, but not by initial organism densities. Results from our population growth simulations indicated that E. lacustris populations introduced at low densities to pH 6 conditions had a higher probability of extinction than those introduced at low densities to a pH 7 environment. Our study indicates that environmental conditions and mate limitation can interact to determine the fate of small populations of sexually reproducing zooplankton species. If a more rapid recovery of acid-damaged zooplankton communities is desired, augmentation of dispersal levels may be needed during the early phases of pH recovery in order to increase the probability of establishment for mate-limited zooplankton species.     

Allee effect limits colonization success of sexually reproducing zooplankton

Understanding the dynamics of populations at low density and the role of Allee effects is a priority due to concern about the decline of rare species and interest in colonization/invasion dynamics. Despite well-developed theory and observational support, experimental examinations of the Allee effect in natural systems are rare, partly because of logistical difficulties associated with experiments at low population density. We took advantage of fish introduction and removal in alpine lakes to experimentally test for the Allee effect at the whole-ecosystem scale. The large copepod Hesperodiaptomus shoshone is often extirpated from the water column by fish and sometimes fails to recover following fish disappearance, despite the presence of a long-lived egg bank. Population growth rate of this dioecious species may be limited by mate encounter rate, such that below some critical density a colonizing population will fail to establish. We conducted a multi-lake experiment in which H. shoshone was stocked at densities that bracketed our hypothesized critical density of 0.5-5 copoepods/m3. Successful recovery by the copepod was observed only in the lake with the highest initial density (3 copepods/m3). Copepods stocked into small cages at 3000 copepods/m3 survived and reproduced at rates comparable to natural populations, confirming that the lakes were suitable habitat for this species. In support of mate limitation as the mechanism underlying recovery failure, we found a significant positive relationship between mating success and density across experimental and natural H. shoshone populations. Furthermore, a mesocosm experiment provided evidence of increased per capita population growth rate with increasing population density in another diaptomid species, Skistodiaptomus pallidus. Together, these lines of evidence support the importance of the Allee effect to population recovery of H. shoshone in the Sierra Nevada, and to diaptomid copepods in general.     

The cost of tolerance: sensitivity of stream benthic communities to UV-B and metals.

The ability to tolerate disturbance is a defense strategy that minimizes the effects of damage to fitness and is essential for sustainability of populations, communities, and ecosystems. Despite the apparent benefits of tolerance, there may be an associated cost that results in a deficiency of a system to respond to additional disturbances. Aquatic ecosystems are often exposed to a variety of natural and anthropogenic disturbances, and the effects of these compound perturbations are not well known. In this investigation, we examine whether tolerance to one stressor, metals, results in a cost of increased sensitivity to an additional stressor, ultraviolet-B (UV-B) radiation. Heavy metal pollution is recognized as a major environmental problem in Rocky Mountain streams. These high-elevation, typically clear streams may be at particular risk to elevated UV-B levels associated with reduced levels of ozone. Microcosm experiments were conducted using natural stream benthic communities collected from a reference site and a site with a long-term history of heavy-metal pollution. Direct and interactive effects of heavy metals and UV-B radiation on structural and functional characteristics of benthic communities were evaluated among four treatments: control, UV-B, metals, and metal and UV-B. Communities from the metal-polluted site were more tolerant of metals but less tolerant to UV-B compared to reference communities. Increased mayfly drift and reduced metabolism in response to metal exposure were observed in reference communities but not in the metal-polluted communities. In contrast to these results, UV-B radiation significantly reduced community metabolism, total macroinvertebrate abundance, and abundances of mayflies, caddisflies, and dipterans from the metal-polluted site, but had no effects on benthic communities from the reference site. ANOSIM results demonstrated that community responses differed among treatments at both sites. Metals had the largest impact on community differences at both sites, while UV-B had greater impacts at the metal-polluted site. This research demonstrates the need to account for potential costs associated with tolerance and that these costs can result in behavioral, structural, and functional impacts to benthic communities.     

Using environmental and geographic data to optimize ex situ collections and preserve evolutionary potential

Maintenance of biodiversity through seed banks and botanical gardens, where the wealth of species’ genetic variation may be preserved ex situ, is a major goal of conservation. However, challenges can persist in optimizing ex situ collections if trade-offs exist among cost, effort, and conserving species evolutionary potential, particularly when genetic data are not available. We evaluated the genetic consequences of population preservation informed by geographic (isolation by distance [IBD]) and environmental (isolation by environment [IBE]) distance for ex situ collections for which population provenance is available. We used 19 genetic and genomic data sets from 15 plant species to assess the proportion of population genetic differentiation explained by geographic and environmental factors and to simulate ex situ collections prioritizing source populations based on pairwise geographic distance, environmental distance, or both. Specifically, we tested the impact prioritizing sampling based on these distances may have on the capture of neutral, functional, or putatively adaptive genetic diversity and differentiation. Individually, IBD and IBE explained limited population genetic differences across all 3 genetic marker classes (IBD, 10–16%; IBE, 1–5.5%). Together, they explained a substantial proportion of population genetic differences for functional (45%) and adaptive (71%) variation. Simulated ex situ collections revealed that inclusion of IBD, IBE, or both increased allelic diversity and genetic differentiation captured among populations, particularly for loci that may be important for adaptation. Thus, prioritizing population collections based on environmental and geographic distance data can optimize genetic variation captured ex situ. For the vast majority of plant species for which there is no genetic information, these data are invaluable to conservation because they can guide preservation of genetic variation needed to maintain evolutionary potential within collections.     

Extirpation of Crayfish in a Lake Affected by Long-Range Anthropogenic Acidification

Plastic Lake, Ontario, remote from any point sources of pollutants, is the most thoroughly investigated anthropogenically acidifying lake in North America. As the lake pH decreased from 5.8 to 5.6 over a period of six years, a resident population of the crayfish Orconectes virilis , which had previously exhibited abundances typical of those of nearby Canadian Shield lakes, became extinct. This is one of only half a dozen documented examples of biotic impoverishment due to the long-range aerial transport of strong acids. The loss of crayfish could not be explained by changes in the predaceous fish community. However, the pH range over which extinction occurred matched that of experimentally acidified Lake 223 where O. virilis began to decline dramatically due to recruitment failures, calcium imbalances, and possibly parasite infection. O. virilis appears to be the third component of the benthic invertebrate fauna of Plastic Lake that has been extirpated by anthropogenic acidification.     

Influence of overstory removal on growth of epiphytic mosses and lichens in western Oregon.

Will old-growth-associated epiphytes survive if the forest canopy is opened around them by thinning or partial harvest? If old-growth association is due to a species' environmental tolerances, it may not survive in the relatively open stands that result from such treatments. If, however, old-growth association is due to dispersal limitations rather than environmental tolerances, retention of host trees as refugia and sources of inoculum might carry populations of old-growth-associated epiphytes into young stands. We studied growth rates of lichen and moss transplants in a Douglas-fir (Pseudotsuga menziesii) forest (tree ages approximately 55 yr) in western Oregon for nine months before and 27 months after moderate thinning, creation of 0.4-ha patch cuts, and in control areas. We also assessed moss sporophyte production. We contrasted responses of one moss species, Isothecium myosuroides sensu lato, which is ubiquitous in forests of varying ages, with those of another moss, Antitrichia curtipendula, and a lichen, Lobaria oregana, which are both associated with old-growth forests. Both old-growth associates grew faster in thinned areas and patch cuts than in controls, while Isothecuim grew most slowly and produced fewest sporophytes in patch cuts. These species are likely to survive in remnants, assuming they can remain attached, and may be successful in young stands if they can disperse and establish there. Our results suggest that logging with green-tree retention and other silvicultural practices that preserve trees or shrubs hosting the species studied here are likely to encourage these species' development in managed forests.     

Escape from UV threats in zooplankton: a cocktail of behavior and protective pigmentation

In order to avoid environmental threats, organisms may respond by altering behavior or phenotype. Using experiments performed in high-latitude Siberia and in temperate Sweden, we show for the first time that, among freshwater crustacean zooplankton, the defense against threats from ultraviolet radiation (UV) is a system where phenotypic plasticity and behavioral escape mechanisms function as complementary traits. Freshwater copepods relied mainly on accumulating protective pigments when exposed to UV radiation, but Daphnia showed strong behavioral responses. Pigment levels for both Daphnia and copepods were generally higher at higher latitudes, mirroring different UV threat levels. When released from the UV threat, Daphnia rapidly reduced (within 10 days) their UV protecting pigmentation-by as much as 40%--suggesting a cost in maintaining UV protective pigmentation. The evolutionary advantage of protective pigments is, likely, the ability to utilize the whole water column during daytime; conversely, since the amount of algal food is generally higher in surface waters, unpigmented individuals are restricted to a less preferred feeding habitat in deeper waters. Our main conclusion is that different zooplankton taxa, and similar taxa at different latitudes, use different mixes of behavior and pigments to respond to UV radiation.     

Seasonal variation in the effects of ocean warming and acidification on a native bryozoan, Celleporaria nodulosa

Ocean warming and acidification are co-occurring stressors likely to affect marine biota through climate-driven change to the ocean. We investigated the effects of increased temperature and lowered pH, solely and in combination, on the growth of the endemic Australian bryozoan, Celleporaria nodulosa. Two temperatures and three pH levels were fully crossed in experimental treatments performed in winter 2008 (August) and summer 2009 (February/March). Fragments of C. nodulosa colonies (clones) were collected from Coffs Harbour, NSW, Australia, (30°18′S, 153°09′E) and elongation of colonies was assessed periodically over a 12-day incubation period. Lowered pH in winter significantly decreased growth. Elevated temperatures during the summer significantly impeded the growth of bryozoan colonies, possibly masking the effect of ocean acidification and discovering a maximal thermal tolerance at around 27 °C for C. nodulosa. The effects of decreased pH and increased temperature may be seasonally dependent and particularly acute during the summer months. Thermal stress may in fact be the initial stressor before ocean acidification, negatively affecting organisms in such a way that they are unable to survive before feeling the effects of ocean acidification.     

Genetic Structure and Migration in Native and Reintroduced Rocky Mountain Wolf Populations

Gray wolf (Canis lupus) recovery in the Rocky Mountains of the U.S. is proceeding by both natural recolonization and managed reintroduction. We used DNA microsatellite analysis of wolves transplanted from Canada to two reintroduction sites in the U.S. to study population structure in native and reintroduced wolf populations. Gene flow due to migration between regions in Canada is substantial, and all three recovery populations in the U.S. had high genetic variation. The reintroduced founders were moderately genetically divergent from the naturally colonizing U.S. population. These findings corroborate that the reintroduction more than meets generally accepted genetic guidelines. Maintaining this variation, however, will depend on ample reproduction in the first few generations. In the long term genetic variation will best be retained if migration occurs among the recolonizing and the two transplanted populations. Evidence from field observation and genetic studies shows extensive dispersal by wolves, and we conclude that exchange among these groups due to natural dispersal is likely if public tolerance and legal protection are adequate outside lands designated for wolf recovery.     

Copper tolerance in a population of Silene vulgaris ssp. maritima (A. & D. Love) at Dolfrwynog Bog near Dolgellau,North Wales

Clones of Silene vulgaris ssp. maritima individuals from Dolfrwynog Bog, North Wales and from a coastal control site were propagated under standardised environmental conditions. After five months, cuttings were taken and tested for copper tolerance by means of root extension analysis in solutions of various different copper concentrations. Indices of tolerance suggested that individuals from the Dolfrwynog population were more resistant to elevated copper concentrations than were the control population individuals. Due to the initial five months under standardised conditions, it was assumed that the cloned plant material would have lost all plastic adaptive responses to its native environment. If this assumption was valid then the differences in copper tolerance observed between the two populations had a genetic basis and the Dolfrwynog Bog population is a copper tolerant ecotype.     

Adaptive introgression as a resource for management and genetic conservation in a changing climate

Current rates of climate change require organisms to respond through migration, phenotypic plasticity, or genetic changes via adaptation. We focused on questions regarding species’ and populations’ ability to respond to climate change through adaptation. Specifically, the role adaptive introgression, movement of genetic material from the genome of 1 species into the genome of another through repeated interbreeding, may play in increasing species’ ability to respond to a changing climate. Such interspecific gene flow may mediate extinction risk or consequences of limited adaptive potential that result from standing genetic variation and mutation alone, enabling a quicker demographic recovery in response to changing environments. Despite the near dismissal of the potential benefits of hybridization by conservation practitioners, we examined a number of case studies across different taxa that suggest gene flow between sympatric or parapatric sister species or within species that exhibit strong ecotypic differentiation may represent an underutilized management option to conserve evolutionary potential in a changing environment. This will be particularly true where advanced‐generation hybrids exhibit adaptive traits outside the parental phenotypic range, a phenomenon known as transgressive segregation. The ideas presented in this essay are meant to provoke discussion regarding how we maintain evolutionary potential, the conservation value of natural hybrid zones, and consideration of their important role in adaptation to climate.     

Simulation of aquatic food web and species interactions by adaptive agents embodied with evolutionary computation: a conceptual framework

Individual-based and state variable-based adaptive agents (AA) are discussed regarding their relevance to different types of ecosystems. Individual-based AA proved applicable to a spatially explicit simulation of highly simplified terrestrial food webs. State variable-based AA with evolutionary computation (EC) embodied are suggested for the simulation of aquatic food webs and plankton species interactions. Embodiment of EC in AA can be achieved by evolving predictive rules (ER), differential equations (EDE) or artificial neural networks (ANN) derived from a diverse lake database. In order to provide ecosystem simulation with resilience to environmental change, agent banks can be created containing alternative agents for same species or functional groups from different lakes. State variable-based AA are currently tested for aquatic ecosytem simulation by means of a diverse lake database. It promises to overcome constraints by the rigidity of traditional lake ecosystem models.     

低浓度溶解氧下猪粪固体有机物水解产酸研究

本文以猪粪为发酵原料,通过批式实验研究不同溶解氧(DO)浓度(0—0.26 mg.L-1)、发酵时间(3—12 d)和挥发性固体(VS)浓度(11.14—111.35 g.L-1)对猪粪固体水解酸化过程的影响,确定了低DO浓度下固体有机物优化水解产酸工艺条件:中温35℃,初始VS浓度37.11 g.L-1,初期的DO浓度0.1—0.26 mg.L-1.在此条件下,发酵时间3 d,DO浓度下降到0.10 mg.L-1以下.猪粪发酵液pH值由7.45±0.10降低到5.86±0.17,VS降解率(25.67±1.20)%,挥发性脂肪酸(VFA)中乙酸(3895±91)mg.L-1、丙酸(2313±82)mg.L-1、正丁酸(1361±17)mg.L-1、正戊酸(540±11)mg.L-1.优化条件下的猪粪水解酸化液进行厌氧产甲烷发酵,发酵10 d内产气停止,低溶解氧水解酸化和厌氧产甲烷发酵累计时间仅为13 d,甲烷体积分数(69.5±0.2)%,VS产气率为(0.282±0.011)L CH.4g-1VS.研究结果表明,适当延长发酵时间能够增加VFA中乙酸的含量,初始VS浓度差异对发酵液VFA浓度和VS降解率的影响较显著,低溶解氧能够促进猪粪固体有机物水解过程.     

Invasive Microstegium populations consistently outperform native range populations across diverse environments

Plant species introduced into novel ranges may become invasive due to evolutionary change, phenotypic plasticity, or other biotic or abiotic mechanisms. Evolution of introduced populations could be the result of founder effects, drift, hybridization, or adaptation to local conditions, which could enhance the invasiveness of introduced species. However, understanding whether the success of invading populations is due to genetic differences between native and introduced populations may be obscured by origin x environment interactions. That is, studies conducted under a limited set of environmental conditions may show inconsistent results if native or introduced populations are differentially adapted to specific conditions. We tested for genetic differences between native and introduced populations, and for origin x environment interactions, between native (China) and introduced (U.S.) populations of the invasive annual grass Microstegium vimineum (stiltgrass) across 22 common gardens spanning a wide range of habitats and environmental conditions. On average, introduced populations produced 46% greater biomass and had 7.4% greater survival, and outperformed native range populations in every common garden. However, we found no evidence that introduced Microstegium exhibited greater phenotypic plasticity than native populations. Biomass of Microstegium was positively correlated with light and resident community richness and biomass across the common gardens. However, these relationships were equivalent for native and introduced populations, suggesting that the greater mean performance of introduced populations is not due to unequal responses to specific environmental parameters. Our data on performance of invasive and native populations suggest that post-introduction evolutionary changes may have enhanced the invasive potential of this species. Further, the ability of Microstegium to survive and grow across the wide variety of environmental conditions demonstrates that few habitats are immune to invasion.     

Spatio-temporal variability of polychaete colonization at volcanic CO2 vents indicates high tolerance to ocean acidification

Ocean acidification is predicted to have negative effects on marine biota, resulting in the loss of biodiversity and changes in marine ecosystem structure and function. However, some species and life stages may be capable of thriving in low pH conditions, either due to their natural ability to tolerate stressful low pH–high pCO₂ conditions and/or alteration of species interactions caused by changes in pH profiles, or due to evolutionary trade-offs. A better understanding of which species may be capable of tolerating ocean acidification can guide future research into the mechanisms for physiological and ecological resilience to future carbon dioxide (CO₂) conditions. We investigated the colonization of selected polychaete species along a pH gradient originating from shallow, coastal volcanic CO₂ vents (Ischia, Italy). Colonization was quantified by exposing artificial invertebrate collectors attached to the substratum for 30 days during different periods of the year (late spring, fall and late winter). Three species, Amphiglena mediterranea, Platynereis dumerilii and Syllis prolifera, were present and abundant along the gradient throughout the year. All three species were significantly more abundant in the most acidified areas, confirming their high tolerance and capacity to cope with very low pH. Abundances of all three species were compared to data previously collected via collectors suspended in the water column. More individuals were found in the collectors attached to the substratum, suggesting that abundances may have previously been underestimated. This is likely due to the close proximity of these collectors with the natural rocky substratum. All three species exhibited similar temporal variability, consistent with their life cycle and reproductive biology. Our results demonstrate high tolerance of the species for low and variable pH and corroborate their use as robust models to explore the capacity to cope with low pH–high pCO₂ conditions, both in the natural vent systems and in the laboratory.     

Populations of the Sydney rock oyster, Saccostrea glomerata, vary in response to ocean acidification

Acidifying oceans are predicted to fundamentally alter marine ecosystems. Over the next century, acute studies suggest that the impacts of climate change on marine organisms and ecosystems may be catastrophic. To date, however, little is known about whether the response of marine organisms varies within a species and whether this provides a potential ??adaptive capacity??. Here, we show that selectively bred lines of the ecologically and economically important estuarine mollusc, the Sydney rock oyster Saccostrea glomerata, are more resilient to ocean acidification than the wild populations. When reared at elevated pCO₂, we found a 25% reduction in shell growth of the selectively bred population of the Sydney rock oyster, Saccostrea glomerata, compared to a 64% reduction in shell growth of wild populations. This study shows that there are significantly different sensitivities to ocean acidification even within the same species, providing preliminary evidence that selective breeding may be a solution for important aquaculture industries to overcome the future effects of ocean acidification.     

Prediction of extinction in plants: interaction of extrinsic threats and life history traits

The global extinction of species proceeds through the erosion of local populations. Using a 60-year time series of annual sighting records of plant species, we studied the correlates of local extinction risk associated with a risk of species extinction in the Park Grass Experiment where plants received long-term exposure to nutrient enrichment, soil acidification, and reductions in habitat size. We used multivariate linear models to assess how extrinsic threats and life history traits influence extinction risk. We investigated effects of four extrinsic threats (nitrogen enrichment, productivity, acidification, and plot size) as well as 11 life history traits (month of earliest flowering, flowering duration, stress tolerance, ruderalness [plant species' ability to cope with habitat disturbance], plant height, diaspore mass, seed bank, life form, dispersal mode, apomixis [the ability for a species to reproduce asexuall through seeds], and mating system). Extinction risk was not influenced by plant family. All of the 11 life history traits except life form and all threat variables influenced extinction risk but always via interactions which typically involved one threat variable and one life history trait. We detected comparatively few significant interactions between life history traits, and the interacting traits compensated for each other. These results suggest that simple predictions about extinction risk based on species' traits alone will often fail. In contrast, understanding the interactions between extrinsic threats and life history traits will allow us to make more accurate predictions of extinctions.