External nutrient inputs into terrestrial ecosystems of the Falkland Islands and the Maritime Antarctic region

Antarctic terrestrial ecosystems are nutrient-poor and depend for their functioning in part on external nutrients. However, little is known about the relative importance of various sources. We measured external mineral nutrient sources (wind blown material, precipitation and guano) at three locations, the cold temperate oceanic Falkland Islands (51°76′S), and the Maritime Antarctic Signy (60°71′S) and Anchorage Islands (67°61′S). These islands differ in the level of vegetation development through different environmental constraints and historical factors. Total mineral nitrogen input differed considerably between the islands. During the 3 month summer period it amounted to 18 mg N m⁻² on the Falkland Islands and 6 and 102 mg N m⁻² at Signy and Anchorage Islands, respectively. The high value for Anchorage was a result of guano deposition. By measuring stable isotopic composition (δ¹⁵N) of the different nitrogen sources and the dominant plant species, we investigated the relative utilisation of each source by the vegetation at each island. We conclude that external mineral nitrogen inputs to Antarctic terrestrial ecosystems show great spatial variability, with the local presence of bird (or other vertebrate) colonies being particularly significant.     

Mercury, cadmium and lead accumulation in Antarctic mosses growing along nutrient and moisture gradients

Accumulation of Hg, Cd and Pb by moss was studied in a coastal ice-free area (Edmonson Point, northern Victoria Land) in relation to the water and nutrient availability and substratum characteristics. Although metal concentrations in surface soils were among the lowest ever reported from remote areas, those of Hg and Cd in mosses were higher, being in the same range as those usually reported in regional surveys in the northern hemisphere. By contrast, Antarctic mosses showed very low Pb concentrations, and no impact from local human activities was detected. Marine aerosols, seabird guano and volcanic emissions appeared to be the more probable sources of Cd and Hg. Besides atmospheric deposition, the main pathway of metals to mosses was probably through evapo-transpiration at their surface which determines an upward migration of ions and their bioaccumulation. Received: 8 September 1997 / Accepted: 4 January 1998     

Nutrient stress gradient in the bottom 5 cm of fast ice, McMurdo Sound, Antarctica

One-centimeter-scale vertical sampling of fast ice from McMurdo Sound, Antarctica reveals evidence of progressive nutrient limitation with distance above the ice/water interface. Over the bottom 6 cm photosynthetically active radiation increases by between 1.8 and 3 times, C:N increases from 6.8 to 19.8 and δ¹³C increases from −18 to −12. Fatty acid composition also changes with a consistent decline in polyunsaturated fatty acids and a rise in saturated fatty acids. These factors all suggest severe and progressive nutrient limitation with distance from the ice/water interface. Accepted: 5 September 1998     

Periphyton nutrient status in a temperate stream with mixed land-uses: implications for watershed nitrogen storage

We sampled periphyton communities in a highly productive stream to characterize how longitudinal changes in watershed geology and land use affect periphyton nutrient status and elemental composition. Nutrient status was evaluated from measures of periphyton nutrient composition (carbon, nitrogen, and phosphorus), stable isotope signatures (δ¹⁵N and δ¹³C), and the response of periphyton to experimental enrichment with nitrogen. Biomass and nutrient content increased dramatically from the headwaters to downstream, while tissue nutrient ratios (C:P and C:N) were more consistent and did not indicate strong N- or P-limitation. Nitrogen enrichment experiments did not exhibit a consistent response upstream or downstream, and periphyton C:N:P stoichiometry showed no significant response to N-enrichment. Absolute densities of periphyton N were 5- to 90-fold greater than the overlying N concentrations in stream water (159- to 353-fold greater for P), and the δ¹⁵N signal indicates downstream enrichment from likely watershed sources (urban and agriculture land-use). These results suggest that periphyton in Spring Creek are not N-limited and store large quantities of both N and P, which in turn can be transported downstream during high flow events. Handling editor: David Hamilton     

Ecological Stoichiometry and Density Responses of Plant-Arthropod Communities on Cormorant Nesting Islands

Seabirds deposit large amounts of nutrient rich guano on their nesting islands. The increased nutrient availability strongly affects plants and consumers. Consumer response differs among taxonomic groups, but mechanisms causing these differences are poorly understood. Ecological stoichiometry might provide tools to understand these mechanisms. ES suggests that nutrient rich taxa are more likely to be nutrient limited than nutrient poorer taxa and are more favored under nutrient enrichment. Here, we quantified differences in the elemental composition of soil, plants, and consumers between islands with and without nesting cormorant colonies and tested predictions made based on ES by relating the elemental composition and the eventual mismatch between consumer and resource stoichiometry to observed density differences among the island categories. We found that nesting cormorants radically changed the soil nutrient content and thereby indirectly plant nutrient content and resource quality to herbivores. In contrast, consumers showed only small differences in their elemental composition among the island categories. While we cannot evaluate the cause of the apparent homeostasis of invertebrates without additional data, we can conclude that from the perspective of the next trophic level, there is no difference in diet quality (in terms of N and P content) between island categories. Thus, bottom-up effects seemed mainly be mediated via changes in resource quantity not quality. Despite a large potential trophic mismatch we were unable to observe any relation between the invertebrate stoichiometry and their density response to nesting cormorant colonies. We conclude that in our system stoichiometry is not a useful predictor of arthropod responses to variation in resource nutrient content. Furthermore, we found no strong evidence that resource quality was a prime determinant of invertebrate densities. Other factors like resource quantity, habitat structure and species interactions might be more important or masked stoichiometric effects.     

Between- and within-population differences in Phragmites australis

Phragmites australis (common reed) is a dominant clonal species in the interface between land and water in many European wetlands. Along the land-water gradient, strong consistently different selective forces might operate to give rise to genetic substructuring. I have investigated the occurrence of genetic substructuring in European P. australis populations. The present paper examines whether seedlings, from seeds collected at both ends of the land-water gradient, showed differences in response to nutrient supply. Under controlled conditions, the relative growth rate (RGR) in the exponential growth phase, and growth characters of 10-week old seedlings were assessed. Among populations, no differences in response to nutrient supply were found. Although total dry weight was not related to the geographic origin of the populations, northern/western compared to southern/eastern European populations (1) formed more but shorter shoots, (2) formed thinner but longer rhizomes, and (3) invested more dry matter in leaves at the expense of stems. It was concluded that these trait differences are likely to originate from differences in the length of the growing season in the native habitat. Within populations, ’water-side’ seedlings had a higher RGR under sub-optimal while for ’land-side’ seedlings this was under optimal nutrient conditions. Ten-week-old ’water-side’ seedlings had a higher total dry weight than ’land-side’ ones, irrespective of nutrient loading. Differences in growth could not clearly be related to differences in single biomass allocation and morphological traits. A discriminant analysis on these traits, however, revealed that ’water-side’ seedlings showed higher plasticity in discriminant scores than ’land-side’ seedlings in response to nutrient supply. Discriminant scores also pointed towards a subtle trade-off between height versus expansion growth of seedlings, from the water to landward side. In the Romanian population, this could be related to morphological differences between ploidy levels. Overall, it was concluded that within populations, selection on growth form rather than on adaptations to the nutrient status of the habitat might have taken place. Received: 20 August 1998 / Accepted: 29 July 1999     

Mini-Review: Nutrient Cycling in Lakes and Streams: Insights from a Comparative Analysis

Understanding of general ecosystem principles may be improved by comparing disparate ecosystems. We compared nutrient cycling in lakes and streams to evaluate whether contrasts in hydrologic properties lead to different controls and different rates of internal nutrient cycling. Our primary focus was nutrient cycling that results in increased productivity, so we quantified nutrient cycling by defining the recycling ratio (ρ) as the number of times a nutrient molecule is sequestered by producers before export. An analytic model of nutrient cycling predicted that in lakes ρ is governed by the processes that promote the mineralization and retard the sedimentation of particulate-bound nutrients, whereas in streams, ρ is governed by processes that promote the uptake and retard the export of dissolved nutrients. These differences were the consequence of contrast between lakes and streams in the mass-specific export rates (mass exported · standing stock^-1· time^-1) of dissolved and particulate nutrients. Although ρ is calculated from readily measured ecosystem variables, we found very few published data sets that provided the necessary data for a given ecosystem. We calculated and compared ρ in two well-studied P-limited ecosystems, Peter Lake and West Fork Walker Branch (WFWB). When ecosystems were scaled so that water residence time was equal between these two ecosystems, ρ was three orders of magnitude greater in WFWB. However, when we scaled by P residence time, ρ was nearly equal between these two ecosystems. This suggests broad similarities in ρ across ecosystem types when ecosystem boundaries are defined so that turnover times of limiting nutrients are the same. Received 19 November 1998; accepted 6 October 1999.     

Low ratios of silica to dissolved nitrogen supplied to rivers arise from agriculture not reservoirs

Coastal marine systems are greatly altered by toxic marine algae, eutrophication and hypoxia. These problems have been linked to decreased ratios of dissolved silica to inorganic nitrogen (Si : DIN) delivered from land. Two mechanisms for this decline under consideration are enhanced nitrogen (N) fertiliser losses from agricultural lands or Si sequestration in reservoirs. Here we examine these mechanisms via nutrient concentrations in impoundments receiving water from 130 watersheds in a landscape representative of the agriculture that often dominates coastal nutrient inputs. Decreased Si : DIN was correlated with agriculture, not impoundment. Watersheds with > 60% agricultural land yielded highest DIN, whereas Si was uncorrelated with agricultural intensity. Furthermore, eutrophic lakes were dominated by Cyanobacteria that use little Si, so reservoirs did not diminish Si : DIN. Instead, Si : DIN increased slightly as reservoir residence time increased. These data suggest that impoundments in agricultural watersheds may enhance the water quality of coastal ecosystems, whereas fertiliser losses are detrimental.     

Monopolization of litter processing by a dominant land crab on a tropical oceanic island

Litter processing by macroinvertebrates typically involves suites of species that act together to determine rates of breakdown and decomposition. However, tropical oceanic islands and coastal fringes on continents are often dominated by one or a few species of omnivorous land crabs that consume leaf litter. We used an exclusion experiment, together with other leaf removal and litter decomposition studies, to assess the role of a single dominant species, the red crab (Gecarcoidea natalis), in litter dynamics in rain forest on Christmas Island, Indian Ocean. In the presence of red crabs, litter cover and biomass varied seasonally, from almost complete cover and high biomass at the end of the dry season to almost total absence of litter at the end of each wet season. When crabs were excluded from both the shaded understory and light gaps in rain forest, litter increased rapidly to almost complete cover, which was then maintained year round. Leaf tether experiments, and measures of litter input and standing crops, indicated that red crabs monopolize litter processing, removing between 39 and 87% of the annual leaf fall from the forest floor. Rates of litter turnover were over twice as high in the presence of land crabs: the decomposition constant, k, was 2.6 in the understory exclusion plots, but rose to 6.0 in the presence of crabs. Red crabs occur at biomass densities (114 g m⁻²) far greater than those reported elsewhere for entire litter faunas. They significantly reduced the abundance of other litter invertebrates, but we did not detect any change in the relative frequencies of the major invertebrate groups (mites, collembolans, pulmonate snails, ants, psocopterans, and spiders). Wherever omnivorous land crabs are abundant, their activities may be paramount in litter decomposition and in regulating the rate of nutrient cycling. In monopolizing litter processing, they may also be important physical “ecosystem engineers”, translocating organic matter and nutrients into the soil and reducing available habitat for other animals. Received: 19 August 1998 / Accepted: 11 January 1999     

Heterogeneity of phosphorus distribution in a patterned landscape,the Florida Everglades

The biologically mediated transfer of nutrients from one part of a landscape to another may create nutrient gradients or subsidize the productivity at specific locations. If limited, this focused redistribution of the nutrient may create non-random landscape patterns that are unrelated to underlying environmental gradients. The Florida Everglades, USA, is a large freshwater wetland that is patterned with tree islands, elevated areas that support woody vegetation. A survey of 12 tree islands found total soil phosphorus levels 3–114 times greater on the island head than the surrounding marsh, indicating that the Florida Everglades is not a homogeneous oligotrophic system. It was estimated that historically 67% of the phosphorus entering the central Everglades was sequestered on tree islands, which are ~3.8% of the total land area. This internal redistribution of phosphorus onto tree islands due to the establishment of trees may be one reason that marshes have remained oligotrophic and may explain the spatial differentiation of the patterned Everglades landscape.     

The importance of guano birds to the Inca Empire and the first conservation measures implemented by humans

The Inca Empire was the largest ancient civilization in South America, extending across almost 4000 km of distinct environments from the Pacific coast to the Andes, and throughout most of the arid desert between. The present study highlights the importance of guano birds (Guanay Cormorant Leucocarbo bougainvillii, Peruvian Pelican Pelecanus thagus and Peruvian Booby Sula variegata) for the expansion and prosperity of the Inca Empire. The use of guano as a fertilizer was fundamental to sustain the agricultural development of the empire and, it has been argued, was the basis for its rapid growth. The access to guano on coastal islands and its subsequent transportation to highlands provided food security for a population of more than 8 million. The importance of guano birds to the Inca Empire led to the development of management plans based on a penal code aiming to preserve these species and their natural habitats. These protective actions may represent the first conservation measures ever implemented by humans based on the importance of species protection for human activities and livelihoods.     

Pattern of nutrient availability and plant community assemblage in Everglades Tree Islands, Florida, USA

We address the relative importance of nutrient availability in relation to other physical and biological factors in determining plant community assemblages around Everglades Tree Islands (Everglades National Park, Florida, USA). We carried out a one-time survey of elevation, soil, water level and vegetation structure and composition at 138 plots located along transects in three tree islands in the Park’s major drainage basin. We used an RDA variance partitioning technique to assess the relative importance of nutrient availability (soil N and P) and other factors in explaining herb and tree assemblages of tree island tail and surrounded marshes. The upland areas of the tree islands accumulate P and show low N concentration, producing a strong island-wide gradient in soil N:P ratio. While soil N:P ratio plays a significant role in determining herb layer and tree layer community assemblage in tree island tails, nevertheless part of its variance is shared with hydrology. The total species variance explained by the predictors is very low. We define a strong gradient in nutrient availability (soil N:P ratio) closely related to hydrology. Hydrology and nutrient availability are both factors influencing community assemblages around tree islands, nevertheless both seem to be acting together and in a complex mechanism. Future research should be focused on segregating these two factors in order to determine whether nutrient leaching from tree islands is a factor determining community assemblages and local landscape pattern in the Everglades, and how this process might be affected by water management.     

Litter decomposition promotes differential feedbacks in an oligotrophic southern Everglades wetland

The differential accumulation or loss of carbon and nutrients during decomposition can promote differentiation of wetland ecosystems, and contribute to landscape-scale heterogeneity. Tree islands are important ecosystems because they increase ecological heterogeneity in the Everglades landscape and in many tropical landscapes. Only slight differences in elevation due to peat accumulation allow the differentiation of these systems from the adjacent marsh. Hydrologic restoration of the Everglades landscape is currently underway, and increased nutrient supply that could occur with reintroduction of freshwater flow may alter these differentiation processes. In this study, we established a landscape-scale, ecosystem-level experiment to examine litter decomposition responses to increased freshwater flow in nine tree islands and adjacent marsh sites in the southern Everglades. We utilized a standard litterbag technique to quantify changes in mass loss, decay rates, and phosphorus (P), nitrogen (N) and carbon (C) dynamics of a common litter type, cocoplum (Chrysobalanus icaco L.) leaf litter over 64 weeks. Average C. icaco leaf degradation rates in tree islands were among the lowest reported for wetland ecosystems (0.23 ± 0.03 yr⁻¹). We found lower mass loss and decay rates but higher absolute mass C, N, and P in tree islands as compared to marsh ecosystems after 64 weeks. With increased freshwater flow, we found generally greater mass loss and significantly higher P concentrations in decomposing leaf litter of tree island and marsh sites. Overall, litter accumulated N and P when decomposing in tree islands, and released P when decomposing in the marsh. However, under conditions of increased freshwater flow, tree islands accumulated more P while the marsh accumulated P rather than mineralizing P. In tree islands, water level explained significant variation in P concentration and N:P molar ratio in leaf tissue. Absolute P mass increased strongly with total P load in tree islands (r ² = 0.81). In the marsh, we found strong, positive relationships with flow rate. Simultaneous C and P accumulation in tree island and mineralization in adjacent marsh ecosystems via leaf litter decomposition promotes landscape differentiation in this oligotrophic Everglades wetland. However, results of this study suggest that variation in flow rates, water levels and TP loads can shift differential P accumulation and loss leading to unidirectional processes among heterogeneous wetland ecosystems. Under sustained high P loading that could occur with increased freshwater flow, tree islands may shift to litter mineralization, further degrading landscape heterogeneity in this system, and signaling an altered ecosystem state.     

INDeGenIUS,a new method for high-throughput identification of specialized functional islands in completely sequenced organisms

Genomic islands (GIs) are regions in the genome which are believed to have been acquired via horizontal gene transfer events and are thus likely to be compositionally distinct from the rest of the genome. Majority of the genes located in a GI encode a particular function. Depending on the genes they encode, GIs can be classified into various categories, such as ‘metabolic islands’, ‘symbiotic islands’, ‘resistance islands’, ‘pathogenicity islands’, etc. The computational process for GI detection is known and many algorithms for the same are available. We present a new method termed as Improved N-mer based Detection of Genomic Islands Using Sequence-clustering (INDeGenIUS) for the identification of GIs. This method was applied to 400 completely sequenced species belonging to proteobacteria. Based on the genes encoded in the identified GIs, the GIs were grouped into 6 categories: metabolic islands, symbiotic islands, resistance islands, secretion islands, pathogenicity islands and motility islands. Several new islands of interest which had previously been missed out by earlier algorithms were picked up as GIs by INDeGenIUS. The present algorithm has potential application in the identification of functionally relevant GIs in the large number of genomes that are being sequenced. Investigation of the predicted GIs in pathogens may lead to identification of potential drug/vaccine candidates.     

Comparisons of N- and P-limited productivity between high granitic islands versus low carbonate atolls in the Seychelles Archipelago: a test of the relative-dominance paradigm

This exploratory study suggests that different geological systems (carbonate vs. granitic) in tropical waters have contrasting patterns of photosynthetic nutrient limitation correlated with inorganic nitrogen (N) and phosphorus (P) availability. Physiological assays for 21 predominant macrophyte species show that inorganic N and P are much less limiting to photosynthesis on granitic islands than is the case on carbonate islands and that, of the two, P is more likely to limit production in carbonate-rich tropical waters. Patterns of nutrient limitation in turn are reflected by differences in the relative dominance of functional groups of sessile, epilithic, photosynthetic organisms. Surveys at 33 sites on 10 islands revealed that nearshore waters on high granitic islands tend to be characterized by large and species-rich standing stocks of frondose macroalgae, often dominated by Sargassum spp., whereas waters around low carbonate islands tend to be dominated by hermatypic corals. Macrophyte tissue and seawater analyses also indicate a possible trend toward higher levels of N and P in granitic vs. carbonate islands. Pagode Island, a low carbonate island influenced by guano from seabird colonies, is an exception, with few corals, relatively high levels of tissue and seawater N and P, and a predominance of macroalgae (mostly Dictyosphaeria cavernosa).     

Purification and characterization of a glucokinase from young tomato (Lycopersicon esculentum L. Mill.) fruit

In order to clearly establish the properties of the enzymes responsible for hexose phosphorylation we have undertaken the separation and characterization of these enzymes present in tomato fruit (Martinez-Barajas and Randall 1996). This report describes the partial purification and characterization of glucokinase (EC. 2.7.1.1) from young green tomato fruit. The procedure yielded a 360-fold enrichment of glucokinase. Tomato fruit glucokinase is a monomer with a molecular mass of 53 kDa. Glucokinase activity was optimal between pH 7.5 and 8.5, preferred ATP as the phosphate donor (K _m = 0.223 mM) and exhibited low activity with GTP or UTP. The tomato fruit glucokinase showed highest affinity for glucose (K _m = 65 μM). Activity observed with glucose was 4-fold greater than with mannose and 50-fold greater than with fructose. The tomato fruit glucokinase was sensitive to product inhibition by ADP (K _i = 36 μM). Little inhibition was observed with glucose 6-phosphate (up to 15 mM) at pH 8.0; however, at pH 7.0 glucokinase activity was inhibited 30–50% by physiological concentrations of glucose 6-phosphate. Received: 4 October 1997 / Accepted: 10 January 1998     

Climate change will alter amphibian‐mediated nutrient pathways: evidence from Rana temporaria tadpoles in experimental ponds

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Effects of nutrient recycling by zooplankton and fish on phytoplankton communities

In laboratory experiments we tested the hypothesis that nutrients supplied by fish and zooplankton affect the structure and dynamics of phytoplankton communities. As expected from their body size differences, fish released nutrients at lower mass-specific rates than Daphnia. On average, these consumers released nutrients at similar N:P ratios, although the ratios released by Daphnia were more variable than those released by fish. Nutrient supply by both fish and Daphnia reduced species richness and diversity of phytoplankton communities and increased algal biomass and dominance. However, nutrient recycling by fish supported a more diverse phytoplankton community than nutrient recycling by Daphnia. We conclude that nutrient recycling by zooplankton and fish have different effects on phytoplankton community structure due to differences in the quality of nutrients released. Received: 21 December 1998 / Accepted: 31 May 1999     

Fire in the Brazilian Amazon 2. Biomass, nutrient pools and losses in cattle pastures

Conversion to cattle pasture is the most common fate of the ≈426,000 km² of tropical forest that has been deforested in the Brazilian Amazon. Yet little is known about the biomass, C, nutrient pools, or their responses to the frequent fires occurring in these pastures. We sampled biomass, nutrient pools and their losses or transformation during fire in three Amazonian cattle pastures with typical, but different, land-use histories. Total aboveground biomass (TAGB) ranged from to 53 to 119 Mg ha⁻¹. Residual wood debris from the forests that formally occupied the sites composed the majority of TAGB (47–87%). Biomass of fine fuels, principally pasture grasses, was ≈16–29 Mg ha⁻¹. Grasses contained as much as 52% of the aboveground K pool and the grass and litter components combined composed as much as 88% of the aboveground P pool. Fires consumed 21–84% of the TAGB. Losses of C to the atmosphere ranged from 11 to 21 Mg ha⁻¹ and N losses ranged from 205 to 261 kg ha⁻¹. Losses of S, P, Ca, and K were <33 kg ha⁻¹. There were no changes in surface soil (0–10 cm) nutrient concentration in pastures compared to adjacent primary forests. Fires occur frequently in cattle pastures (i.e., about every 2 years) and pastures are now likely the most common type of land burned in Amazonia. The first 6 years of a pastures existence would likely include the primary forest slash fire and three pasture fires. Based upon our results, the cumulative losses of N from these fires would be 1935 kg ha⁻¹ (equivalent to 94% of the aboveground pool of primary forest). Postfire aboveground C pools in old pastures are as low as 3% of those in adjacent primary forest. The initial primary forest slash fire and the repeated fires occurring in the pastures result in the majority of aboveground C and nutrient pools being released via combustion processes rather than decomposition processes. Received: 6 January 1997 / Accepted: 2 September 1997     

Conserving the endangered Mexican fishing bat (<Emphasis Type="Italic">Myotis vivesi</Emphasis>): genetic variation indicates extensive gene flow among islands in the Gulf of California

The endangered Mexican fishing bat, Myotis vivesi, appears to have suffered widespread extinction and population decline on islands throughout the Gulf of California, largely due to predation by introduced cats and rats. To restore populations of fishing bats and other native species, conservation efforts have focused on eradicating introduced vertebrates from several Gulf islands. These efforts assume that individuals from existing populations will recolonize islands and that continued dispersal will help sustain vulnerable populations thereafter. However, the extent of inter-island dispersal in fishing bats is unknown. In this study we analyzed patterns of genetic variation to gauge the extent of gene flow and, thus, potential dispersal among islands. DNA was sampled from 257 fishing bats on 11 Gulf islands (separated by ca. 6–685 km of open water), and individuals were genotyped at six microsatellite loci and haplotyped at a 282 bp fragment of the mtDNA control region. With microsatellites, we found weak population genetic structure and a pattern of isolation by distance, while with mtDNA we found strong structure but no isolation by distance. Our results indicate that island subpopulations separated by large expanses of open water are nonetheless capable of maintaining high genetic diversity and high rates of gene flow. Unfortunately, little is known about the spatial patterns of dispersal or mating system of fishing bats, and these behavioral factors, in particular female philopatry, might reduce the probability of the species recolonizing Gulf islands.