Scale-dependent response diversity of seabirds to prey in the North Sea

Functional response diversity is defined as the diversity of responses to environmental change among species that contribute to the same ecosystem function. Because different ecological processes dominate on different spatial and temporal scales, response diversity is likely to be scale dependent. Using three extensive data sets on seabirds, pelagic fish, and zooplankton, we investigate the strength and diversity in the response of seabirds to prey in the North Sea over three scales of ecological organization. Two-stage analyses were used to partition the variance in the abundance of predators and prey among the different scales of investigation: variation from year to year, variation among habitats, and variation on the local patch scale. On the year-to-year scale, we found a strong and synchronous response of seabirds to the abundance of prey, resulting in low response diversity. Conversely, as different seabird species were found in habitats dominated by different prey species, we found a high diversity in the response of seabirds to prey on the habitat scale. Finally, on the local patch scale, seabirds were organized in multispecies patches. These patches were weakly associated with patches of prey, resulting in a weak response strength and a low response diversity. We suggest that ecological similarities among seabird species resulted in low response diversity on the year-to-year scale. On the habitat scale, we suggest that high response diversity was due to interspecific competition and niche segregation among seabird species. On the local patch scale, we suggest that facilitation with respect to the detection and accessibility of prey patches resulted in overlapping distribution of seabirds but weak associations with prey. The observed scale dependencies in response strength and diversity have implications for how the seabird community will respond to different environmental disturbances.     

Seasonal changes in the diet of a critically endangered seabird and the importance of trawling discards

Pelagic seabirds obtain food from oceans where the availability of their prey changes rapidly both seasonally and spatially. Here, we investigated changes in the trophic habits of the critically endangered Balearic shearwater (Puffinus mauretanicus) through the breeding season and tested for dietary differences between sexes and age classes. We analysed δ¹⁵N and δ¹³C values in blood of adults during the pre-incubation, incubation and chick-rearing periods and of their chicks. Using a two-isotope mixing model, we estimated dietary contributions based on isotope values from potential prey species which included small pelagic species available naturally and demersal fish species available only from trawling discards. Balearic shearwaters showed clear isotopic and dietary variation through the breeding season. During pre-incubation, breeding adults appeared to exploit demersal fish, whereas during the incubation and chick-rearing period, they fed mainly on pelagic anchovies (Engraulis encrasicolus) and pilchards (Sardina pilchardus). Similarly, chicks were fed mainly with anchovies, a resource with a high energetic value. This variation in the dietary habits of adult shearwaters during the breeding season was probably related to both natural and fishery-induced seasonal changes in the availability of potential prey species within their main feeding grounds. However, changes in the nutritional requirements of the shearwaters could also play an important role. Indeed, diet differed between sexes during pre-incubation: females fed less on trawling discards and more on small pelagic fish than males. This sexual segregation in diet could be the consequence of higher nutritional requirements of females during this period. Our study reveals the differential importance of both trawling discards and small pelagic fish species for a pelagic seabird depending on the breeding period and illustrates the importance of considering the entire breeding season when making inferences about the importance of specific prey in seabird dietary studies.     

Underwater and above-water search patterns of an Arctic seabird: reduced searching at small spatiotemporal scales

How predators vary search patterns in response to prey predictability is poorly known. For example, marine invertebrates may be predictable but of low energy value, while fish may be of higher energy value but unpredictable at large (pelagic schools) or small (solitary benthics) spatial scales. We investigated the search patterns of the thick-billed murre (Uria lomvia), an Arctic seabird feeding on invertebrates, pelagic fish, or benthic fish. Foraging ranges at the Coats Island colony are generally smaller (<240 min per trip) than at larger colonies, and many birds specialize in foraging tactics and diet. Underwater search times for benthic fish were higher than for pelagic fish or invertebrates while above-water search times for pelagic fish were higher than for benthic fish or invertebrates. There were few stops during trips. Total trip time, flying time, number of flights, and number of dives were intercorrelated and increased with prey energy content, suggesting that longer trips involved fewer prey encounters due to selection of higher-quality, but rarer, prey items. Flight times were not Lévy-distributed and seabirds may have used area-restricted searches. The high degree of specialization, apparent absence of information center effects, and reduced above-water searching times may be linked to the relatively small colony size and the resulting short commuting distances to feeding areas, leading to greater prey predictability. We concluded that prey predictability over various scales affected predator search patterns.     

Introduced trout sever trophic connections in watersheds: consequences for a declining amphibian

Trophic linkages between terrestrial and aquatic ecosystems are increasingly recognized as important yet poorly known features of food webs. Here we describe research to understand the dynamics of lake food webs in relation to a native riparian amphibian and its interaction with introduced trout. The mountain yellow-legged frog Rana muscosa is endemic to alpine watersheds of the Sierra Nevada Mountains and the Transverse Ranges of California, but it has declined to a small fraction of its historical distribution and abundance. Although remaining frogs and introduced trout feed in different habitats of alpine lakes, our stable-isotope analyses clearly show that the same resource base of benthic invertebrates sustains their growth. During one period, insect emergence from naturally fishless lakes was nearly 20-fold higher compared to adjacent lakes with trout, showing that fish reduce availability of aquatic prey to amphibious and terrestrial consumers. Although trout cannot prey on adult frogs due to gape limitation, foraging post-metamorphic frogs are 10 times more abundant in the absence of trout, suggesting an important role for competition for prey by trout in highly unproductive alpine watersheds. Most Sierran lakes contain fish, and those that do not are usually small isolated ponds; in our study, these two lake types supported the lowest densities of post-metamorphic frogs, and these frogs were less reliant on local, benthic sources of productivity. Since Rana muscosa was formerly the most abundant vertebrate in the Sierra Nevada, the reduction in energy flow from lake benthos to this consumer due to fish introductions may have had negative consequences for its numerous terrestrial predators, many of which have also declined. We suggest that disruptions of trophic connections between aquatic and terrestrial food webs are an important but poorly understood consequence of fish introduction to many thousands of montane lakes and streams worldwide and may contribute to declines of native consumers in riparian habitats.     

Behavioural inertia places a top marine predator at risk from environmental change in the Benguela upwelling system

In variable environments, organisms are bound to track environmental changes if they are to survive. Most marine mammals and seabirds are colonial, central-place foragers with long-term breeding-site fidelity. When confronted with environmental change, such species are potentially constrained in their ability to respond to these changes. For example, if environmental conditions deteriorate within their limited foraging range, long-lived species favour adult survival and abandon their current breeding effort, which ultimately influences population dynamics. Should poor conditions persist over several seasons, breeding-site fidelity may force animals to continue breeding in low-quality habitats instead of emigrating towards more profitable grounds. We assessed the behavioural response of a site-faithful central-place forager, the Cape gannet Morus capensis, endemic to the Benguela upwelling system, to a rapid shift in the distribution and abundance of its preferred prey, small pelagic shoaling fish. We studied the distribution and the abundance of prey species, and the diet, foraging distribution, foraging effort, energy requirements, and breeding success of gannets at Malgas Island (South Africa) over four consecutive breeding seasons. Facing a rapid depletion of preferred food within their foraging range, Cape gannets initially increased their foraging effort in search of their natural prey. However, as pelagic fish became progressively scarcer, breeding birds resorted to scavenging readily available discards from a nearby demersal fishery. Their chicks cannot survive on such a diet, and during our 4-year study, numbers of breeding birds at the colony decreased by 40% and breeding success of the remaining birds was very low. Such behavioural inflexibility caused numbers of Cape gannets breeding in Namibia to crash by 95% following over-fishing of pelagic fish in the 1970s. In the context of rapid environmental changes, breeding-site fidelity of long-lived species may increase the risk of local or even global extinction, rendering these species particularly vulnerable to global change.     

Evaluating impacts of forage fish abundance on marine predators

Forage fish—small, low trophic level, pelagic fish such as herrings, sardines, and anchovies—are important prey species in marine ecosystems and also support large commercial fisheries. In many parts of the world, forage fish fisheries are managed using precautionary principles that target catch limits below the maximum sustainable yield. However, there are increasing calls to further limit forage fish catch to safeguard their fish, seabird, and marine mammal predators. The effectiveness of these extra-precautionary regulations, which assume that increasing prey abundance increases predator productivity, are under debate. In this study, we used prey-linked population models to measure the influence of forage fish abundance on the population growth rates of 45 marine predator populations representing 32 fish, seabird, and mammal species from 5 regions around the world. We used simulated data to confirm the ability of the statistical model to accurately detect prey influences under varying levels of influence strength and process variability. Our results indicate that predator productivity was rarely influenced by the abundance of their forage fish prey. Only 6 predator populations (13% of the total) were positively influenced by increasing prey abundance and the model exhibited high power to detect prey influences when they existed. These results suggest that additional limitation of forage fish harvest to levels well below sustainable yields would rarely result in detectable increases in marine predator populations.     

Age and sexual differences in the exploitation of two anthropogenic food resources for an opportunistic seabird

The availability of food resources has been suggested as a major factor in the substantial increase in reproductive output, survival, recruitment and, ultimately, population growth rates in most organisms. In fact, the artificial increase in food availability resulting from human activities has been suggested as a factor in the substantial increase in population size of several seabirds in recent decades. In the present study, our primary aim was to estimate the importance of the main natural prey and two alternative feeding resources, fishery discards and the invasive American crayfish Procambarus clarkii, for an opportunistic seabird, the Audouin’s gull Larus audouinii. We also assessed the influence of age and sex in the use of those three types of food. For this purpose, we compared the analyses of δ¹⁵N and δ¹³C in blood of male and female adults of known age and chicks with those in their potential prey. Our results reveal sex-related and age-related differences in the consumption of fish discards, small pelagic fish and American crayfish. Differences in the diet of males and females and also between adults and chicks could be related to different nutritional requirements. Age differences were probably related to their different foraging proficiency and the tendency of young breeders to opportunistically exploit anthropogenically derived food. This study illustrates the importance of considering the age and sex of individuals to obtain feasible dietary information and to understand how the exploitation of food of human origin could affect population growth.     

Importance of jumbo squid Dosidicus gigas (Orbigny, 1835) in the pelagic ecosystem of the central Gulf of California

The Humboldt squid is an important predator in the pelagic ecosystem of the central Gulf of California and the commercial catch of this species has increased over the past decade, probable due to a decrease of several top predators (sharks, large pelagic fish and the marine mammals) and the optimal feeding conditions in this area. Its high abundance and important position in the pelagic food web was quantified through two trophic models of the pelagic ecosystem of the central Gulf of California. Models represented conditions in 1980 and 2002, to document the decadal changes in ecosystem structure and function. The models were composed of 18 functional groups, including marine mammals, birds, fish, mollusks, crustaceans, and primary producers. Model results show direct negative effects on principal prey groups such as myctophids and pelagic red crab and positive effects on sharks, marine mammals and specifically sperm whales. It thus appears that the jumbo squid has an important role in the ecosystem and plays a central part in the overall energy flow as main food item for most top predators, and due to its predation of organisms on lower tropic levels.     

Measuring terrestrial subsidies to aquatic food webs using stable isotopes of hydrogen

Understanding river food webs requires distinguishing energy derived from primary production in the river itself (autochthonous) from that produced externally (allochthonous), yet there are no universally applicable and reliable techniques for doing so. We compared the natural abundance stable isotope ratios of hydrogen (deltaD) of allochthonous and autochthonous energy sources in four different aquatic ecosystems. We found that autochthonous organic matter is uniformly far more depleted in deuterium (lower deltaD values) than allochthonous: an average difference of approximately 100% per hundred. We also found that organisms at higher trophic levels, including both aquatic invertebrates and fish, have deltaD values intermediate between aquatic algae and terrestrial plants. The consistent differences between leaves and algae in deltaD among these four watersheds, along with the intermediate values in higher trophic levels, indicate that natural abundance hydrogen isotope signatures are a powerful tool for partitioning energy flow in aquatic ecosystems.     

A major increase in snake pipefish (<Emphasis Type="Italic">Entelurus aequoreus</Emphasis>) in northern European seas since 2003: potential implications for seabird breeding success

Since the early 2000s routine fish surveys have recorded increasing numbers of snake pipefish, Entelurus aequoreus, in the northeast Atlantic. Fishermen and divers have also commented on this increase and pipefish have started to appear in the diet of seabirds and other marine predators. This paper collates information from these diverse sources and assesses the current status of snake pipefish. We found compelling evidence of a dramatic increase in the abundance of snake pipefish starting around 2003 and continuing up to the present (2006) and a range expansion northwards to Spitzbergen and the Barents Sea. Since 2004 snake pipefish have been increasingly recorded in the diet of many species of seabird breeding in colonies around the coast of the UK, and in Norway, Iceland and the Faeroe Islands. Information on the nutrient value of snake pipefish is currently lacking but their rigid, bony structure makes them difficult for young seabirds to swallow and there are numerous records of chicks choking to death. Thus, in the case of avian predators during the breeding season, it appears unlikely that increased abundance of snake pipefish will provide a useful alternative prey. The reason for the rapid and dramatic increase in numbers of snake pipefish is currently unclear but such events are characteristic of marine ecosystems and will almost certainly have an effect on food web dynamics.     

Seabird use of discards from a nearshore shrimp fishery in the South Atlantic Bight,USA

Shrimp trawling is common throughout the southeastern and Gulf of Mexico coasts of the USA and is the primary contributor to fisheries discards in these regions. Tens of thousands of nearshore seabirds nest near shrimp trawling grounds in the USA, but to date, there has been no assessment of the relationship between seabirds and shrimp trawlers. We examined the taxonomic composition of bycatch, rate at which seabirds scavenged bycatch, and energy density of discarded bycatch in a nearshore commercial shrimp fishery. Bycatch was primarily comprised of demersal fish that are not typically accessible to the plunge-diving and surface-feeding seabirds that occur in the area. Hence, seabird diets in the region appear to be broadened taxonomically by the availability of discards. Results from discard experiments indicated that 70% of the nearly 5,500 items discarded by hand were scavenged by seabirds and that the fate of a discarded item was most strongly predicted by its taxonomic order. Laughing gulls scavenged the greatest proportion of discards, although brown pelicans were the only species to scavenge more discards than predicted based upon their abundance. Because this is the first such study in the region, it is difficult to ascertain the extent or intensity of the impact that discards have on nearshore seabirds. Nonetheless, our results suggest that it will be difficult for managers to clearly understand fluctuations in local seabird population dynamics without first understanding the extent to which these species rely upon discards. This may be especially problematic in situations where seabird populations are recovering following natural or anthropogenic stressors.     

Trends in seabird breeding populations across the Great Barrier Reef

The Great Barrier Reef is an iconic ecosystem, known globally for its rich marine biodiversity that includes many thousands of tropical breeding seabirds. Despite indications of localized declines in some seabird species from as early as the mid-1990s, trends in seabird populations across the reef have never been quantified. With a long history of human impact and ongoing environmental change, seabirds are likely sentinels in this important ecosystem. Using 4 decades of monitoring data, we estimated site-specific trends for 9 seabird species from 32 islands and cays across the reef. Trends varied markedly among species and sites, but probable declines occurred at 45% of the 86 species-by-site combinations analyzed compared with increases at 14%. For 5 species, we combined site-specific trends into a multisite trend in scaled abundance, which revealed probable declines of Common Noddy (Anous stolidus), Sooty Tern (Onychoprion fuscatus), and Masked Booby (Sula dactylatra), but no long-term changes in the 2 most widely distributed species, Greater Crested Tern (Thalasseus bergii) and Brown Booby (Sula leucogaster). For Brown Booby, long-term stability largely resulted from increases at a single large colony on East Fairfax Island that offset declines at most other sites. Although growth of the Brown Booby population on East Fairfax points to the likely success of habitat restoration on the island, it also highlights a general vulnerability wherein large numbers of some species are concentrated at a small number of key sites. Identifying drivers of variation in population change across species and sites while ensuring long-term protection of key sites will be essential to securing the future of seabirds on the reef.     

Parent–offspring dietary segregation of Cory’s shearwaters breeding in contrasting environments

In pelagic seabirds, who often explore distant food resources, information is usually scarce on the level of trophic segregation between parents and their offspring. To investigate this issue, we used GPS tracking, stable isotopes and dietary information of Cory’s shearwaters Calonectris diomedea breeding in contrasting environments. Foraging trips at Selvagem Grande (an oceanic island) mainly targeted the distant African coast, while at Berlenga island (located on the continental shelf), shearwaters foraged mainly over nearby shelf waters. The degree of isotopic segregation between adults and chicks, based on δ¹³C, differed markedly between the two sites, indicating that adult birds at Selvagem fed their chicks with a mixture of shelf and offshore pelagic prey but assimilated more prey captured on coastal shelf waters. Isotopic differences between age classes at Berlenga were much smaller and may have resulted from limited dietary segregation or from age-related metabolic differences. The diet of shearwaters was also very different between the two colonies, with offshore pelagic prey only being detected at Selvagem Grande. Our findings suggest that spatial foraging constraints influence resource partitioning between pelagic seabirds and their offspring and can lead to a parent–offspring dietary segregation.     

Population dynamics of Hawaiian seabird colonies vulnerable to sea-level rise

Globally, seabirds are vulnerable to anthropogenic threats both at sea and on land. Seabirds typically nest colonially and show strong fidelity to natal colonies, and such colonies on low-lying islands may be threatened by sea-level rise. We used French Frigate Shoals, the largest atoll in the Hawaiian Archipelago, as a case study to explore the population dynamics of seabird colonies and the potential effects sea-level rise may have on these rookeries. We compiled historic observations, a 30-year time series of seabird population abundance, lidar-derived elevations, and aerial imagery of all the islands of French Frigate Shoals. To estimate the population dynamics of 8 species of breeding seabirds on Tern Island from 1980 to 2009, we used a Gompertz model with a Bayesian approach to infer population growth rates, density dependence, process variation, and observation error. All species increased in abundance, in a pattern that provided evidence of density dependence. Great Frigatebirds (Fregata minor), Masked Boobies (Sula dactylatra), Red-tailed Tropicbirds (Phaethon rubricauda), Spectacled Terns (Onychoprion lunatus), and White Terns (Gygis alba) are likely at carrying capacity. Density dependence may exacerbate the effects of sea-level rise on seabirds because populations near carrying capacity on an island will be more negatively affected than populations with room for growth. We projected 12% of French Frigate Shoals will be inundated if sea level rises 1 m and 28% if sea level rises 2 m. Spectacled Terns and shrub-nesting species are especially vulnerable to sea-level rise, but seawalls and habitat restoration may mitigate the effects of sea-level rise. Losses of seabird nesting habitat may be substantial in the Hawaiian Islands by 2100 if sea levels rise 2 m. Restoration of higher-elevation seabird colonies represent a more enduring conservation solution for Pacific seabirds.     

Tuna food habits related to the micronekton distribution in French Polynesia

Stomach content analyses are commonly used to study both fish feeding behaviour and trophic conditions. However, the interpretation of such data depends on fish foraging behaviour for a given environment and how representative the stomach contents are to the prey distribution. Tuna feeding behaviour was studied within the context of a research programme conducted in French Polynesia. Tuna prey distribution was characterised using acoustic measurements and pelagic trawls; thereafter, this distribution was compared with the stomach contents of tuna caught using an instrumented longline. Acoustic, pelagic trawling and stomach content analyses give complementary elements to describe the pelagic trophic habitat and to better understand tuna-prey relationships. The classic concept of a reduced food availability for tunas in the tropical pelagic environment seems relative. Tunas able to dive enough during daytime to exploit the migrant micronektonic species secure a source of regular food. This is particularly true of bigeye tuna (Thunnus obesus), which have ecophysiological capacities for this purpose. The behaviour of albacore tuna (T. alalunga), which dive >400 m in depth, remains less clear, as little is known about their vertical behaviour. Lastly, yellowfin tuna (T. albacares), which are distributed in more superficial waters, can better exploit the biomass of juvenile fish and crustaceans exported from the reefs. Analysis of the stomach fullness of tuna caught by longline, a passive gear, generally showed an empty state. This result suggests that most tuna foraging on large prey aggregations present in the study area are quickly satiated and escape longline capture and sampling. A consequence is that studies of tuna feeding behaviour based on longlining may be biased, particularly when large aggregations of prey are present such as in convergence zones. Another potential consequence is that longline tuna catch rates could differ according to prey richness. Longline tuna catch rates may sometimes reflect the relative abundance of prey rather than relative tuna abundance. Electronic supplementary material to this paper can be obtained by using the Springer LINK server located at http://dx.doi.org/10.1007/s00227-001-0776-3.     

A simulation model to explore the response of the Gulf of Maine food web to large-scale environmental and ecological changes

A dynamic simulation model was constructed using outputs from a balanced Gulf of Maine (GOM) energy budget model as the initial parameter set. The model was structured to provide a recipient control set of dynamics, largely based off of flows to and from different biological groups. The model was used to produce Monte Carlo simulations that were compared (percent change in biomass) with basecase simulations for a variety of scenarios. Changes in primary production, large increases in pelagic and demersal fish biomass, increases in fishing mortality, and large increases in top predators such as baleen whales and pinnepids were simulated. These scenarios roughly simulated the potential impacts of climate change, altered fishing pressure, additional protected species mitigations, and combinations thereof. Results suggest that the GOM system is primarily influenced by bottom-up processes involving phytoplankton, zooplankton, and bacterial biomass. Pelagic and demersal fish were important in determining trends in some of the scenarios. Marine mammals, large pelagic fish, and seabirds have a minor role in the GOM system in terms of biomass flows among the ecosystem components. The system is resilient to large-scale change due, in part to many predator–prey linkages. However, major alterations could occur from sustained climate change, high fishing rates, and by combinations of these types of external forcing mechanisms.     

Aquatic and terrestrial organic matter in the diet of stream consumers: implications for mercury bioaccumulation

The relative contribution of aquatic vs. terrestrial organic matter to the diet of consumers in fluvial environments and its effects on bioaccumulation of contaminants such as mercury (Hg) remain poorly understood. We used stable isotopes of carbon and nitrogen in a gradient approach (consumer isotope ratio vs. periphyton isotope ratio) across temperate streams that range in their pH to assess consumer reliance on aquatic (periphyton) vs. terrestrial (riparian vegetation) organic matter, and whether Hg concentrations in fish and their prey were related to these energy sources. Taxa varied in their use of the two sources, with grazing mayflies (Heptageniidae), predatory stoneflies (Perlidae), one species of water strider (Metrobates hesperius), and the fish blacknose dace (Rhinichthys atratulus) showing strong connections to aquatic sources, while Aquarius remigis water striders and brook trout (Salvelinus fontinalis) showed a weak link to in-stream production. The aquatic food source for consumers, periphyton, had higher Hg concentrations in low-pH waters, and pH was a much better predictor of Hg in predatory invertebrates that relied mainly on this food source vs. those that used terrestrial C. These findings suggest that stream biota relying mainly on dietary inputs from the riparian zone will be partially insulated from the effects of water chemistry on Hg availability. This has implications for the development of a whole-system understanding of nutrient and material cycling in streams, the choice of taxa in contaminant monitoring studies, and in understanding the fate of Hg in stream food webs.     

Nutritional geometry and macronutrient variation in the diets of gannets: the challenges in marine field studies

Foraging theory proposes that the nutritional driver of food choice and foraging in carnivores is energy gain. In contrast, recent laboratory experiments have shown that several species of carnivore select prey that provides a diet with a specific balance of macronutrients, rather than the highest energy content. It remains, however, to be determined how nutritionally variable the foods of predators in the wild are, and whether they feed selectively from available prey to balance their diet. Here, we used a geometric method named the right-angled mixture triangle (RMT) for examining nutritional variability in the prey and selected diets of a group of wild carnivores and marine top predators, the gannets (Morus spp.). A prey-level diet analysis was performed on Australasian gannets (M. serrator) from two New Zealand locations, and the macronutrient composition of their chosen prey species was measured. We use RMT to extend the comparison in the compositions of foods and diets from Australasian gannets from Australia as well as Northern gannets (M. bassanus) and Cape gannets (M. capensis). We found nutritional variability at multiple scales: intra- and interspecific variability in the pelagic fish and squid prey themselves; and intra- and interspecific variability in the diets consumed by geographically disparate populations of gannets. This nutritional variability potentially presents these predatory seabirds with both opportunity to select an optimal diet, and constraint if prevented from securing an optimal diet.     

Utilization of nitrogen derived from seabird guano by terrestrial and marine plants at St. Paul, Pribilof Islands, Bering Sea, Alaska

Dense, conspicuous colonies of seabirds and pinnipeds breed on ocean islands throughout the world. Such colonies have been shown to have local impacts on prey populations, but whether or not they affect nutrient cycling has been debated. We determined the natural abundance levels of the stable isotopes (C and N) of primary producers, seabirds and other consumers at and near St. Paul Island, Pribilof Islands, Bering Sea, in summer 1993. Marine primary producers (phytoplankton, as particulate organic matter, and kelp) collected near seabird colonies were ca. 6.5‰ enriched in both ¹⁵N and ¹³C relative to those collected further from shore. Terrestrial plants collected near the seabird colonies were enriched in ¹⁵N (δ¹⁵N ca. 22‰) compared with conspecifics collected away from the colonies (δ¹⁵N ca. 11‰). The trend towards higher δ¹⁵N values in marine and terrestrial plants near bird colonies is consistent with their uptake of ornithogenic N. This ¹⁵N-enrichment of plants using ornithogenic N can be attributed to a combination of two processes: trophic enrichment, and volatilization of ammonia produced during degradation of terrestrially deposited guano. Seabird breeding colonies at St. Paul Island appear to be significant sources of recycled nitrogen for terrestrial plants in the vicinity of colonies and for phytoplankton in the nearshore zone. Received: 4 July 1997 / Accepted: 20 November 1997     

Role of the fish Astyanax aeneus (Characidae) as a keystone nutrient recycler in low-nutrient neotropical streams

Nutrient recycling by animals is a potentially important biogeochemical process in both terrestrial and aquatic ecosystems. Stoichiometric traits of individual species may result in some taxa playing disproportionately important roles in the recycling of nutrients relative to their biomass, acting as keystone nutrient recyclers. We examined factors controlling the relative contribution of 12 Neotropical fish species to nutrient recycling in four streams spanning a range of phosphorus (P) levels. In high-P conditions (135 microg/L soluble reactive phosphorus, SRP), most species fed on P-enriched diets and P excretion rates were high across species. In low-P conditions (3 microg/L SRP), aquatic food resources were depleted in P, and species with higher body P content showed low rates of P recycling. However, fishes that were subsidized by terrestrial inputs were decoupled from aquatic P availability and therefore excreted P at disproportionately high rates. One of these species, Astyanax aeneus (Characidae), represented 12% of the total population and 18% of the total biomass of the fish assemblage in our focal low-P study stream but had P excretion rates > 10-fold higher than other abundant fishes. As a result, we estimated that P excretion by A. aeneus accounted for 90% of the P recycled by this fish assemblage and also supplied approximately 90% of the stream P demand in this P-limited ecosystem. Nitrogen excretion rates showed little variation among species, and the contribution of a given species to ecosystem N recycling was largely dependent upon the total biomass of that species. Because of the high variability in P excretion rates among fish species, ecosystem-level P recycling could be particularly sensitive to changes in fish community structure in P-limited systems.