Isotopic Differences between Forage Consumed by a Large Herbivore in Open,Closed, and Coastal Habitats: New Evidence from a Boreal Study System

Documenting habitat-related patterns in foraging behaviour at the individual level and over large temporal scales remains challenging for large herbivores. Stable isotope analysis could represent a valuable tool to quantify habitat-related foraging behaviour at the scale of individuals and over large temporal scales in forest dwelling large herbivores living in coastal environments, because the carbon (δ¹³C) or nitrogen (δ¹⁵N) isotopic signatures of forage can differ between open and closed habitats or between terrestrial and littoral forage, respectively. Here, we examined if we could detect isotopic differences between the different assemblages of forage taxa consumed by white-tailed deer that can be found in open, closed, supralittoral, and littoral habitats. We showed that δ¹³C of assemblages of forage taxa were 3.0‰ lower in closed than in open habitats, while δ¹⁵N were 2.0‰ and 7.4‰ higher in supralittoral and littoral habitats, respectively, than in terrestrial habitats. Stable isotope analysis may represent an additional technique for ecologists interested in quantifiying the consumption of terrestrial vs. marine autotrophs. Yet, given the relative isotopic proximity and the overlap between forage from open, closed, and supralittoral habitats, the next step would be to determine the potential to estimate their contribution to herbivore diet.     

Location Is Everything: Evaluating the Effects of Terrestrial and Marine Resource Subsidies on an Estuarine Bivalve

Estuaries are amongst the world’s most productive ecosystems, lying at the intersection between terrestrial and marine environments. They receive substantial inputs from adjacent landscapes but the importance of resource subsidies is not well understood. Here, we test hypotheses for the effects of both terrestrial- and salmon-derived resource subsidies on the diet (inferred from stable isotopes of muscle tissue), size and percent nitrogen of the soft-shell clam (Mya arenaria), a sedentary estuarine consumer. We examine how these relationships shift across natural gradients among 14 estuaries that vary in upstream watershed size and salmon density on the central coast of British Columbia, Canada. We also test how assimilation and response to subsidies vary at smaller spatial scales within estuaries. The depletion and enrichment of stable isotope ratios in soft-shell clam muscle tissue correlated with increasing upstream watershed size and salmon density, respectively. The effects of terrestrial- and salmon-derived subsidies were also strongest at locations near stream outlets. When we controlled for age of individual clams, there were larger individuals with higher percent nitrogen content in estuaries below larger watersheds, though this effect was limited to the depositional zones below river mouths. Pink salmon exhibited a stronger effect on isotope ratios of clams than chum salmon, which could reflect increased habitat overlap as spawning pink salmon concentrate in lower stream reaches, closer to intertidal clam beds. However, there were smaller clams in estuaries that had higher upstream pink salmon densities, possibly due to differences in habitat requirements. Our study highlights the importance of upstream resource subsidies to this bivalve species, but that individual responses to subsidies can vary at smaller scales within estuaries.     

Salmon species, density and watershed size predict magnitude of marine enrichment in riparian food webs

Resource subsidies across habitat boundaries can structure recipient communities and food webs. In the northern Pacific region, bears Ursus spp. foraging on anadromous salmon Oncorhynchus spp. provide a key link between marine and terrestrial ecosystems, with salmon density, fish size and watershed size as potential predictors of the magnitude of marine subsidy to terrestrial habitats. We use nitrogen and carbon stable isotopes to provide an assessment of the patterns of marine‐enrichment in riparian plants (11 species, 4 guilds) and litter invertebrates (4 guilds) sampled from 27 watersheds in coastal British Columbia, Canada. Watersheds occurred in three geographical regions (Vancouver Island, mainland midcoast and Haida Gwaii) and varied in size, and in biomass (kg m^?1 of spawning length) and species of salmon (chum O. keta, pink O. gorbuscha and coho O. kisutch). δ¹⁵N values in all plant species and invertebrate guilds were positively predicted by total salmon biomass (kg m^?1) and negatively predicted by watershed size. We observed replicated parallel slopes among plant species and invertebrate guilds across the gradient in salmon biomass, with differences in means hypothesized to be due to plant fractionation and animal trophic position. As such, we derived a watershed δ¹⁵N‐index averaged across guilds, and using an information theoretic approach we find that the biomass of chum salmon is a much stronger predictor of the δ¹⁵N‐index than either pink or coho salmon, or the sum biomass of all species. The top linear model contained chum biomass and watershed size. Chum salmon biomass independently predicted δ¹⁵N‐index variation in all three regions of British Columbia. Chum salmon are larger than pink or coho and provide an energetic reward for bears that facilitates carcass transfer, tissue selective foraging, and nutrient distribution by insect scavengers. Analyses of biodiversity and habitat data across many watersheds moves towards a long‐term goal in fisheries ecology to better integrate ecosystem values in salmon conservation.     

Trophic ecology of Pacific salmon (<Emphasis Type="Italic">Oncorhynchus</Emphasis> spp.) in the ocean: a synthesis of stable isotope research

Increasing interest in the marine trophic dynamics of Pacific salmon has been motivated by the recognition of their sensitivity to changing climate and to the competitive effects of hatchery fish on wild stocks. It has become more common to use stable isotopes to supplement traditional diet studies of salmon in the ocean; however, there have been no integrated syntheses of these data to determine whether stable isotope analyses support the existing conventional wisdom of feeding strategies of the Pacific salmon. We performed a meta-analysis of stable isotope data to examine the extent of trophic partitioning among five species of Pacific salmon during their marine lives. Pink, sockeye, and chum salmon showed very high overlap in resource use and there was no consistent evidence for chum relying on alternative food webs dominated by gelatinous zooplankton. δ¹⁵N showed that Chinook and coho salmon fed at trophic levels higher than the other three species. In addition, these two species were distinctly enriched in ¹³C, suggesting more extensive use of coastal food webs compared to the more depleted (pelagic) signatures of pink, sockeye, and chum salmon. This paper presents the first synthesis of stable isotope work on Pacific salmon and provides δ¹⁵N and δ¹³C values applicable to research on the fate of the marine derived nutrients these organisms transport to freshwater and riparian ecosystems.     

Food spectra of the farm juveniles of the Chum Salmon <Emphasis Type="Italic">Oncorhynchus keta</Emphasis> (Salmoniformes,Salmonidae) in the Ryazanovka River (Southern Primor’e)

Qualitative data are presented on the diet of juvenile chum salmon Oncorhynchus keta released from the Ryazanovka experimental fish hatchery farm situated in the south of Primor’e Territory. The stomach fullness, frequency of occurrence of food items, their average weight value, and selectivity are estimated. Food spectra of chum salmon juveniles during downstream migration comprised representatives of 51 taxa of bottom freshwater invertebrates and terrestrial insects. The bulk of the diet consisted of larvae of aquatic insects—chironomids, other Diptera, and mayflies. Their diversity increased with the growth of fry. Fry of chum salmon had a wide spectrum of feeding preference, favorite food items were larvae and pupae of chironomids.     

Do Low-Mercury Terrestrial Resources Subsidize Low-Mercury Growth of Stream Fish? Differences between Species along a Productivity Gradient

Low productivity in aquatic ecosystems is associated with reduced individual growth of fish and increased concentrations of methylmercury (MeHg) in fish and their prey. However, many stream-dwelling fish species can use terrestrially-derived food resources, potentially subsidizing growth at low-productivity sites, and, because terrestrial resources have lower MeHg concentrations than aquatic resources, preventing an increase in diet-borne MeHg accumulation. We used a large-scale field study to evaluate relationships among terrestrial subsidy use, growth, and MeHg concentrations in two stream-dwelling fish species across an in-stream productivity gradient. We sampled young-of-the-year brook trout (Salvelinus fontinalis) and Atlantic salmon (Salmo salar), potential competitors with similar foraging habits, from 20 study sites in streams in New Hampshire and Massachusetts that encompassed a wide range of aquatic prey biomass. Stable isotope analysis showed that brook trout used more terrestrial resources than Atlantic salmon. Over their first growing season, Atlantic salmon tended to grow larger than brook trout at sites with high aquatic prey biomass, but brook grew two-fold larger than Atlantic salmon at sites with low aquatic prey biomass. The MeHg concentrations of brook trout and Atlantic salmon were similar at sites with high aquatic prey biomass and the MeHg concentrations of both species increased at sites with low prey biomass and high MeHg in aquatic prey. However, brook trout had three-fold lower MeHg concentrations than Atlantic salmon at low-productivity, high-MeHg sites. These results suggest that differential use of terrestrial resource subsidies reversed the growth asymmetry between potential competitors across a productivity gradient and, for one species, moderated the effect of low in-stream productivity on MeHg accumulation.     

Tracing biogeochemical subsidies from glacier runoff into Alaska's coastal marine food webs

Nearly half of the freshwater discharge into the Gulf of Alaska originates from landscapes draining glacier runoff, but the influence of the influx of riverine organic matter on the trophodynamics of coastal marine food webs is not well understood. We quantified the ecological impact of riverine organic matter subsidies to glacier‐marine habitats by developing a multi‐trophic level Bayesian three‐isotope mixing model. We utilized large gradients in stable (δ¹³C, δ¹⁵N, δ²H) and radiogenic (Δ¹⁴C) isotopes that trace riverine and marine organic matter sources as they are passed from lower to higher trophic levels in glacial‐marine habitats. We also compared isotope ratios between glacial‐marine and more oceanic habitats. Based on isotopic measurements of potential baseline sources, ambient water and tissues of marine consumers, estimates of the riverine organic matter source contribution to upper trophic‐level species including fish and seabirds ranged from 12% to 44%. Variability in resource use among similar taxa corresponded to variation in species distribution and life histories. For example, riverine organic matter assimilation by the glacier‐nesting seabirds Kittlitz's murrelet (Brachyramphus brevirostris) was greater than that of the forest‐nesting marbled murrelet (B. marmoratus). The particulate and dissolved organic carbon in glacial runoff and near surface coastal waters was aged (12100–1500 years BP ¹⁴C‐age) but dissolved inorganic carbon and biota in coastal waters were young (530 years BP ¹⁴C‐age to modern). Thus terrestrial‐derived subsidies in marine food webs were primarily composed of young organic matter sources released from glacier ecosystems and their surrounding watersheds. Stable isotope compositions also revealed a divergence in food web structure between glacial‐marine and oceanic sites. This work demonstrates linkages between terrestrial and marine ecosystems, and facilitates a greater understanding of how climate‐driven changes in freshwater runoff have the potential to alter food web dynamics within coastal marine ecosystems in Alaska.     

The survivability of the ectoparasitic flagellate Ichthyobodo necator on chum salmon fry (Oncorhynchus keta) in seawater and comparison to Ichthyobodo sp. on Japanese flounder (Paralichthys olivaceus)

Experimental studies revealed that a freshwater ectoparasitic flagellate Ichthyobodo necator (Henneguy, 1883) could survive and reproduce in seawater after infected chum salmon fry, Oncorhynchus keta (Walbaum), were transferred directly from fresh water to 33% seawater. Minor morphological changes (slight reduction in body width, loss of twistlike wrinkles on body surface, and reduction in contractile vacuoles) were observed in the attached form of I. necator following transfer to seawater. The field survey also confirmed that I. necator occurs on chum salmon fry in seawater estuaries (salinity 17-34%) and in freshwater habitats. It was assumed that I. necator acquired salinity tolerance as a result of adapting to the migratory behavior of its anadromous host. Two morphologically similar bodonids, I. necator from chum salmon and Ichthyobodo sp. from marine Japanese flounder, Paralichthys olivaceus (Temminck and Schlegel), were differentiated by cross-infection experiments. Thus, the parasite from marine flounder should be regarded as a separate species from I. necator.     

Marine subsidy promotes spatial and dietary niche variation in an omnivore,the Keen’s mouse (Peromyscus keeni)

Marine‐derived resource subsidies can generate intrapopulation variation in the behaviors and diets of terrestrial consumers. How omnivores respond, given their multiple trophic interactions, is not well understood. We sampled mice (Peromyscus keeni) and their food sources at five sites on three islands of the Central Coast of British Columbia, Canada, to test predictions regarding variation in the spatial behavior and consumption of marine‐subsidized foods among individuals. About 50% of detections (n = 27 recaptures) occurred at traps closest to shoreline (25 m), with capture frequencies declining significantly inland (up to 200 m). Stable isotope signatures (δ ¹³C and δ ¹⁵N), particularly δ ¹⁵N, in plant foods, forest arthropod prey, and mouse feces were significantly enriched near shorelines compared with inland, while δ ¹³C patterns were more variable. Bayesian isotope mixing models applied to isotope values in mouse hair indicated that over one‐third (35–37%) of diet was comprised of beach‐dwelling arthropods, a marine‐derived food source. Males were more abundant near the shoreline than females and consumed more marine‐derived prey, regardless of reproductive status or availability of other food sources. Our results identify how multiple pathways of marine nutrient transfer can subsidize terrestrial omnivores and how subsets of recipient populations can show variation in spatial and dietary response.     

Causes and consequences of thermal tolerance limits in rocky intertidal porcelain crabs, genus petrolisthes

Vertical zonation of intertidal organisms, from the shallowsubtidal to the supralittoral zones, is a ubiquitous featureof temperate and tropical rocky shores. Organisms that livehigher on the shore experience larger daily and seasonal fluctuationsin microhabitat conditions, due to their greater exposure toterrestrial conditions during emersion. Comparative analysesof the adaptive linkage between physiological tolerance limitsand vertical distribution are the most powerful when the studyspecies are closely related and occur in discrete vertical zonesthroughout the intertidal range. Here, I summarize work on thephysiological tolerance limits of rocky intertidal zone porcelaincrab species of the genus Petrolisthes to emersion-related heatstress. In the eastern Pacific, Petrolisthes species live throughouttemperate and tropical regions, and are found in discrete verticalintertidal zones in each region. Whole organism thermal tolerancelimits of Petrolisthes species, and thermal limits of heartand nerve function reflect microhabitat conditions. Speciesliving higher in the intertidal zone are more eurythermal thanlow-intertidal congeners, tropical species have the highestthermal limits, and the differences in thermal tolerance betweenlow- and high-intertidal species is greatest for temperate crabs.Acclimation of thermal limits of high-intertidal species isrestricted as compared to low-intertidal species. Thus, becausethermal limits of high-intertidal species are near current habitattemperature maxima, global warming could most strongly impactintertidal species.     

Functional response of juvenile pink and chum salmon: effects of consumer size and two types of zooplankton prey

Feeding rate experiments were conducted for pink salmon Oncorhynchus gorbuscha fry [mean fork length ( L _F) 39 mm], juveniles (103–104 mm L _F) and juvenile chum salmon Oncorhynchus keta (106–107 mm L _F). Fishes were presented with small copepod ( Tisbi sp.) or larger mysid shrimp ( Mysidopsis bahia ) prey at varying densities ranging from 1 to 235 prey l⁻¹ in feeding rate experiments conducted at water temperatures ranging from 10·5 to 12·0° C under high light levels and low turbidity conditions. Juvenile pink and chum salmon demonstrated a type II functional response to mysid and copepod prey. Mysid prey was readily selected by both species whereas the smaller bodied copepod prey was not. When offered copepods, pink salmon fry fed at a higher maximum consumption rate (2·5 copepods min⁻¹) than larger juvenile pink salmon (0·4 copepods min⁻¹), whereas larger juvenile chum salmon exhibited the highest feeding rate (3·8 copepods min⁻¹). When feeding on mysids, the maximum feeding rate for larger juvenile pink (12·3 mysids min⁻¹) and chum (11·5 mysids min⁻¹) salmon were similar in magnitude, and higher than feeding rates on copepods. Functional response models parameterized for specific sizes of juvenile salmon and zooplankton prey provide an important tool for linking feeding rates to ambient foraging conditions in marine environments, and can enable mechanistic predictions for how feeding and growth should respond to spatial-temporal variability in biological and physical conditions during early marine life stages.     

Habitat-related differences in recruitment and survival of early recruits of the invasive <Emphasis Type="Italic">Sargassum muticum</Emphasis> (Phaeophyta,Sargassaceae) in northern Portugal

Habitat-related differences in recruitment and early survival of the invasive seaweed Sargassum muticum were investigated in relation to the lunar phase (gamete release trigger) and meso-grazing between mid-intertidal and low-intertidal pools in northern Portugal. In situ experiments were performed to determine recruitment and to test the effect of intertidal level (mid-intertidal and low-intertidal pools), grazing (no cage, cage and open cage), and lunar phase (full, new and each quarter moon) on recruit survival during reproductive seasons in two consecutive years. Our results show clearly that recruitment and survival were higher in the mid-intertidal than in low-intertidal environment in both study periods. Caging treatments affected recruit survival in both environments, however, differences in patterns suggesting differences in grazing pressure due to differences in grazer density and/or the assemblages. It therefore seems that grazing on early recruits of this invader might contribute to the resistance to proliferation. Mid-intertidal pools constitute a more favourable environment for recruits of S. muticum than the low-intertidal, which may explain the mainly mid-intertidal distribution of the species in Portugal.     

Movers and shakers: nutrient subsidies and benthic disturbance predict biofilm biomass and stable isotope signatures in coastal streams

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Carcass analogues provide marine subsidies for macroinvertebrates and juvenile Atlantic salmon in temperate oligotrophic streams

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Stable Isotope Ratios as a Tool for Assessing Changes in Carbon and Nutrient Sources in Antarctic Terrestrial Ecosystems

Samples of an angiosperm species, nine lichen species and a terrestrial alga, were collected from a variety of Antarctic terrestrial habitats, and were analysed for C and N stable isotope composition. Collections were made along natural gradients, the marine gradient, running from the sea coast inland and the moisture gradient, determined by melt water and precipitation runoff, and running towards the sea coast. Considerable variation in stable isotope ratios was found; δ¹³C values ranged between −16 and −32‰ and δ¹⁵N values between −23 and +23‰ The variation in stable carbon isotope ratios could be attributed in part to species specific differences, but differences in water availability also played a role, as was shown for the terrestrial alga Prasiola crispa and the lichen species Usnea antarctica. The differences in the isotope ratios of nitrogen could be retraced to the origin of nitrogen: marine or terrestrial. The nitrogen stable isotope ratios were influenced by both the marine gradient from the sea inland and the melt water and precipitation flow running in the opposite direction, towards the sea. This was shown for the lichen species Turgidosculum complicatulum and the angiosperm species Deschampsia antarctica. The variation in the C and N stable isotope ratios can be used to determine sources and pathways of N and changes in the water availability in Antarctic terrestrial ecosystems. Contrary to earlier reports the use of stable N isotope ratios is possible in this case because of the relative simplicity of the structure of the Antarctic terrestrial ecosystems.     

Effects of spawning Pacific salmon on terrestrial invertebrates: Insects near spawning habitat are isotopically enriched with nitrogen‐15 but display no differences in body size

When Pacific salmon (Oncorhynchus spp.) spawn and die, they deliver marine‐derived nutrient subsidies to freshwater and riparian ecosystems. These subsidies can alter the behavior, productivity, and abundance of recipient species and their habitats. Isotopes, such as nitrogen‐15 (¹⁵N), are often used to trace the destination of marine‐derived nutrients in riparian habitats. However, few studies have tested for correlations between stable isotopes and physiological responses of riparian organisms. We examined whether increases in δ ¹⁵N in terrestrial insect bodies adjacent to salmon spawning habitat translate to changes in percent nitrogen content and body size. This involved comparisons between distance from a salmon‐bearing river, marine‐derived nutrients in soils and insects, soil moisture content, and body size and nitrogen content in two common beetle families (Coleoptera: Curculionidae, Carabidae). As predicted, δ¹⁵N in riparian soils attenuated with distance from the river but was unaffected by soil moisture. This gradient was mirrored by δ¹⁵N in the herbivorous curculionid beetles, whereas carabid beetles, which feed at a higher trophic level and are more mobile, did not show discernable patterns in their δ¹⁵N content. Additionally, neither distance from the river nor body δ¹⁵N content was related to beetle body size. We also found that nitrogen‐15 was not correlated with total percent nitrogen in insect bodies, meaning that the presence of spawning salmon did not increase the percent nitrogen content of these insects. We conclude that while salmon‐derived nutrients had entered terrestrial food webs, the presence of δ¹⁵N alone did not indicate meaningful physiological changes in these insects in terms of percent nitrogen nor body size. While stable isotopes may be useful tracers of marine‐derived nutrients, they cannot necessarily be used as a proxy for physiologically important response variables.     

Marine subsidies alter the diet and abundance of insular and coastal lizard populations

We evaluated the impact of marine materials on the diet and abundance of arthropodivorous lizards inhabiting islands and the coast of the Gulf of California. Here, marine materials are brought onto land by seabirds and by tidal action, and both subsidy pathways cause arthropod abundance to increase. We evaluated Uta stansburiana (side-blotched lizard) diets in three habitats defined by having: (1) no marine subsidies available, (2) only seabird-derived subsidies, and (3) only tidal-derived subsidies. Stable isotope data indicated that lizard diets are subsidized indirectly by seabird and tidal activity. For example, in coastal areas we determined that 40% of a lizard's diet contains arthropods that have consumed algae. Such subsidies may explain why we found that lizards in coastal areas occupy a significantly higher trophic position than lizards in unsubsidized areas. We analyzed eight years of survey data on all arthropodivorous lizards to determine if diet subsidies result in increased lizard abundance. We found that lizards were more abundant in coastal areas than they were in inland habitats, and that they were more abundant on islands with seabirds than on islands without seabird populations. This study provides insight into the importance and effect of marine-derived nutrients from multiple sources on vertebrate consumers inhabiting islands and coastal areas.     

Spatial variability of stable carbon and nitrogen isotope ratios in a Mediterranean coastal lagoon

Exploring the trophic pathway of organic matter within the Mauguio lagoon (southern France, western Mediterranean), we found spatial differences in the isotopic composition (both δ¹³C and δ¹⁵N values) of organic matter sources (primary producers, particulate and sedimentary organic matter), which were mirrored in the upper trophic levels (invertebrates and fish). On average, δ¹³C was heavier by about 1.5–2‰ in the location under marine influence than in the sites influenced by freshwater discharge. The opposite trend was found for δ¹⁵N, which attained maximum values in the north-central zone influenced by freshwater delivery. For both C and N stable isotope ratios, the highest spatial variability was found in organic matter sources (2–3‰), while invertebrates and fish exhibited less variability (\~1–2‰). The differences observed may be related to both anthropogenic (wastewater input) and natural (marine vs. terrestrial inputs) factors. Discharge of wastewater, which affects the innermost location, generally determines an increase in the relative abundance of ¹⁵N. In addition, terrestrially derived nutrients and organic matter, which also affect the innermost location, are known to determine a shift towards ¹³C-depleted values. Our results substantiate the finding that the analysis of carbon and nitrogen stable isotopes can help in elucidating origin and fate of organic matter in coastal lagoons, which are characterised by a great spatial variability and complexity.     

Relationships between growth traits and scale stable isotopes (δ13C, δ15N) of adult chum salmon Oncorhynchus keta in Hokkaido,Japan

Stable isotope analysis (SIA) in combination with growth analysis using scales collected from adult chum salmon Oncorhynchus keta migrating back to Hokkaido, Japan, was performed to describe the variation of isotopic composition of carbon (δ¹³C) and nitrogen (δ¹⁵N) in scales and to examine relationships with growth traits [age, fork length (FL), and relative growth ratio in the last growth period [(RGR^last)]. Scale stable isotope (SI) values in 3‐ to 6‐year‐old fish were highly variable, ranging from ?17.6‰ to ?14.3‰ for δ¹³C and 9.5‰ to 13.4‰ for δ¹⁵N. The δ¹⁵N was positively correlated with FL, and this tendency may indicate changes in trophic level with growth. Significant effect was not detected between δ¹⁵N and RGR^last, it can be inferred that factors potentially yielding high δ¹⁵N may not necessarily result in higher growth rates. No trend found between FL and δ¹³C may imply that there is no clear segregation in feeding locations between the 3‐ to 6‐year groups. This study provided basic information for scale SI values of chum salmon adults and indicated that SIA using scales could be a new approach to elucidating the trophic ecology of chum salmon.     

Marine resources subsidize insular rodent populations in the Gulf of California,Mexico

Inputs of energy and nutrients from one ecosystem may subsidize consumers in adjacent ones, with significant consequences for local communities and food webs. We used stable isotope and faecal pellet analysis to quantify use of ocean-derived resources by small mammals on islands in the Gulf of California, Mexico. Rodents were live-trapped on grids originating near shore and extending 125-200 m inland to evaluate the extent to which rodents transport marine nutrients inland, and to determine whether marine foods subsidize island populations, permitting higher densities than would be possible based on terrestrial resources alone. Both faeces and stable carbon and nitrogen isotopes revealed that omnivorous mice (Peromyscus maniculatus) consume ocean-derived prey, including littoral and supralittoral invertebrates, and that their diets differed markedly from those of granivorous rodents (Chaetodipus rudinoris). On a small, seabird roosting island, marine prey were important in the diet of mice regardless of their proximity to shore, underscoring the pervasive influence of the ocean on small islands with relatively large coastline area. On a large island, however, consumption of marine foods declined sharply > or =50 m from shore, which suggests that mice are poor conduits of inland movement of energy and nutrients from the sea. Marine resources seemed to act as subsidies for omnivorous rodents: more P. maniculatus were captured near shore than farther inland and there was an inverse relationship between island area and rodent abundance, suggesting that small islands with large amounts of marine inputs support the highest population densities. Patterns of local and island-wide abundance of P. maniculatus are likely the result of several interacting factors, including frustrated dispersal, competition with C. rudinoris, and the absence of predators. We speculate, however, that the availability of marine resources allows P. maniculatus to reach high densities and to persist on small islands in the Gulf despite low and unpredictable terrestrial productivity. Spatial trophic subsidies thus provide a possible mechanistic explanation for the widely reported inverse relationship between population density and island or habitat area.