Agricultural food subsidies, migratory connectivity and large-scale disturbance in arctic coastal systems: a case study.

An allochthonous input can modify trophic relationships, by providing an external resource that is normally limiting within a system. The subsidy may not only elicit a growth response of the primary producers via a bottom-up effect, but it also may lead to runaway herbivore growth in the absence of increased predation. If the consumer is migratory and predation is similarly dampened in the alternative system, the increased numbers may produce a top-down cascade of direct and indirect effects on an ecosystem that may be a great distance from the source of the subsidy. In an extreme case, it can lead to a catastrophic shift in ecosystem functioning as a result of biotic exploitation that produces an alternative stable state. The loss of resilience is particularly sensitive to herbivore density which can result in two different outcomes to the vegetation on which the consumer feeds. Over-compensatory growth of above-ground biomass gives way to sward destruction and near irreversible changes in soil properties as density of a herbivore increases. A striking temporal asymmetry exists between a reduction in the consumer population and recovery of damaged vegetation and degraded soils.

The embarrassment of riches: atmospheric deposition of nitrogen and community and ecosystem processes.

Anthropogenic sources of nitrogen have exceeded, and will continue to exceed, annual inputs of nitrogen produced by natural processes. Nitrogen enrichment may in plant tissue chemistry and microbial decomposition processes, as well as affecting rates of herbivory, all of which may be expected to result in changes in plant species assemblages Individual concepts, such as nitrogen saturation and critical load, used to describe the effects of enrichment on soil, community, ecosystm processes and species assemblages, cannot accomodate easily the range of interactions and different environmental processes. A number of approaches need to be used in tandem. Major gaps in knowledge are rates of transfer of anthropogenic nitrogen within and between different ecosystem and how these rates affect population dynamic of individual species and trophic relationships. Without this information, predictions of biological effects of enrichment are difficult to make.

High mortality, fluctuation in numbers, and heavy subterranean insect herbivory in bush lupine, Lupinus arboreus.

Sporadic patchy die-off of bush lupine, Lupinus arboreus, has long been known. We describe in detail a series of these incidents on the central California coast, based upon observational and comparative evidence. Stands of thousands of plants die, while nearby mature plants live on. In some sites, repeated die-off followed by regeneration from the seed bank has led to the cover and density of this woody, perennial plant fluctuating widely over the 40 year period for which records exist. Root damage by caterpillars of the ghost moth or "swift" Hepialus californicus (Lepidoptera, Hepialidae) is a major cause of individual bush death and a probable cause of die-off of stands of lupine. Hidden from view underground, a few of these insects readily kill a juvenile or young mature plant by girdling and reaming-out roots. The mass mortality of L. arboreus that we observed involved heavy root damage by these caterpillars in evenaged stands of plants in their first (1.5-year-old) or second (2.5-year-old) flowering season. The injured plants set seed before dying. Older, larger bush lupines better withstood root damage. In plants aged 3 or more years, damage and mortality were correlated with the intensity of ghost moth caterpillars in the roots. At the highest intensity (mean = 37.5, maximum = 62 caterpillars/root), a stand of large, old L. arboreus suffered 41% mortality; 45% of root cambium (median value) was destroyed by feeding caterpillars. Mass death of mature L. arboreus was not correlated with folivory, and leaf damage ranged from nil to moderate in instances of die-off. The western tussock moth, Orgyia vetusta, accounted for the highest levels of folivory, but this insect was rare when die-offs occurred. The lowest lupine mortality rates in our study occurred where tussock caterpillar intensities were high and where plants were repeatedly defoliated by this insect. However, experimental defoliation by high, but realistic, intensities of tussock moth caterpillars resulted in some mortality of mature bushes, and the combined effects of leaf and root herbivory have yet to be assessed. In its natural range on the California coast, bush lupine has several additional species of insect herbivores that can be locally abundant and injurious to the plant, although none is associated with die-off. Subterranean natural enemies of ghost moth caterpillars may play a role in the patchy waxing and waning of this shrub. Locally, a new species of entomophagous nematode (Heterorhabditis sp.) cause high mortality in the soil, before ghost moth caterpillars have entered the root. This natural enemy may thus afford lupines protection from heavy underground herbivory.

Environmental change and the cost of philopatry: an example in the lesser snow goose.

The consequences of philopatric and dispersal behaviours under changing environmental conditions were examined using data from the colony of Lesser Snow Geese (Anser caerulescens caerulescens) breeding at La Pérouse Bay, Manitoba, Canada. In response to increased population size and decreased food abundance over time, increasing numbers of family groups have been dispersing from the traditional feeding areas. Goslings from dispersed broods were significantly heavier (7.3%), and had longer culmens (3.1%), head lengths (2.6%) and marginally longer tarsi (1.9%) on average than goslings that remained within La Pérouse Bay itself. These differences were consistent in each of 5 years. There was no evidence that the larger size of dispersed goslings was due to either a tendency for larger adults to disperse to alternative sites, or increased mortality of smaller goslings among dispersed broods. The most likely cause for the larger size of goslings from dispersed broods was the significantly greater per capita availability of the preferred salt-marsh forage species at non-traditional brood-rearing areas. The larger goslings in non-traditional feeding areas showed significantly higher firstyear survival, suggesting that the use of deteriorating traditional feeding areas may currently be maladaptive in this population.

Growth responses of arctic graminoids following grazing by captive lesser snow geese.

The effects of grazing by captive goslings of the Lesser Snow Goose on coastal vegetation at La Pérouse By. Manitoba were investigated. Swards of Carex subspathacea, Festuca rubra and Calamagrostis deschampsioides were grazed once for different periods (0-180 min) and regrowth of vegetation determined, based on measurements of standing crop, net above-ground primary production (NAPP) and forage quality (leaf nitrogen content). The amounts of foliage removed from swards of Carex subspathacea increased with the length of the grazing period, but after 44 days of regrowth there were no significant differences in above-ground biomass between control and grazed plots. While the amount of foliage removed by goslings from swards of Festuca rubra increased with the length of the grazing period (except after 150 min of grazing), the increase in biomass following defoliation was similar among treatments. Goslings removed little biomass from swards of Calamagrostis deschampsioides, even when the opportunity for grazing was 180 min. No significant differences in standing-crop or NAPP between grazed and ungrazed plots were detected by the end of summer. Grazing had no significant effect on amounts of nitrogen in leaf tissue of all species, suggesting that faecal nitrogen was not rapidly incorporated into plant biomass within the growing season. Patterns of regrowth of these species are compared to that of Puccinellia phryganodes. An increase in goose numbers in recent years has led to birds foraging on less preferred species, such as Calamagrostis deschampsiodes and Festuca rubra. Their poor nutritional quality and a lack of a rapid growth response following defoliation may explain, in part, the decline in the weight of wild goslings recorded over the last decade.

Effect of increased salinity on CO2 assimilation, O 2 evolution and the δ (13)C values of leaves of Plantago maritima L. developed at low and high NaCl levels.

The effect of increased salinity on photosynthesis was studied in leaves of Plantago maritima L. that developed while plants were at low and high NaCl levels. In leaves that developed while plants were grown at 50 mol·m(-3), exposure to 200 and 350 mol·m(-3) NaCl resulted in reductions in net CO2 assimilation and stomatal conductance. The decline in CO2 assimilation in plants at 200 and 350 mol·m(-3) NaCl occurred almost exclusively at high intercellular CO2 concentrations. The initial slope of the CO2 assimilation-intercellular CO2 (A-C i) curve, determined after salinity was increased, was identical or very similar to that measured initially. In contrast to the reductions observed in CO2 assimilation, there were no significant differences in O2 evolution rates measured at 5% CO2 among leaves from plants exposed to higher salinity and plants remaining at low salinity.Leaves that developed while plants were at increased salinity levels also had significantly lower net CO2 assimilation rates than plants remaining at 50 mol·m(-3) NaCl. The lower CO2 assimilation rates in plants grown at 200 and 350 mol·m(-3) NaCl were a result of reduced stomatal conductance and low intercellular CO2 concentration. There were no significant differences among treatments for O2 evolution rates measured at high CO2 levels. The increased stomatal limitation of photosynthesis was confirmed by measurements of the (13)C/(12)C composition of leaf tissue. Water-use efficiency was increased in the plants grown at high salinity.

The role of lesser snow geese as nitrogen processors in a sub-arctic salt marsh.

Ammonia volatilization losses from faeces of Lesser Snow Geese were measured during the summer of 1987 on the salt-marsh flats at La Pérouse Bay. Amounts of ammonia volatilized increased with increasing ambient temperature, and ranged from 1.0 to 15.1 mg N per 100 mg of nitrogen present as soluble ammonium ions at the start of the 8-h experiment. Using estimates of faecal deposition reported previously, the annual loss via volatilization was estimated at 0.08 g N m-2, or 7.9% of the nitrogen present in goose faeces. Percent change in soluble ammonium ions in fresh faeces after 8 h ranged from -51.1% to +41.1%, indicating that net mineralization of organic nitrogen occurred in some of the faeces. Microbial respiration of fresh goose faeces increased exponentially with temperature. However, variable rates of net mineralization per unit rate of respiration indicated that the substrate quality affected microbial immobilization and thus net nitrogen mineralization. In feeding experiments, captive goslings grazed different types of vegetation, each with distinctive nutritional qualities. Forage quality had significant effects on goose feeding behavior and subsequent rates of nitrogen mineralization in fresh faeces. Net nitrogen mineralization rates in faeces from geese which grazed the three vegetation types ranged from 1.31 to 4.97 mg NH 4+ -N gDW-1 24 h-1. Because plant growth in this salt marsh is nitrogen-limited, where swards are grazed, mineralization of organic faecal nitrogen represents an essential link in the maintenance of the flow of nitrogen into the sediments and the sustained growth of vegetation at a time when most required by the geese.

Responses of halophytes to high salinities and low water potentials.

The effects of nonsaline polyethylene glycol (PEG)-6000 and saline seawater solutions of comparable osmotic potential on the concentrations of organic solutes and inorganic ions in the tissues of halophytes (Plantago maritima L., Triglochin maritima L., Limonium vulgare Mill., Halimione portulacoides (L.) Aell) have been investigated. Studies were made to determine whether high salinities induce specific ion effects that are absent in plants grown in nonsaline solutions of comparable osmotic potential. Over-all, the responses of each species to the two different treatments (seawater or PEG) are similar; the accumulation of organic solutes (compatible osmotica) in tissues is primarily correlated with a decrease in the osmotic potential of culture solutions. Depending on the species, sorbitol, proline, reducing sugars, quaternary ammonium compounds, and alpha-amino nitrogen accumulate in tissues as the water potential of the tissues falls. Within a species there are differences in the concentrations of inorganic ions and organic solutes between roots and shoots of plants grown at high salinities or at high concentrations of PEG.

Carbon dioxide exchange in leaves of Spartina anglica Hubbard.

The gas-exchange method has been used to measure net carbon dioxide assimilation at different temperatures and irradiances in leaves of salt-treated and untreated plants of Spartina anglica. The results together with those of the CO2 compensation point and leaf anatomical data clearly indicate that Spartina anglica is a high photosynthetic capacity plant. At high temperatures and irradiances leaves of the salt-treated plants were able to maintain net photosynthesis, in contrast to the behaviour of leaves of untreated plants.The ecological significance of these results is discussed.