Species shifts induce soil organic matter priming and changes in microbial communities.

Invasion of plant species with functional traits that influences the rhizosphere can have significant effects on soil organic matter (SOM) dynamics if the invasive species stimulates soil microbial communities with, for example, an enhanced supply of labile carbon and oxygen. We evaluated these effects along a Phragmites invasion chronosequence spanning over 40 years. Using a δ13C and δ15N enriched substrate, we separated SOM-derived and substrate-derived carbon (C) and nitrogen (N) mineralization in surface (top 15 cm), shallow (30-45 cm), and deep (65-80 cm) soils collected from established, newly invaded, and native plant communities. We found all soils were susceptible to SOM priming, but priming profiles differed between vegetation communities, being highest at the surface in native assemblage soils, whereas highest at depth under invasive plants. Changes in functional microbial community composition at depth in Phragmites soils, evidenced by an increase in relative fungal laccase abundance, explained the SOM priming in these deep invaded soils. Our results show that invasive Phragmites maintains a microbial community at depth able to degrade SOM faster than that under native vegetation, evidencing that plant species shifts can fundamentally change soil biogeochemistry, altering element cycling and decreasing SOM residence time. Furthermore, our experimental design allowed to quantify real-time SOM-C and SOM-N gross mineralization, resulting in a new model relating C and N mineralization in these wetland soils and providing new insights on how SOM decomposition impacts N availability and cycling across wetland N pools.

Rapid evolution of a coastal marsh ecosystem engineer in response to global change.

There is increasing evidence that global change can alter ecosystems by eliciting rapid evolution of foundational plants capable of shaping vital attributes and processes. Here we describe results of a field-scale exposure experiment and multilocus assays illustrating that elevated CO2 (eCO2) and nitrogen (N) enrichment can result in rapid shifts in genetic and genotypic variation in Phragmites australis, an ecologically dominant plant that acts as an ecosystem engineer in coastal marshes worldwide. Compared to control treatments, genotypic diversity declined over three years of exposure, especially to N enrichment. The magnitude of loss also increased over time under conditions of N enrichment. Comparisons of genotype frequencies revealed that proportional abundances shifted with exposure to eCO2 and N in a manner consistent with expected responses to selection. Comparisons also revealed evidence of tradeoffs that constrained exposure responses, where any particular genotype responded favorably to one factor rather than to different factors or to combinations of factors. These findings challenge the prevailing view that plant-mediated ecosystem outcomes of global change are governed primarily by differences in species responses to shifting environmental pressures and highlight the value of accounting for organismal evolution in predictive models to improve forecasts of ecosystem responses to global change.

Responses of stomatal features and photosynthesis to porewater N enrichment and elevated atmospheric CO2 in Phragmites australis, the common reed.

PREMISE: Biological invasions increasingly threaten native biodiversity and ecosystem services. One notable example is the common reed, Phragmites australis, which aggressively invades North American salt marshes. Elevated atmospheric CO2 and nitrogen pollution enhance its growth and facilitate invasion because P. australis responds more strongly to these enrichments than do native species. We investigated how modifications to stomatal features contribute to strong photosynthetic responses to CO2 and nitrogen enrichment in P. australis by evaluating stomatal shifts under experimental conditions and relating them to maximal stomatal conductance (gwmax ) and photosynthetic rates. METHODS: Plants were grown in situ in open-top chambers under ambient and elevated atmospheric CO2 (eCO2 ) and porewater nitrogen (Nenr ) in a Chesapeake Bay tidal marsh. We measured light-saturated carbon assimilation rates (Asat ) and stomatal characteristics, from which we calculated gwmax and determined whether CO2 and Nenr altered the relationship between gwmax and Asat . RESULTS: eCO2 and Nenr enhanced both gwmax and Asat , but to differing degrees; gwmax was more strongly influenced by Nenr through increases in stomatal density while Asat was more strongly stimulated by eCO2 . There was a positive relationship between gwmax and Asat that was not modified by eCO2 or Nenr , individually or in combination. CONCLUSIONS: Changes in stomatal features co-occur with previously described responses of P. australis to eCO2 and Nenr . Complementary responses of stomatal length and density to these global change factors may facilitate greater stomatal conductance and carbon gain, contributing to the invasiveness of the introduced lineage.

Rapid recovery of carbon cycle processes after the cessation of chronic nutrient enrichment.

Salt marshes provide critical ecosystem services including some of the highest rates of carbon storage on Earth. However, many salt marshes receive very high nutrient loads and there is a growing body of evidence indicating that this nutrient enrichment alters carbon cycle processes. While many restoration plans prioritize nutrient management in their efforts to conserve salt marsh ecosystems, there has been little empirical investigation of the capacity for carbon cycle processes to recover once nutrient loading is reduced. To address this, we compared rates of greenhouse gas fluxes (i.e., CO2 and methane) measured using static chambers, and soil organic matter decomposition, using both litter bags and the Tea Bag Index (TBI), during the last two years of a long-term, ecosystem-scale nutrient enrichment experiment (2015-2016) as well as in the first two years of recovery post-enrichment (2017-2018). We found that both ecosystem respiration (Reco) and decomposition processes (i.e., rhizome decomposition and soil organic matter stabilization) were enhanced by nutrient enrichment, but returned to reference ecosystem levels within the first year following the cessation of nutrient enrichment and remained at reference levels in the second year. These results suggest that management practices intended to reduce nutrient loads in coastal systems may, in fact, allow for rapid recovery of carbon cycle processes, potentially restoring the high carbon sequestration rates of these blue carbon ecosystems.

Not All Nitrogen Is Created Equal: Differential Effects of Nitrate and Ammonium Enrichment in Coastal Wetlands.

Excess reactive nitrogen (N) flows from agricultural, suburban, and urban systems to coasts, where it causes eutrophication. Coastal wetlands take up some of this N, thereby ameliorating the impacts on nearshore waters. Although the consequences of N on coastal wetlands have been extensively studied, the effect of the specific form of N is not often considered. Both oxidized N forms (nitrate, NO3-) and reduced forms (ammonium, NH4+) can relieve nutrient limitation and increase primary production. However, unlike NH4+, NO3- can also be used as an electron acceptor for microbial respiration. We present results demonstrating that, in salt marshes, microbes use NO3- to support organic matter decomposition and primary production is less stimulated than when enriched with reduced N. Understanding how different forms of N mediate the balance between primary production and decomposition is essential for managing coastal wetlands as N enrichment and sea level rise continue to assail our coasts.

Plant species determine tidal wetland methane response to sea level rise.

Blue carbon (C) ecosystems are among the most effective C sinks of the biosphere, but methane (CH4) emissions can offset their climate cooling effect. Drivers of CH4 emissions from blue C ecosystems and effects of global change are poorly understood. Here we test for the effects of sea level rise (SLR) and its interactions with elevated atmospheric CO2, eutrophication, and plant community composition on CH4 emissions from an estuarine tidal wetland. Changes in CH4 emissions with SLR are primarily mediated by shifts in plant community composition and associated plant traits that determine both the direction and magnitude of SLR effects on CH4 emissions. We furthermore show strong stimulation of CH4 emissions by elevated atmospheric CO2, whereas effects of eutrophication are not significant. Overall, our findings demonstrate a high sensitivity of CH4 emissions to global change with important implications for modeling greenhouse-gas dynamics of blue C ecosystems.

Unraveling the Gordian Knot: Eight testable hypotheses on the effects of nutrient enrichment on tidal wetland sustainability.

The position of tidal wetlands at the land-sea interface makes them especially vulnerable to the effects of nutrient discharges and sea level rise (SLR). Experimental studies of coastal wetland nutrient additions report conflicting results among and within habitats, highlighting the importance of site-specific factors, and how spatial and temporal scaling modulates responses. This suite of influences as SLR accelerates creates a "Gordian Knot" that may compromise coastal habitat integrity. We present eight testable hypotheses here to loosen this knot by identifying critical modulators about nutrient form, soil type and porosity, physiochemical gradients, and eco-evolutionary responses that may control the impacts of nutrient enrichment on coastal wetland sustainability: (1) the delivery and form of the nutrient shapes the ecosystem response; (2) soil type mediates the effects of nutrient enrichment on marshes; (3) belowground responses cannot be solely explained by phenotypic responses; (4) shifting zones of redox and salinity gradients modulate nutrient enrichment impacts; (5) eco-evolutionary processes can drive responses to nutrient availability; (6) nutrient enrichment leads to multiple changed ecosystem states; (7) biogeography trumps a plant's plastic responses to nutrient enrichment; and, (8) nutrient-enhanced wetlands are more susceptible to additional (and anticipated) anthropogenic changes. They provide a framework to investigate and integrate the urgently needed research to understand how excess nutrients threaten the sustainability of coastal wetlands, and wetlands in general. While there is no single 'right way' to test these hypotheses, including a combination of complex and simple, highly-replicated experiments is essential.

The concentration distribution and pollution assessment of heavy metals in surface sediments of the Bohai Bay, China.

Three hundred five surface sediment samples from the Bohai Bay in northeastern China were examined for grain size, organic carbon (Corg) concentration, and concentrations of heavy metals (Pb, Zn, Cu, As, Cr, Cd, and Hg). Average metal concentrations were 33 mg/kg (Cu), 27 mg/kg (Pb), 95 mg/kg (Zn), 75 mg/kg (Cr), 0.3 mg/kg (Cd), 13 mg/kg (As), and 72 µg/kg (Hg). In most cases, these concentrations were lower than the China Marine Sediment Quality criteria. Enrichment factors, however, suggested moderate to strong Cd and Hg contamination of the Bohai Bay. The fact that 68.6% of Pollution Load Index (PLI) values exceeded 2 demonstrated strong pollution of the Bohai Bay, Hg contributed the most to the PLI.

Nitrogen uptake kinetics and saltmarsh plant responses to global change.

Coastal wetlands are important carbon sinks globally, but their ability to store carbon hinges on their nitrogen (N) supply and N uptake dynamics of dominant plant species. In terrestrial ecosystems, uptake of nitrate (NO3-) and ammonium (NH4+) through roots can strongly influence N acquisition rates and their responses to environmental factors such as rising atmospheric CO2 and eutrophication. We examined the 15N uptake kinetics of three dominant plant species in North American coastal wetlands (Spartina patens, C4 grass; Phragmites australis, C3 grass; Schoenoplectus americanus, C3 sedge) under ambient and elevated CO2 conditions. We further related our results to the productivity response of these species in two long-term field experiments. S. patens had the greatest uptake rates for NO3- and NH4+ under ambient conditions, suggesting that N uptake kinetics may underlie its strong productivity response to N in the field. Elevated CO2 increased NH4+ and NO3- uptake rates for S. patens, but had negative effects on NO3- uptake rates in P. australis and no effects on S. americanus. We suggest that N uptake kinetics may explain differences in plant community composition in coastal wetlands and that CO2-induced shifts, in combination with N proliferation, could alter ecosystem-scale productivity patterns of saltmarshes globally.

Cosmopolitan Species As Models for Ecophysiological Responses to Global Change: The Common Reed Phragmites australis.

Phragmites australis is a cosmopolitan grass and often the dominant species in the ecosystems it inhabits. Due to high intraspecific diversity and phenotypic plasticity, P. australis has an extensive ecological amplitude and a great capacity to acclimate to adverse environmental conditions; it can therefore offer valuable insights into plant responses to global change. Here we review the ecology and ecophysiology of prominent P. australis lineages and their responses to multiple forms of global change. Key findings of our review are that: (1) P. australis lineages are well-adapted to regions of their phylogeographic origin and therefore respond differently to changes in climatic conditions such as temperature or atmospheric CO2; (2) each lineage consists of populations that may occur in geographically different habitats and contain multiple genotypes; (3) the phenotypic plasticity of functional and fitness-related traits of a genotype determine the responses to global change factors; (4) genotypes with high plasticity to environmental drivers may acclimate or even vastly expand their ranges, genotypes of medium plasticity must acclimate or experience range-shifts, and those with low plasticity may face local extinction; (5) responses to ancillary types of global change, like shifting levels of soil salinity, flooding, and drought, are not consistent within lineages and depend on adaptation of individual genotypes. These patterns suggest that the diverse lineages of P. australis will undergo intense selective pressure in the face of global change such that the distributions and interactions of co-occurring lineages, as well as those of genotypes within-lineages, are very likely to be altered. We propose that the strong latitudinal clines within and between P. australis lineages can be a useful tool for predicting plant responses to climate change in general and present a conceptual framework for using P. australis lineages to predict plant responses to global change and its consequences.

Nutrient foraging strategies are associated with productivity and population growth in forest shrubs.

Background and Aims: Temperate deciduous forest understoreys are experiencing widespread changes in community composition, concurrent with increases in rates of nitrogen supply. These shifts in plant abundance may be driven by interspecific differences in nutrient foraging (i.e. conservative vs. acquisitive strategies) and, thus, adaptation to contemporary nutrient loading conditions. This study sought to determine if interspecific differences in nutrient foraging could help explain patterns of shrub success and decline in eastern North American forests. Methods: Using plants grown in a common garden, fine root traits associated with nutrient foraging were measured for six shrub species. Traits included the mean and skewness of the root diameter distribution, specific root length (SRL), C:N ratio, root tissue density, arbuscular mycorrhizal colonization and foraging precision. Above- and below-ground productivity were also determined for the same plants, and population growth rates were estimated using data from a long-term study of community dynamics. Root traits were compared among species and associations among root traits, measures of productivity and rates of population growth were evaluated. Key Results: Species fell into groups having thick or thin root forms, which correspond to conservative vs. acquisitive nutrient foraging strategies. Interspecific variation in root morphology and tissue construction correlated with measures of productivity and rates of cover expansion. Of the four species with acquisitive traits, three were introduced species that have become invasive in recent decades, and the fourth was a weedy native. In contrast, the two species with conservative traits were historically dominant shrubs that have declined in abundance in eastern North American forests. Conclusions: In forest understoreys of eastern North America, elevated nutrient availability may impose a filter on species success in addition to above-ground processes such as herbivory and overstorey canopy conditions. Shrubs that have root traits associated with rapid uptake of soil nutrients may be more likely to increase in abundance, while species without such traits may be less likely to keep pace with more productive species.

An invasive wetland grass primes deep soil carbon pools.

Understanding the processes that control deep soil carbon (C) dynamics and accumulation is of key importance, given the relevance of soil organic matter (SOM) as a vast C pool and climate change buffer. Methodological constraints of measuring SOM decomposition in the field prevent the addressing of real-time rhizosphere effects that regulate nutrient cycling and SOM decomposition. An invasive lineage of Phragmites australis roots deeper than native vegetation (Schoenoplectus americanus and Spartina patens) in coastal marshes of North America and has potential to dramatically alter C cycling and accumulation in these ecosystems. To evaluate the effect of deep rooting on SOM decomposition we designed a mesocosm experiment that differentiates between plant-derived, surface SOM-derived (0-40 cm, active root zone of native marsh vegetation), and deep SOM-derived mineralization (40-80 cm, below active root zone of native vegetation). We found invasive P. australis allocated the highest proportion of roots in deeper soils, differing significantly from the native vegetation in root : shoot ratio and belowground biomass allocation. About half of the CO2 produced came from plant tissue mineralization in invasive and native communities; the rest of the CO2 was produced from SOM mineralization (priming). Under P. australis, 35% of the CO2 was produced from deep SOM priming and 9% from surface SOM. In the native community, 9% was produced from deep SOM priming and 44% from surface SOM. SOM priming in the native community was proportional to belowground biomass, while P. australis showed much higher priming with less belowground biomass. If P. australis deep rooting favors the decomposition of deep-buried SOM accumulated under native vegetation, P. australis invasion into a wetland could fundamentally change SOM dynamics and lead to the loss of the C pool that was previously sequestered at depth under the native vegetation, thereby altering the function of a wetland as a long-term C sink.

Saltmarsh plant responses to eutrophication.

In saltmarsh plant communities, bottom-up pressure from nutrient enrichment is predicted to increase productivity, alter community structure, decrease biodiversity, and alter ecosystem functioning. Previous work supporting these predictions has been based largely on short-term, plot-level (e.g., 1-300 m2 ) studies, which may miss landscape-level phenomena that drive ecosystem-level responses. We implemented an ecosystem-scale, nine-year nutrient experiment to examine how saltmarsh plants respond to simulated conditions of coastal eutrophication. Our study differed from previous saltmarsh enrichment studies in that we applied realistic concentrations of nitrate (70-100 µM NO3- ), the most common form of coastal nutrient enrichment, via tidal water at the ecosystem scale (~60,000 m2 creeksheds). Our enrichments added a total of 1,700 kg N·creek-1 ·yr-1 , which increased N loading 10-fold vs. reference creeks (low-marsh, 171 g N·m-2 ·yr-1 ; high-marsh, 19 g N·m-2 ·yr-1 ). Nutrients increased the shoot mass and height of low marsh, tall Spartina alterniflora; however, declines in stem density resulted in no consistent increase in aboveground biomass. High-marsh plants S. patens and stunted S. alterniflora did not respond consistently to enrichment. Nutrient enrichment did not shift community structure, contrary to the prediction of nutrient-driven dominance of S. alterniflora and Distichlis spicata over S. patens. Our mild responses may differ from the results of previous studies for a number of reasons. First, the limited response of the high marsh may be explained by loading rates orders of magnitude lower than previous work. Low loading rates in the high marsh reflect infrequent inundation, arguing that inundation patterns must be considered when predicting responses to estuarine eutrophication. Additionally, we applied nitrate instead of the typically used ammonium, which is energetically favored over nitrate for plant uptake. Thus, the form of nitrogen enrichment used, not just N-load, may be important in predicting plant responses. Overall, our results suggest that when coastal eutrophication is dominated by nitrate and delivered via flooding tidal water, aboveground saltmarsh plant responses may be limited despite moderate-to-high water-column N concentrations. Furthermore, we argue that the methodological limitations of nutrient studies must be considered when using results to inform management decisions about wetlands.

Belowground advantages in construction cost facilitate a cryptic plant invasion.

The energetic cost of plant organ construction is a functional trait that is useful for understanding carbon investment during growth (e.g. the resource acquisition vs. tissue longevity tradeoff), as well as in response to global change factors like elevated CO2 and N. Despite the enormous importance of roots and rhizomes in acquiring soil resources and responding to global change, construction costs have been studied almost exclusively in leaves. We sought to determine how construction costs of aboveground and belowground organs differed between native and introduced lineages of a geographically widely dispersed wetland plant species (Phragmites australis) under varying levels of CO2 and N. We grew plants under ambient and elevated atmospheric CO2, as well as under two levels of soil nitrogen. We determined construction costs for leaves, stems, rhizomes and roots, as well as for whole plants. Across all treatment conditions, the introduced lineage of Phragmites had a 4.3 % lower mean rhizome construction cost than the native. Whole-plant construction costs were also smaller for the introduced lineage, with the largest difference in sample means (3.3 %) occurring under ambient conditions. In having lower rhizome and plant-scale construction costs, the introduced lineage can recoup its investment in tissue construction more quickly, enabling it to generate additional biomass with the same energetic investment. Our results suggest that introduced Phragmites has had an advantageous tissue investment strategy under historic CO2 and N levels, which has facilitated key rhizome processes, such as clonal spread. We recommend that construction costs for multiple organ types be included in future studies of plant carbon economy, especially those investigating global change.

Phragmites australis management in the United States: 40 years of methods and outcomes.

Studies on invasive plant management are often short in duration and limited in the methods tested, and lack an adequate description of plant communities that replace the invader following removal. Here we present a comprehensive review of management studies on a single species, in an effort to elucidate future directions for research in invasive plant management. We reviewed the literature on Phragmites management in North America in an effort to synthesize our understanding of management efforts, identify gaps in knowledge and improve the efficacy of management. Additionally, we assessed recent ecological findings concerning Phragmites mechanisms of invasion and integrated these findings into our recommendations for more effective management. Our overall goal is to examine whether or not current management approaches can be improved and whether they promote reestablishment of native plant communities. We found: (i) little information on community-level recovery of vegetation following removal of Phragmites; and (ii) most management approaches focus on the removal of Phragmites from individual stands or groups of stands over a relatively small area. With a few exceptions, recovery studies did not monitor vegetation for substantial durations, thus limiting adequate evaluation of the recovery trajectory. We also found that none of the recovery studies were conducted in a landscape context, even though it is now well documented that land-use patterns on adjacent habitats influence the structure and function of wetlands, including the expansion of Phragmites. We suggest that Phragmites management needs to shift to watershed-scale efforts in coastal regions, or larger management units inland. In addition, management efforts should focus on restoring native plant communities, rather than simply eradicating Phragmites stands. Wetlands and watersheds should be prioritized to identify ecosystems that would benefit most from Phragmites management and those where the negative impact of management would be minimal.

Tidal marsh plant responses to elevated CO2 , nitrogen fertilization, and sea level rise.

Elevated CO2 and nitrogen (N) addition directly affect plant productivity and the mechanisms that allow tidal marshes to maintain a constant elevation relative to sea level, but it remains unknown how these global change drivers modify marsh plant response to sea level rise. Here we manipulated factorial combinations of CO2 concentration (two levels), N availability (two levels) and relative sea level (six levels) using in situ mesocosms containing a tidal marsh community composed of a sedge, Schoenoplectus americanus, and a grass, Spartina patens. Our objective is to determine, if elevated CO2 and N alter the growth and persistence of these plants in coastal ecosystems facing rising sea levels. After two growing seasons, we found that N addition enhanced plant growth particularly at sea levels where plants were most stressed by flooding (114% stimulation in the + 10 cm treatment), and N effects were generally larger in combination with elevated CO2 (288% stimulation). N fertilization shifted the optimal productivity of S. patens to a higher sea level, but did not confer S. patens an enhanced ability to tolerate sea level rise. S. americanus responded strongly to N only in the higher sea level treatments that excluded S. patens. Interestingly, addition of N, which has been suggested to accelerate marsh loss, may afford some marsh plants, such as the widespread sedge, S. americanus, the enhanced ability to tolerate inundation. However, if chronic N pollution reduces the availability of propagules of S. americanus or other flood-tolerant species on the landscape scale, this shift in species dominance could render tidal marshes more susceptible to marsh collapse.

Jack-and-master trait responses to elevated CO2 and N: a comparison of native and introduced Phragmites australis.

Global change is predicted to promote plant invasions world-wide, reducing biodiversity and ecosystem function. Phenotypic plasticity may influence the ability of introduced plant species to invade and dominate extant communities. However, interpreting differences in plasticity can be confounded by phylogenetic differences in morphology and physiology. Here we present a novel case investigating the role of fitness trait values and phenotypic plasticity to global change factors between conspecific lineages of Phragmites australis. We hypothesized that due to observed differences in the competitive success of North American-native and Eurasian-introduced P. australis genotypes, Eurasian-introduced P. australis would exhibit greater fitness in response to global change factors. Plasticity and plant performance to ambient and predicted levels of carbon dioxide and nitrogen pollution were investigated to understand how invasion pressure may change in North America under a realistic global change scenario. We found that the introduced Eurasian genotype expressed greater mean trait values in nearly every ecophysiological trait measured--aboveground and belowground--to elevated CO(2) and nitrogen, outperforming the native North American conspecific by a factor of two to three under every global change scenario. This response is consistent with "jack and master" phenotypic plasticity. We suggest that differences in plant nitrogen productivity, specific leaf area, belowground biomass allocation, and inherently higher relative growth rate are the plant traits that may enhance invasion of Eurasian Phragmites in North America. Given the high degree of genotypic variability within this species, and our limited number of genotypes, our results must be interpreted cautiously. Our study is the first to demonstrate the potential importance of jack-and-master phenotypic plasticity in plant invasions when facing imminent global change conditions. We suggest that jack-and-master invasive genotypes and/or species similar to introduced P. australis will have an increased ecological fitness, facilitating their invasion in both stressful and resource rich environments.

Effects of cadmium and zinc on larval growth and survival in the ground beetle, Pterostichus oblongopunctatus.

Carabid beetles, like Pterostichus oblongopunctatus, living in metal contaminated areas may be exposed to elevated levels of metals within their diets. However, when compared to other second order consumers, they have one of the lowest observed levels of metals, indicating methods of detoxification to deal with such toxicants. In this study, we investigated if chronic, multigenerational exposure to metals leads to resistance to toxic metal concentrations, and if so, what are the costs associated with them. Adult organisms were collected from two sites, a polluted and a reference site near Olkusz, in southern Poland. These adults were immediately mated, and eggs were collected twice weekly to assess the effects in the larvae of the F(1) generation. Larvae were randomly exposed to one of four artificial mediums: control, 50 mg kg(-1) Cd, 500 mg kg(-1) Zn, and a combined treatment of 50 mg kg(-1) Cd and 500 mg kg(-1) Zn to investigate possible interactions. Individuals were sacrificed at 10, 30, and 40 days. Although metals were not accumulated in larvae (p>0.05), larvae fed the Cd or the Zn treatment grew significantly slower, and had the lowest survival rate (p<0.05) in respect to control. Out of metal treated animals, those on the combined treatment of CdZn grew the quickest and had the highest observed survival (p<0.05). Although previous studies have demonstrated changes in adult population parameters under chronic, multigeneration exposure to toxic metal concentrations, our study did not reveal any changes in the larval stage.