Rhizodeposition of organic carbon by plants with contrasting traits for resource acquisition: responses to different fertility regimes

Background and aims Rhizodeposition plays an important role in mediating soil nutrient availability in ecosystems. However, owing to methodological difficulties (i.e., narrow zone of soil around roots, rapid assimilation by soil microbes) fertility-induced changes in rhizodeposition remain mostly unknown. Methods We developed a novel long-term continuous 13C labelling method to address the effects of two levels of nitrogen (N) fertilization on rhizodeposited carbon (C) by species with different nutrient acquisition strategies. Results Fertility-induced changes in rhizodeposition were modulated by root responses to N availability rather than by changes in soil microbial biomass. Differences among species were mostly related to plant biomass: species with higher total leaf and root biomass also had higher total rhizodeposited C, whereas species with lower root biomass had higher specific rhizodeposited C (per gram root mass). Experimental controls demonstrated that most of the biases commonly associated with this type of experiment (i.e., long-term steady-state labelling) were avoided using our methodological approach. Conclusions These results suggest that the amount of rhizodeposited C from plants grown under different levels of N were driven mainly by plant biomass and root morphology rather than microbial biomass. They also underline the importance of plant characteristics (i.e., biomass allocation) as opposed to traits associated with plant resource acquisition strategies in predicting total C rhizodeposition.

Past, present and future - assessing changes in aquatic environments using subfossil diatoms

The deterioration of surface waters is one of the most important issues in the environmental management of the European Union. Thus, the EU Water Framework Directive 2000/60/EC (WFD) requires “good ecological and chemical status” of surface waters by 2015 allowing only a slight departure from ecological reference conditions characterized by the biological communities typical for the conditions of minimal anthropogenic impact. The WFD requires the determination of ecological reference conditions and the present ecological status of surface waters. To meet this legislative demand, sedimentary diatom assemblages were used in these studies with various methods 1) to assess natural and human activity induced environmental changes, 2) to characterize background conditions 3) to evaluate the present ecological status and 4) to predict the future of the water bodies in the light of palaeolimnological data. As the WFD refers to all surface waters, both coastal and inland sites were included. Two long and two short sediment cores from the Archipelago Sea in the northern Baltic Sea were examined for their siliceous microfossils in order to assess (1) the Holocene palaeoenvironmental history and (2) the recent eutrophication of the area. The diatom record was divided into local diatom assemblage zones (LDAZ, long cores) and diatom assemblage zones (DAZ, short cores). Locally weighted weighted averaging regression and calibration (LWWA) was applied for the quantitative reconstruction of past TN concentrations (short cores). An age model for the long cores was constructed by using independent palaeomagnetic and AMS-14C methods. The short cores were dated using radiometric (210Pb, 226Ra and 137Cs) methods. The long cores date back to the early history of the Archipelago Sea, which was freshwater – no salinity increase referable to the brackish phase of the Yoldia Sea is recognized. The nutrient status of the lacustrine phase was slightly higher in the Archipelago Sea than in the Baltic Proper. Initial brackish-water influence is observed at 8 150 ±80 cal. BP (LDAZ4), but fully brackish conditions were established at 7 700 ±80 cal. BP (LDAZ5). The diatom assemblages indicate increasing salinity, warming climate and possible eutrophic conditions during the lacustrine to brackish-water transition. The decreasing abundance of Pseudosolenia calcar-avis (Schultze) Sundström and the increasing abundance of the ice-cover indicator species Pauliella taeniata (Grunow) Round and Basson indicate decreasing salinity and climatic cooling after ca. 5 000 cal. BP. Signs of eutrophication are visible in the most recent diatom assemblage zones of both short cores. Diatom-inferred total nitrogen (DI-TN) reconstructions partially fail to trace the actual measured total nitrogen concentrations especially from the late 1980s to the mid 1990s. This is most likely due to the dominating diatom species Pauliella taeniata, Thalassiosira levanderi Van Goor and Fragilariopsis cylindrus (Grunow) W. Krieger being more influenced by factors such as the length of the ice-season rather than nutrient concentrations. It is concluded that the diatom assemblages of the study sites are principally governed by climate fluctuations, with a slight influence of eutrophication visible in the most recent sediments. There are indications that global warming, with reduced ice cover, could impact the spring blooming diatom species composition in the Archipelago Sea. In addition, increased sediment accumulation in the early 90s coincides with the short ice-seasons suggesting that warming climate with decreasing ice-cover may increase sedimentation in the study area. The diverse diatom assemblages dominated by benthic species (54 %) in DAZ1 in the Käldö Fjärd core can be taken as background diatom assemblages for the Archipelago Sea. Since then turbidity has increased and the diatom assemblages have been dominated by planktonic diatoms from around the mid 1800s onwards. The reconstructed reference conditions for the total nitrogen concentrations fluctuate around 400 μg l-1. Altogether two short sediment cores and eight short cores for top-bottom analysis were retrieved from Lake Orijärvi and Lake Määrjärvi to assess the impact of the acid mine drainage (AMD) derived metals from the Orijärvi mine tailings on the diatom communities of the lakes. The Cu (Pb, Zn) mine of Orijärvi (1757 – 1956) was the first one in Finland where flotation techniques (1911 – 1955) were used to enrich ore and large quantities of tailings were produced. The AMD derived metal impact to the lakes was found to be among the heaviest thus far recorded in Finland. Concentrations of Cu, Pb and Zn in Lake Orijärvi sediments are two to three orders of magnitude higher than background values. The metal inputs have affected Lake Orijärvi and Lake Määrjärvi diatom communities at the community levels through shifts in dominant taxa (both lakes) and at the individual level through alteration in frustule morphology (Lake Orijärvi). At present, lake water still has elevated heavy metal levels, indicating that the impact from the tailings area continues to affect both lakes. Lake Orijärvi diatom assemblages are completely dominated by benthic species and are lacking planktonic diatoms. In Lake Määrjärvi the proportion of benthic and tychoplanktonic diatoms has increased and the planktonic taxa have decreased in abundance. Achnanthidium minutissimum Kützing and Brachysira vitrea (Grun.) R. Ross in Hartley were the most tolerant species to increased metal concentrations. Planktonic diatoms are more sensitive to metal contamination than benthic taxa, especially species in the genus Cyclotella (Kützing) Brébisson. The ecological reference conditions assessed in this study for Lake Orijärvi and Lake Määrjärvi comprise diverse planktonic and benthic communitites typical of circumneutral oligotrophic lakes, where the planktonic diatoms belonging to genera Cyclotella , Aulacoseira Thwaites, Tabellaria Ehrenberg and Asterionella Hassall dominate in relative abundances up to ca. 70%. The benthic communities are more diverse than the planktonic consisting of diatoms belonging to the genera Achnanthes Bory, Fragilaria Lyngbye and Navicula St. Vincent. This study clearly demonstrates that palaeolimnological methods, especially diatom analysis, provide a powerful tool for the EU Water Frame Work Directive for defining reference conditions, natural variability and current status of surface waters. The top/bottom approach is a very useful tool in larger-scale studies needed for management purposes. This “before and after” type of sediment sampling method can provide a very time and cost effective assessment of ecological reference conditions of surface waters.

Dry biomass distribution in a cerrado sensu stricto site in Brazil central

The Cerrado has been the main source of firewood and charcoal in Brazil, but despite being one of the hot spots for conservation of the world's biodiversity, neither plantations of native species nor sustainable management has been adopted in the region. The aim of this work was to investigate the biomass distribution and the potential for energy production of the cerrado species. The study was conducted in a cerrado sensu stricto site at the Água Limpa Farm (15º 56'14'' S and 47º 46'08'' W) in the Cerrado Biosphere Reserve. An area of 63.54ha was divided in 20 x 50m plots and, a random sample consisting of ten of these plots, representing 1.56% of the study-site, was assessed. All woody individuals from 5 cm diameter at 30 cm above ground level were identified and measured. Each individual was felled, the twigs thinner than 3cm were discarded while the larger branches and the trunks, both with bark, were weighted separately. After that, 2.5cm transverse sections of the trunk with bark were taken at 0, 25, 50, 75 and 100% of the length. A similar sample was also taken at the base of each branch. A total of 47 species in 35 genera and 24 families were found, with an average density of 673 individuals per ha. The diameter distribution showed a reversed-J shape with 67% of the individuals up to 13cm, while the maximum diameter was 32.30cm. Seven species represented 72% of the total biomass. In general, the species with higher production per tree were among those with higher production per ha. This content was distributed by diameter classes, reaching a maximum of 2.5ton/ha between 9 to 13cm and then, decreasing to 0.96 ton/ha between 29 to 33cm diameter. Carbon sequestering was 6.2ton/ha (until the actual stage of cerrado) based on an average 50% carbon content in the dry matter. The heat combustion of the wood varied from 18,903kj/kg to 20,888kj/kg with an average of 19,942kj/kg. The smaller diameter classes fix more carbon due to the large number of small plants per ha. But, for a species that reached larger dimensions and contained individuals in all diameter classes, Vochysia thyrsoidea, one can verify an increase in carbon fixation from 1.41 kg/ha in the first class (5 to 9cm) to 138,3kg/ha in the last (25 to 33cm). That indicates that it is possible to select species that reach larger size with a higher capacity of carbon accumulation per plant. The species that reached larger dimensions, with a production per tree above average and had high calorific power values were Dalbergia miscolobium, Pterodon pubescens and Sclerolobium paniculatum. These species have potential for use in fuelwood plantations and sustainable management.

Estrutura e relações ambientais de grupos florísticos em fragmento de floresta estacional subtropical

Este estudo teve por objetivos analisar a estrutura de grupos florísticos na vegetação arbórea-arbustiva de um trecho de encosta de Floresta Estacional Subtropical, no Parque Estadual Quarta Colônia, RS, e determinar fatores ambientais importantes na ocorrência das espécies, a fim de subsidiar programas de restauração de áreas degradadas como ecossistema de referência. Para isso, foi realizado estudo fitossociológico em 12 parcelas de 200 m² (10 x 20 m), distantes 20 m entre si, distribuídas sistematicamente em quatro faixas de 100 m entre si, seguindo gradiente de topografia. Em cada parcela foram realizadas a identificação e a medição da circunferência à altura do peito (CAP) e da altura total dos indivíduos arbóreos e arbustivos, ou seja, com CAP > 15 cm, denominado componente arbóreo. Os dados do sub-bosque foram obtidos em parcelas circulares com raio de 1,78 m, instaladas no centro das parcelas de 10 x 20 m, medindo-se indivíduos com altura > 30 cm e CAP <15 cm. As variáveis ambientais foram compostas pela declividade, pela intensidade luminosa, pelas características químicas e textura do solo. Os dados foram analisados segundo a estrutura e diversidade da vegetação por meio da análise de agrupamento pelo método TWINSPAN (Two-way Indicator Species Analysis) e da ordenação pelo método CCA (Análise de Correspondência Canônica). Foi identificada a formação de dois grupos florísticos. A composição de espécies de cada grupo pertence a diferentes estágios de sucessão natural, conforme histórico de uso agrícola, e, segundo suas características funcionais, podem ser indicadas para programas de restauração florestal.

Leaching of Picloram in ultisol under different rainfall volumes

The present study aimed to evaluate the leaching potential of Picloram in Ultisol columns under different rainfall amounts. For such, 30 treatments were evaluated (one soil associated with three levels of rainfall and ten depths).The experiments were arranged in a split-plot design, in a completely randomized design, with four replications. PVC columns of 10 cm in diameter and 50 cm in length were filled with these soils, moistened, and placed upright for 48 hours to drain the excess water. The herbicide was applied and rainfall simulations were carried out at specified intensities, according to the treatments, to force Picloram leaching. After 72 hours, all the columns were arranged in a horizontal position and opened lengthwise. Then, soil sampling was carried out every 5 cm of depth for subsequent herbicide extraction and quantification and analysis by high performance liquid chromatography. The remaining soil samples were placed in plastic pots, and, at the respective depths, the indicator species Cucumis sativus was sown. Twenty-one days after the emergence (DAE) of the indicator plants, evaluations were conducted to verify the symptoms of toxicity caused by Picloram in the plants. It was concluded that Picloram leaching is directly dependent on the volume of rain applied. The herbicide reached the deepest regions in the soil with the highest intensity of rain. The results obtained by bioassay were in agreement with those found by liquid chromatography.

Sorption of oxadiazon in soils cultivated in the brazilian cerrado

The objective of this study was to evaluate oxadiazon sorption in different soils of the Brazilian Cerrado, highlighting the correlations of lethal doses of this herbicide capable of inhibiting 50% of the dry matter accumulation of the bio-indicator (LD50) among the chemical characteristics of the soil and its direct and indirect effects. The experiment was carried out in a greenhouse in a randomized block design and four repetitions. Each experimental unit consisted of a pot with increasing rates of oxadiazon and oat (Avena sativa), as the bio-indicator species. For sorption evaluation, washed sand and 22 soils (substrates) from Cerrado Brazilian's Alliaceae cultivated areas were used. LD50 and sorption ratio (SR) = [(LD50soil - LD50sand)/LD50sand] to the substrates were determined. Pearson correlation analysis was performed between the chemical characteristics of the substrates and the LD50 of oxadiazon. A path analysis was quantified, to deploy only the significant correlations estimated in direct and indirect effects of the characters on LD50, which is a basic variable. A more pronounced LD50 (528.09 g ha-1) for the Cerrado soil sample resulted in higher SR (> 53.00), while in the washed sand substrate, LD50 corresponded only to 9.74 g ha-1 of the oxadiazon (available in soil). It was concluded that oxadiazon sorption is influenced by the chemical characteristics of the soils, highlighting the correlation with pH (CaCl2), magnesium content, aluminum, organic matter, organic carbon, and aluminum saturation.

The use of diatom assemblages to determine paleotrophic changes in lakes

The purpose of this study was to develop a classifi cation scheme for l ake trophic status based on the relative abundance of l ake sediment diatom trophic indicator species. A total of 600 diatom frustules were counted from the surface sediments of e a ch of 30 lakes selected to repr e seni~ a continuum from u.lt ra-oligotrophic t,o fairly eutrophic but not hype r-' eutrophic conditions. Published autecological information was used to determine the trophic indicator status of each of the s pecies. A quotieht was derived by dividing the s um of all the e utrophic indicator species by the sum of all oligotrophic indicai.-:.or species. Oligo'- mesotrophic. mesotrophic and meso-eutrophic species were added to both the numerator and denomina tor. Five categories of diatom i.nferred trophic status were recognized : ultra-oligotrophic - 0'-0.2:3, oligotrophic::: 0.24-0 . 70, mesotrophic :: 0.'71 -0.99, meso-elxtrophic :: 1. 00-1. '78 and eutrophic:: 1.. 79-2. 43. But only three of these (oligotrophic:: 0-0.69, mesotrophic ::: 0 . 70'-1.69 j and eutrophic:: 1.70-2.50) proved usef ul. The present study of the relationship between diatom inferred trophic status and the literature-derived trophic status of SO lake s (which were purposely chosen to represent a broad spectrum of lakes in Canada) indicated that: 1) Based on diatom species (assemblages ) it is possible to segregate the lakes from which. th",)se diatoms were taken into three basic categories : o ligotrophic, mesotrophic and eutrophic lake types. ~~) It was not possible t,o separate meso-eutrophic and o l igo-mesotrophic lakes f rom mesotrophic l akes as the the degree of overlap betwee n the diat,om species in these lake types was extremely high. 3) Ha d mo r e ul tra-oligotroph,ic lakes been sampled it might have been possible to more a ccurately s eparate them f rom oligotrophic Jakes. 4 ) Had. more humic lakes been sampled in this s tudy I f eel it would have been possible to identify a unique diatom a ssemblage which would h a ve chara cterized t his lake type . Re gression analyses were performed using the aforementioned diatom inferred trophic index as a f unction of 1) log Sec chi transparency (r = - 0.70) 2) total phosphorus (r = 0. 77 ) and 3) chlorophyll-a (r = 0.74). Once e ach of these rel ationships had been established , it was possible to infer paleotrophic (downcore) changes in an oligotrophic lake (Barbara Lake) and in a eutrophic lake (Chemung Lake) . Barbara Llake was dominated by oligotrophic s pecies and remained oligotrophic throughout the 200-·year history r epresented by i t s 32 em long sediment core. Chemung Lake is currently dominated. by eutrophic species but went through a mesotrophic st,age which was associated with a rise in the water level of the lake followi n g dam construction in its watershed in the early 1.900 J ::;. This was followed by its reversion to it,s present day eutrophic stage (dominated by eutrophic species) possibly as a r esult of shallowing process which can be attributed to " silting' up" of the reservoir and the invasion of the l ake by Myriophyllum spjcatum (Eurasian milfoil) i n the 1970's . In addition, nutrient .:r':l.ch run"'offwhich resulted from increased human a.ctivities associated with cottage development along its shores has contribut ed to its eutrophication. There is some evidence that the rat,e o :f its prog ressive eutrophication has declined during the last decade. This was correlated with legislation enacted in the 60's and 70's in Ontario which was aimed at reducing nutrient loading from cottages.

Short-term Response of Breeding Barred Owls to Forestry in a Boreal Mixedwood Forest Landscape

Forestry and other activities are increasing in the boreal mixedwood of Alberta, with a concomitant decrease in older forest. The Barred Owl (Strix varia) is an old-growth indicator species in some jurisdictions in North America. Hence, we radio-tagged Barred Owls in boreal mixedwood in Alberta to determine whether harvesting influenced habitat selection. We used three spatial scales: nest sites, i.e., nest tree and adjacent area of 11.7 m radius around nests, nesting territory of 1000 m radius around nests, and home range locations within 2000 m radius of the home range center. Barred Owls nested primarily in balsam poplar (Populus balsamifera) snags > 34 cm dbh and nest trees were surrounded by large, > 34 cm dbh, balsam poplar trees and snags. Nesting territories contained a variety of habitats including young < 80-yr-old, deciduous-dominated stands, old deciduous and coniferous-dominated stands, treed bogs, and recent clear-cuts. However, when compared to available habitat in the study area, they were more likely to contain old conifer-dominated stands and recent cutblocks. We assumed this is because all of the recent harvest occurred in old stands, habitat preferred by the owls. When compared with random sites, locations used for foraging and roosting at the home range scale were more likely to be in young deciduous-dominated stands, old conifer-dominated stands and cutblocks > 30 yr old, and less likely to occur in old deciduous-dominated stands and recent cutblocks. Hence, although recent clearcuts occurred in territories, birds avoided these microhabitats during foraging. To meet the breeding requirements of Barred Owls in managed forests, 10–20 ha patches of old deciduous and mixedwood forest containing large Populus snags or trees should be maintained. In our study area, nest trees had a minimum dbh of 34 cm. Although cut areas were incorporated into home ranges, the amount logged was low, i.e., 7%, in our area. Hence more research is required to determine harvest levels tolerated by owls over the long term.

Off-road sampling reveals a different grassland bird community than roadside sampling: implications for survey design and estimates to guide conservation

Grassland bird species continue to decline steeply across North America. Road-based surveys such as the North American Breeding Bird Survey (BBS) are often used to estimate trends and population sizes and to build species distribution models for grassland birds, although roadside survey counts may introduce bias in estimates because of differences in habitats along roadsides and in off-road surveys. We tested for differences in land cover composition and in the avian community on 21 roadside-based survey routes and in an equal number of adjacent off-road walking routes in the grasslands of southern Alberta, Canada. Off-road routes (n = 225 point counts) had more native grassland and short shrubs and less fallow land and road area than the roadside routes (n = 225 point counts). Consequently, 17 of the 39 bird species differed between the two route types in frequency of occurrence and relative abundance, measured using an indicator species analysis. Six species, including five obligate grassland species, were more prevalent at off-road sites; they included four species listed under the Canadian federal Species At Risk Act or listed by the Committee on the Status of Endangered Wildlife in Canada: Sprague’s Pipit (Anthus spragueii), Baird’s Sparrow (Ammodramus bairdii), the Chestnut-collared Longspur (Calcarius ornatus), and McCown’s Longspur (Rhynchophanes mccownii). The six species were as much as four times more abundant on off-road sites. Species more prevalent along roadside routes included common species and those typical of farmland and other human-modified habitats, e.g., the European Starling (Sturnus vulgaris), the Black-billed Magpie (Pica hudsonia), and the House Sparrow (Passer domesticus). Differences in avian community composition between roadside and off-road surveys suggest that the use of BBS data when generating population estimates or distribution models may overestimate certain common species and underestimate others of conservation concern. Our results highlight the need to develop appropriate corrections for bias in estimates derived from roadside sampling, and the need to design surveys that sample bird communities across a more representative cross-section of the landscape, both near and far from roads. 

Does changing the taxonomical resolution alter the value of soil macroinvertebrates as bioindicators of metal pollution?

Ecological indicators are taxa that are affected by, and indicate effects of, anthropogenic environmental stress or disturbance on ecosystems. There is evidence that some species of soil macrofauna (i.e. diameter > 2 min) constitute valuable biological indicators of certain types of soil perturbations. This study aims to determine which level of taxonomic resolution, (species, family or ecological group) is the best to identify indicator of soil disturbance. Macrofauna were sampled in a set of sites encompassing different land-use systems (e.g. forests, pastures, crops) and different levels of pollution. Indicator taxa were sought using the IndVal index proposed by Dufrene and Legendre [Dufrene, M., Legendre, P., 1997. Species assemblages and indicator species: the need for a flexible asymetrical approach. Ecological Monographs 67, 345-3661. This approach is based on a hierarchical typology of sites. The index value changes along the typology and decreases (increases) for generalist (specialist) faunal units (species, families or ecological groups). Of the 327 morphospecies recorded, 19 were significantly associated with a site type or a group of sites (5.8%). Similarly, species were aggregated to form 59 families among which 17 (28.8%) displayed a significant indicator value. Gathering species into 28 broad ecological assemblages led to 14 indicator groups (50%). Beyond the simple proportion of units having significant association with a given level of the site typology, the proportion of specialist and generalist groups changed dramatically when the level of taxonomic resolution was altered. At the species level 84% of the indicator units were specialist, whereas this proportion decreased to 70 and 43% when families and ecological groups were considered. Because specialist groups are the most interesting type of indicators either in terms of conservation or for management purposes we come to the conclusion that the species level is the most accurate taxonomic level in bioindication studies although it requires a high amount of labour and operator knowledge and is time-consuming. (c) 2005 Published by Elsevier Ltd.

Landscape typology in the Mediterranean context: a tool for habitat restoration

Despite the wide use of Landscape Character Assessment (LCA) as a tool for landscape planning in NW Europe, there are few examples of its application in the Mediterranean. This paper reports on the results from the development of a typology for LCA in a study area of northern Sardinia, Italy to provide a spatial framework for the analysis of current patterns of cork oak distribution and future restoration of this habitat. Landscape units were derived from a visual interpretation of map data stored within a GIS describing the physical and cultural characteristics of the study area. The units were subsequently grouped into Landscape Types according to the similarity of shared attributes using Two Way Indicator Species Analysis (TWINSPAN). The preliminary results showed that the methodology classified distinct Landscape Types but, based on field observations, there is a need for further refinement of the classification. The distribution and properties of two main cork oak habitats types was examined within the identified Landscape Types namely woodlands and wood pastures using Patch Analyst. The results show very clearly a correspondence between the distribution of cork oak pastures and cork oak woodland and landscape types. This forms the basis of the development of strategies for the maintenance, restoration and recreation of these habitat types within the study area, ultimately for the whole island of Sardinia. Future work is required to improve the landscape characterisation , particularly with respect to cultural factors, and to determine the validity of the landscape spatial framework for the analysis of cork oak distribution as part of a programme of habitat restoration and re-creation.

Linking molecular and population stress responses in Daphnia magna exposed to cadmium

DNA microarrays can be used to measure environmental stress responses. If they are to be predictive of environmental impact, we need to determine if altered gene expression translates into negative impacts on individuals and populations. A large cDNA microarray (14000 spots) was created to measure molecular stress responses to cadmium in Daphnia magna,the most widely used aquatic indicator species, and relate responses to population growth rate (pgr). We used the array to detect differences in the transcription of genes in juvenile D. magna (24 h old) after 24 h exposure to a control and three cadmium concentrations (6, 20, and 37 mu g Cd2+ L-1). Stress responses at the population level were estimated following a further 8 days exposure. Pgr was approximately linear negative with increasing cadmium concentration over this range. The microarray profile of gene expression in response to acute cadmium exposure begins to provide an overview of the molecular responses of D. magna, especially in relation to growth and development. Of the responding genes, 29% were involved with metabolism including carbohydrate, fat and peptide metabolism, and energy production, 31% were involved with transcription/translation, while 40% of responding genes were associated with cellular processes like growth and moulting, ion transport, and general stress responses (which included oxidative stress). Our production and application of a large Daphnia magna microarray has shown that measured gene responses can be logically linked to the impact of a toxicant such as cadmium on somatic growth and development, and consequently pgr.

Towards a population ecology of stressed environments: the effects of zinc on the springtail Folsomia candida

1. To understand population dynamics in stressed environments it is necessary to join together two classical lines of research. Population responses to environmental stress have been studied at low density in life table response experiments. These show how the population's growth rate (pgr) at low density varies in relation to levels of stress. Population responses to density, on the other hand, are based on examination of the relationship between pgr and population density. 2. The joint effects of stress and density on pgr can be pictured as a contour map in which pgr varies with stress and density in the same way that the height of land above sea level varies with latitude and longitude. Here a microcosm experiment is reported that compared the joint effects of zinc and population density on the pgr of the springtail Folsomia candida (Collembola). 3. Our experiments allowed the plotting of a complete map of the effects of density and a stressor on pgr. Particularly important was the position of the pgr= 0 contour, which suggested that carrying capacity varied little with zinc concentration until toxic levels were reached. 4. This prediction accords well with observations of population abundance in the field. The method also allowed us to demonstrate, simultaneously, hormesis, toxicity, an Allee effect and density dependence. 5. The mechanisms responsible for these phenomena are discussed. As zinc is an essential trace element the initial increase in pgr is probably a consequence of dietary zinc deficiency. The Allee effect may be attributed to productivity of the environment increasing with density at low density. Density dependence is a result of food limitation. 6. Synthesis and applications. We illustrate a novel solution based on mapping a population's growth rate in relation to stress and population density. Our method allows us to demonstrate, simultaneously, hormesis, toxicity, an Allee effect and density dependence in an important ecological indicator species. We hope that the approach followed here will prove to have general applicability enabling predictions of field abundance to be made from estimates of the joint effects of the stressors and density on population growth rate.

The joint effects of larval density and C-14-cypermethrin on the life history and population growth rate of the midge Chironomus riparius

1. Chemical effects on organisms are typically assessed using individual-level endpoints or sometimes population growth rate (PGR), but such measurements are generally made at low population densities. In contrast most natural populations are subject to density dependence and fluctuate around the environmental carrying capacity as a result of individual competition for resources. As ecotoxicology aims to make reliable population projections of chemical impacts in the field, an understanding of how high-density or resource-limited populations respond to environmental chemicals is essential. 2. Our objective was to determine the joint effects of population density and chemical stress on the life history and PGR of an important ecotoxicological indicator species, Chironomus riparius, under controlled laboratory conditions. Populations were fed the same ration but initiated at different densities and exposed to a solvent control and three concentrations of C-14-cypermethrin in a sediment-water test system for 67 days at 20 +/- 1 degreesC. 3. Density had a negative effect on all the measured life-history traits, and PGR declined with increasing density in the controls. Exposure to C-14-cypermethrin had a direct negative effect on juvenile survival, presumably within the first 24 h because the chemical rapidly dissipated from the water column. Reductions in the initial larval densities resulted in an increase in the available resources for the survivors. Subsequently, exposed populations emerged sooner and started producing offspring earlier than the controls. C-14-cypermethrin had no effect on estimated fecundity and adult body weight but interacted with density to reduce the time to first emergence and first reproduction. As a result, PGR increased with cypermethrin concentration when populations were initiated at high densities. 4. Synthesis and applications. The results showed that the effects of C-14-cypermethrin were buffered at high density, so that the joint effects of density and chemical stress on PGR were less than additive. Low levels of chemical stressors may increase carrying capacity by reducing juvenile competition for resources. More and perhaps fitter adults may be produced, similar to the effects of predators and culling; however, toxicant exposure may result in survivors that are less tolerant to changing conditions. If less than additive effects are typical in the field, standard regulatory tests carried out at low density may overestimate the effects of environmental chemicals. Further studies over a wide range of chemical stressors and organisms with contrasting life histories are needed to make general recommendations.

The influence of larval density, food availability and habitat longevity on the life history and population growth rate of the midge Chironomus riparius

Despite long-standing interest in the forms and mechanisms of density dependence, these are still imperfectly understood. However, in a constant environment an increase in density must reduce per capita resource availability, which in turn leads to reduced survival, fecundity and somatic growth rate. Here we report two population experiments examining the density dependent responses under controlled conditions of an important indicator species, Chironomus riparius. The first experiment was run for 35 weeks and was started at low density with replicate populations being fed three different rations. Increased ration reduced generation time and increased population growth rate (pgr) but had no effect on survival, fecundity and female body weight in the first generation. In the second generation there was a six-fold increase in generation time, presumably due to the greatly reduced per capita resource availability as the estimated initial densities of the second generation were 300 times greater than the first. Juvenile survival to emergence, fecundity, adult body weight and pgr declined by 90%, 75%, 35% and 99%, respectively. These large between-generation effects may have obscured the effects of the threefold variation in ration, as only survival to emergence significantly increased with ration in the second generation. These results suggest that some chironomid larvae survive a reduction in resource availability by growing more slowly. In the ephemeral habitats sometimes occupied by C. riparius, the effects of population density may depend crucially on the longevity of the environment. A second experiment was therefore performed to measure pgr from six different starting densities over an eight-week period. The relationship between pgr and density was concave, viewed from above. At densities above 16 larvae per cm(2), less than 1% of the population emerged and no offspring were produced. Under the conditions of experiment 2 - an 8-week habitat lifespan carrying capacity was estimated as 8 larvae per cm(2).