An experimental investigation of interactions in snail-macrophyte-epiphyte systems

Summary An experimental investigation under field conditions of enclosures containing freshwater pulmonate snails, the macrophyteCeratophyllum demersum and epiphytes, produced evidence of beneficial interactions.Ceratophyllum growth, measured in terms of stem length, numbers of leaf-nodes and growing tips and leaf survival was significantly enhanced in the presence of snails. This effect was attributed to the increased availability of plant nutrients of snail origin, such as phosphates and ammonia, as well as to the snails' action as “cleaning symbionts” in reducing the density of bacterial and algal epiphyton potentially deleterious to macrophytes. Principal component analysis revealed both seasonal and treatment effects of snail grazing on algal epiphyton. Small adnate algal species (e.g.Cocconeis placentula) survived grazing and benefited from the removal of larger, competitor, species. Snail densities increased in all treatments, despite high (86%) juvenile mortality. It is concluded that freshwater pulmonate snails are strong interactors in lentic habitats, enhancing the growth ofCeratophyllum and producing characterisic epiphyte communities. This benefits not only the snails, but also the plants and epiphytes that are associated with them. Thus the interactions between these component parts of the community can be considered as mutualistic.     

Development of a diverse epiphyte community in response to phosphorus fertilization

The role of terrestrial soil nutrient supply in determining the composition and productivity of epiphyte communities has been little investigated. In a montane Hawaiian rainforest, we documented dramatic increases in the abundance and species richness of canopy epiphytes in a forest that had been fertilized annually with phosphorus (P) for 15 years; there was no response in forest that had been fertilized with nitrogen (N) or other nutrients. The response of N-fixing lichens to P fertilization was particularly strong, although mosses and non-N-fixing lichens also increased in abundance and diversity. We show that enhancement of canopy P availability is the most likely factor driving the bloom in epiphytes. These results provide strong evidence that terrestrial soil fertility may structure epiphyte communities, and in particular that the abundance of N-fixing lichens – a functionally important epiphyte group – may be particularly sensitive to ecosystem P availability.     

EPIPHYTE-GRAZER INTERACTIONS ON ZOSTERA MARINA (ANTHOPHYTA: MONOCOTYLEDONES): EFFECTS OF DENSITY ON COMMUNITY FUNCTION1

Interactions between algal epiphytes and their grazers can have a significant impact on the structure and function of eelgrass (Zostera marina L.) meadows. In Puget Sound, the herbivorous gastropod Lacuna variegata Carpenter and its congeners appear to remove large quantities of the epiphytic community from eelgrass blades. When snails at typical field densities were used in microcosms, Lacuna significantly reduced epiphytic biomass and areal productivity. Biomass-specific productivity of the epiphytic community showed an increasing trend with increasing snail density. Epiphytic productivity increased nonlinearly with increasing epiphytic biomass. The commonly used logistic population growth formula adequately described this relationship. Grazing rate also increased nonlinearly with increasing epiphytic biomass. The Hailing equation adequately described the relationship between grazing rate and epiphytic biomass. The proportion of the epiphytic biomass found on the oldest blade of an eelgrass shoot was related linearly to epiphytic biomass, suggesting that a constant fraction of the epiphytic community is lost regardless of epiphytic density. Lacuna clearly removed large quantities of epiphytic material from eelgrass blades, significantly altering community function. Modified Lotka-Volterra equations, incorporating the logistic growth form and Hailing grazing equation, should prove useful in modeling the epiphyte–grazer interaction.     

The effects of predation by the mummichog, Fundulus heteroclitus (L.), on the abundance and distribution of the salt marsh snail, Melampus bidentatus (Say)

Enclosure and exclosure experiments were conducted in Canary Creek marsh to examine how predation by a killifish, Fundulus heteroclitus (L.), affects the abundance and size distribution of the salt marsh snail, Melampus bidentatus (Say). Enclosures (7.6 × 19.7 m) were stocked with Fundulus heteroclitus at densities of one-half normal, normal, twice normal, and four times normal density. Fish exclusion pens were also built. In both years of study, the mean density of snails increased significantly in pens where fish were excluded or their density reduced. During the same period in each year, the density of snails in pens containing higher than normal fish density fell by ≈50%.

Fish density also affected the size distribution of snails within pens. In both years, mean shell length of snails in the pen with the highest density of fish was significantly greater, and mean shell length of snails within fish exclusion pens was significantly lower than in all other treatments. Gape size limitation of F. heteroclitus causes selective predation on small snails and apparently is responsible for the difference in mean shell length among treatments.

Density and size distribution measurements of the natural Melampus bidentatus population in Canary Creek marsh were also taken. Larger snails were found to occur in the low marsh zone, whereas smaller snails occurred in the high marsh zone. Since the high marsh area is flooded less often than the low marsh, and the grass types found in the high marsh zone are known to afford protection from fish predation, the distribution of snails in the marsh is consistent with the idea that fish predation is an important factor influencing the distribution of snails in Canary Creek marsh.     

Richness and species composition of arboreal arthropods affected by nutrients and predators: a press experiment

A longstanding goal for ecologists is to understand the processes that maintain biological diversity in communities, yet few studies have investigated the combined effects of predators and resources on biodiversity in natural ecosystems. We fertilized nutrient limited plots and excluded insectivorous birds in a randomized block design, and examined the impacts on arthropods associated with the dominant tree in the Hawaiian Islands, Metrosideros polymorpha (Myrtaceae). After 33 months, the species load (per foliage mass) of herbivores and carnivores increased with fertilization, but rarified richness (standardized to abundance) did not change. Fertilization depressed species richness of arboreal detritivores, and carnivore richness dropped in caged, unfertilized plots, both because of the increased dominance of common, introduced species with treatments. Herbivore species abundance distributions were more equitable than other trophic levels following treatments, and fertilization added specialized native species without changing relativized species richness. Overall, bird removal and nutrient addition treatments on arthropod richness acted largely independently, but with countervailing influences that obscured distinct top-down and bottom-up effects on different trophic levels. This study demonstrates that species composition, biological invasions, and the individuality of species traits may complicate efforts to predict the interactive effects of resources and predation on species diversity in food webs.     

Top-down and bottom-up control of infauna varies across the saltmarsh landscape

Responses of infaunal saltmarsh benthic invertebrates to whole-ecosystem fertilization and predator removal were quantified in Plum Island Estuary, Massachusetts, USA. Throughout a growing season, we enriched an experimental creek on each flooding tide to 70 μM NO₃⁻ and 4 μM PO₄^− 3 (a 10× increase in loading above background), and we reduced Fundulus heteroclitus density by 60% in a branch of the fertilized and a reference creek. Macroinfauna and meiofauna were sampled in creek (mudflat and creek wall), marsh edge (tall-form Spartina alterniflora) and marsh platform (Spartina patens and stunted S. alterniflora) habitats before and after treatments were begun; responses were tested with BACI-design statistics. Treatment effects were most common in the mid-range of the inundation gradient. Most fertilization effects were on creek wall where ostracod abundance increased, indices of copepod reproduction increased and copepod and annelid communities were altered. These taxa may use epiphytes (that respond rapidly to fertilization) of filamentous algae as a food source. Killifish reduction effects on meiobenthic copepod abundance were detected at the marsh edge and suggest predator limitation. Fish reduction effects on annelids did not suggest top-down regulation in any habitat; however, fish reduction may have stimulated an increased predation rate on annelids by grass shrimp. Interactions between fertilization and fish reduction occurred under S. patens canopy where indirect predator reduction effects on annelids were indicated. No effects were observed in mudflat or stunted S. alterniflora habitats. Although the responses of infauna to fertilization and predator removal were largely independent and of similar mild intensity, our data suggests that the effects of ecological stressors vary across the marsh landscape.     

Intra- and interspecific competition among coexisting lotic snails

The competitive interactions of two lotic snails, Elimia cahawbensis and Elimia carinifera , were examined in a second-order spring-fed stream. We first demonstrated food limitation in laboratory microcosms where snails grew faster when exposed to enhanced periphyton levels. We then tested the magnitude and relative strengths of intra- and interspecific competition in similar stream and laboratory mesocosm experiments. Treatments were maintained in Plexiglas enclosures over a 7-week period with 0, 1×, 2× and 4× ambient biomass of each species alone, as well as mixed species treatments at 2× and 4× ambient. Snail responses to treatments were almost identical in field and laboratory experiments. Growth rates of both species were reduced by increased density of snails indicating strong intra- and interspecific competition among E. cahawbensis and E. carinifera . An analysis of the strengths of intra- and interspecific competition indicated minimal differences for either species, implying a lack of competitive dominance. Although periphyton biomass was generally highest without snails, there was little difference in periphyton biomass and snail production over the four-fold density range, regardless of species composition. These results suggest that E. cahawbensis and E. carinifera are functionally redundant with density-dependent responses in growth rate resulting in similar grazing pressure across a density gradient. This clearly demonstrates that species impact is not necessarily reflected by measures of abundance or biomass, and that secondary production should be considered.     

Effects of phosphorus enrichment and grazing snails on modern stromatolitic microbial communities

1. The effects of phosphorus enrichment and grazing snails on a benthic microbial community that builds stromatolic oncolites were examined in an experiment at Rio Mesquites, Cuatro Ciénegas, Mexico. Chemical analyses of stream water samples indicated that overall atomic ratios of total nitrogen (N) to total phosphorus (P) were approximately 110, indicating a strong potential for P‐limitation of microbial growth. 2. Phosphorus enrichment involved addition of 5 μmol Na₂HPO₄ L^?1 to streamside microcosms receiving intermittent inputs of stream water while grazer manipulation involved removal of the dominant grazer, the snail Mexithauma quadripaludium. After 7 weeks, we examined responses in organic matter content, C : N : P ratios, metabolism (P removal, primary production, dark respiration, and calcification), and microbial community structure using molecular fingerprinting of 16S rRNA genes. 3. Manipulation of snails did not affect response variables measured in these treatments (organic matter, C : P ratio, P removal rate). However, P enrichment significantly decreased the C : P and N : P ratios of surficial materials in the oncolites (organic matter content was unchanged), increased net and gross photosynthesis (oxygen consumption in the dark was unchanged), increased rates of calcification, and increased diatoms relative to cyanobacteria. Heterotrophic Eubacteria and Archaea were only modestly affected. Thus, our results indicate weak grazing effects but strong impacts of P in this benthic system. 4. We hypothesise that a state of severe P‐limitation is imposed on autotrophic production in this food web due, at least in part, to co‐precipitation of phosphate during calcite deposition. This produces severe P‐limitation of the benthic algae and cyanobacteria, resulting in high C : P ratio of microbial mats relative to the biomass of photoautotrophs (phytoplankton, terrestrial foliage) in other ecosystems. In turn, this high C : P ratio is likely to generate severe stoichiometric constraints on the herbivores, thus limiting their populations and resulting in weak overall grazing impacts.     

NUTRIENT EFFECTS ON SEAGRASS EPIPHYTE COMMUNITY STRUCTURE IN FLORIDA BAY1

A field experiment was employed in Florida Bay investigating the response of seagrass epiphyte communities to nitrogen (N) and phosphorus (P) additions. While most of the variability in epiphyte community structure was related to uncontrolled temporal and spatial environmental heterogeneity, P additions increased the relative abundance of the red algae–cyanobacterial complex and green algae, with a concomitant decrease in diatoms. When N was added along with P, the observed changes to the diatoms and the red algae–cyanobacterial complex were in the same direction as P‐only treatments, but the responses were decreased in magnitude. Within the diatom community, species relative abundances, species richness, and diversity responded weakly to nutrient addition. P additions produced changes in diatom community structure that were limited to summer and were stronger in eastern Florida Bay than in the western bay. These changes were consistent with well‐established temporal and spatial patterns of P limitation. Despite the significant change in community structure resulting from P addition, diatom communities from the same site and time, regardless of nutrient treatment, remained more similar to one another than to the diatom communities subject to identical nutrient treatments from different sites and times. Overall, epiphyte communities exhibited responses to P addition that were most evident at the division level.     

Trophic facilitation by the oystercatcher Haematopus palliatus Temminick on the scavenger snail Buccinanops globulosum Kiener in a Patagonian bay

This study investigated the role of the American oystercatcher (Haematopus palliatus) as a resource subsidizer for the scavenger snail Buccinanops globulosum in a northern Patagonian bay (40°45′S, 64°56′W, San Antonio Bay, Argentina). The most frequent food item for the snails was dead crabs Cyrtograpsus angulatus Dana, and the snails preferred this item. In the field, most dead crabs (78%) resulted from oystercatcher predation. Field densities of dead crabs were within the 95% confidence limits of the estimated densities produced by oystercatcher foraging activity, suggesting that a large proportion of carrion available for snails is a byproduct of oystercatcher predation. Dead crabs with injuries were more rapidly detected and consumed by snails, probably because injuries produced by oystercatchers increase leakage of body fluids and facilitate penetration of the proboscis of the snails. Our results suggest that oystercatcher predation subsidize this scavenger snail by increasing availability of food, decreasing variability in their provision and facilitating their consumption.     

Relative effects of nutrient enrichment and grazing on epiphyte-macrophyte (Zostera marina L.) dynamics

The independent and interactive effects of nutrient concentration and epiphyte grazers on epiphyte biomass and macrophyte growth and production were examined in Zostera marina L. (eelgrass) microcosms. Experiments were conducted during early summer, late summer, fall, and spring in a greenhouse on the York River estuary of Chesapeake Bay. Nutrient treatments consisted of ambient or enriched (3× ambient) concentrations of inorganic nitrogen (ammonium nitrate) and phosphate. Grazer treatments consisted of the presence or absence of field densities of isopods, amphipods, and gastropods. epiphyte biomass increased with both grazer removal and nutrient enrichment during summer and spring experiments. The effect of grazers was stronger than that of nutrients. There was little epiphyte response to treatment during the fall, a result possibly of high ambient nutrient concentrations and low grazing pressure. Under low grazer densities of early summer, macrophyte production (g m^–2 d^–1) was reduced by grazer removal and nutrient enrichment independently. Under high grazer densities of late summer, macrophyte production was reduced by enrichment only with grazers absent. During spring and fall there were no macrophyte responses to treatment. The relative influence of epiphytes on macrophyte production may have been related to seasonally changing water temperature and macrophyte requirements for light and inorganic carbon.     

Can the freshwater snail Bellamya aeruginosa (Mollusca) affect phytoplankton community and water quality?

The purpose of this microcosm experiment was to determine whether the freshwater snail Bellamya aeruginosa affected phytoplankton community and water quality. Three treatments of different snail densities (low, medium, and high) and a control (no snails) were set up in twelve enclosures. Chlorophyll a (chl a), transparency (SD), water temperature (WT), dissolved oxygen (DO), pH, nutrients, and abundance of phytoplankton were determined on days 0, 3, 7, 11, 15, 23, 31, 46, and 61. The total chl a concentration decreased and SD increased in the treatments, whereas the proportion of cyanobacteria chl a increased. Bellamya seemed to result indirectly in a decrease of DO, by reducing the algal biomass rather than by respiration. A significant decrease in nitrogen–phosphorus ratios was observed in the treatments. In the enclosed system the abundance of colonial green algae decreased over time whereas that of cyanobacteria and flagellates increased. Principal response curves showed that both phytoplankton community and water quality in the medium and high-density treatments were consistently different from in the control. These results indicate that the presence of snails resulted in a significant change of water physicochemical properties and phytoplankton community.     

The Conservation Value of Secondary Forests for Vascular Epiphytes in Central Panama

Secondary forests that develop following land abandonment could compensate for the losses of diversity and structure that accompany deforestation of old‐growth forests in tropical regions. Whether secondary forests can harbor similar species richness, density, and composition of old‐growth forests for vascular epiphytes remains largely unknown for secondary forests older than 50 yr. We examined community structure (species richness, density, and species composition) of vascular epiphytes in older secondary forests between 35 and 115 yr after land abandonment and nearby old‐growth forests to determine if the community structure of epiphytes in secondary forests approaches that of old‐growth forests over time. The recovery of epiphyte species richness was rapid with 55‐year‐old forests containing 65 percent of old‐growth epiphyte species richness. Secondary forest epiphyte communities were found to be statistically nested within secondary forests older in age and within old‐growth forests. Similarity of epiphyte communities to old‐growth forests increased to 75 percent, 115 yr after abandonment. This study suggests that secondary forests will likely recover old‐growth epiphyte richness and composition given enough time. Epiphyte densities did not recover quickly with 55‐year‐old forests having 14 percent and 115‐year‐old forests having only 49 percent of the density of old‐growth forest epiphytes. The low density of epiphytes in secondary forests could impact rainforest diversity and function. We conclude that in less than 115 yr, although secondary moist forests have high conservation value for some aspects of community structure, they are unlikely to compensate biologically for the loss of diversity and ecosystem function that high epiphyte densities provide.     

Community regulation by herbivore parasitism and density: Trait-mediated indirect interactions in the intertidal

The abundant herbivorous mud-snail Hydrobia ulvae is an ecosystem engineer in soft-bottom intertidal habitats due to its grazing and bioturbation activity. However, mud snails are commonly infected by trematodes that reduce their overall activity, which in turn may affect their impact on the surrounding benthic community. To test this hypothesis, we performed field experiments manipulating both the abundance of uninfected snails (0, 7500 and 15.000 ind. m^- 2) and the level of snail parasitism (0, 33 and 100% trematode prevalence) on a Danish mud-flat. The results showed that increasing snail abundance and parasitism generally had opposite effects on the community of microphytobenthos and zoobenthos. Increasing snail density increased the chlorophyll-a concentration in the substrate (enhancement), whereas increasing parasitism decreased it. In accordance, the benthic primary producers were generally less nutrient limited at high snail density and mostly so at high levels of snail parasitism. Moreover, epipsammic diatoms were favoured over epipelic diatoms at increasing snail density, whereas the opposite was evident at increasing snail parasitism. At the community level, increasing snail density increased evenness among epipelic diatoms, whereas increasing snail parasitism decreased evenness and species diversity. Probably through the action of trophic cascades and varying levels of disturbance, the zoobenthic community was influenced by experimental treatments as well. The indirect effects of snail parasitism influenced significantly the abundance of more faunal species (seven) than did snail density (two). At the community level, increasing snail density decreased evenness and lowest species richness coincided with intermediate snail density. In contrast, increasing snail parasitism resulted in increasing evenness and peaking species richness at intermediate level of parasitism. Together, the results show that parasites solely through their impact on the behaviour of a single community member can be significant indirect determinants of community organisation and function.     

The influence of temporally variable predation risk on indirect interactions in an aquatic food chain

We know little about how temporally variable predation risk influences prey behavior. The risk allocation hypothesis predicts that prey facing more frequent risk should show weak anti-predator responses, and should be particularly active foragers during rare periods of safety, compared to prey facing infrequent risk. Several studies offer support for the risk allocation hypothesis, but how these responses might propagate through the larger ecological community remains largely unknown. We experimentally investigated the relative strength of trait- and density-mediated indirect effects of a predator on its prey’s resource across predation treatments that varied the lethality (caged or free-swimming predators) and temporal variability (always, often, or sometimes present) of predation. We performed this experiment in pond mesocosms using a giant water bug predator (Belostoma lutarium), an herbivorous pond snail (Physa gyrina), and algae as the basal resource. Snails greatly reduced the abundance of their algal resource when in the absence of predation. Lethal predation at low and medium intensities had significant positive indirect effects on the abundance of algae, mostly by reducing snail density. Snails responded behaviorally to high levels of deadly predation by foraging more and hiding less than in other situations, as predicted by the risk allocation hypothesis, and thus ameliorated the density-mediated indirect effects of predators on algae. Behavioral responses to caged predators, and the subsequent trait-mediated indirect effects, were negligible regardless of predation intensity. Our previous work has demonstrated that trait-mediated indirect effects are weak when resources are abundant, as they were in this experiment. This work demonstrates that temporal variation in predation intensity plays a key role in determining the relative strength of TMIIs and DMIIs in an aquatic food chain.     

Field evidence of trait-mediated indirect interactions in a rocky intertidal food web

Studies on the implications of food web interactions to community structure have often focused on density-mediated interactions between predators and their prey. This approach emphasizes the importance of predator regulation of prey density via consumption (i.e. lethal effects), which, in turn, leads to cascading effects on the prey's resources. A more recent and contrasting view emphasizes the importance of non-lethal predator effects on prey traits (e.g. behaviour, morphology), or trait-mediated interactions. On rocky intertidal shores in New England, green crab ( Carcinus maenas ) predation is thought to be important to patterns of algal abundance and diversity by regulating the density of herbivorous snails ( Littorina littorea ). We found, however, that risk cues from green crabs can dramatically suppress snail grazing, with large effects on fucoid algal communities. Our results suggest that predator-induced changes in prey behaviour may be an important and under-appreciated component of food web interactions and community dynamics on rocky intertidal shores.     

The effects of nutrient addition on floral characters and pollination in two subalpine plants, <Emphasis Type="Italic">Ipomopsis aggregata</Emphasis> and <Emphasis Type="Italic">Linum lewisii</Emphasis>

The availability of soil and pollination resources are main determinants of fitness in many flowering plants, but the degree to which each is limiting and how they interact to affect plant fitness is unknown for many species. We performed resource (water and nutrients) and pollination (open and supplemental) treatments on two species of flowering plants, Ipomopsis aggregata and Linum lewisii, that differed in life-history, and we measured how resource addition affected floral characters, pollination, and reproduction (both male and female function). We separated the direct effects of resources versus indirect effects on female function via changes in pollination using a factorial experiment and path analysis. Resource addition affected I. aggregata and L. lewisii differently. Ipomopsis aggregata, a monocarp, responded to fertilization in the year of treatment application, increasing flower production, bloom duration, corolla width, nectar production, aboveground biomass, and pollen receipt relative to control plants. Fertilization also increased total seed production per plant, and hand-pollination increased seeds per fruit in I. aggregata, indicating some degree of pollen limitation of seed production. In contrast, fertilization had no effect on growth or reproductive output in the year of treatment on L. lewisii, a perennial, except that fertilization lengthened bloom duration. However, delayed effects of fertilization were seen in the year following treatment, with fertilized plants having greater aboveground biomass, seeds per fruit, and seeds per plant than control plants. In both species, there were no effects of resource addition on male function, and the direct effects of fertilization on female function were relatively stronger than the indirect effects via changes in pollination. Although we studied only two plant species, our results suggest that life-history traits may play an important role in determining the reproductive responses of plants to soil nutrient and pollen additions.     

A manipulative study of macroinvertebrate grazers in Hong Kong streams: do snails compete with insects?

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Phlorotannins versus other factors affecting epiphyte abundance on the kelp Ecklonia radiata

We examined factors affecting the abundance and distribution of epiphytes (fouling) on the sublittoral kelp Ecklonia radiata. We first assessed the importance of phlorotannins as chemical defences against epiphytes by (a) correlating epiphyte loads on different parts of the thallus with the phlorotannin content of those tissues, and (b) experimentally testing the effects of variation in phlorotannin concentration against the settlement and growth of gametes of Ulva lactuca, a common epiphyte in the system. Tissue phlorotannin content was, at best, only weakly related to epiphyte loads, with r ² values typically <0.10. Inhibition of Ulva gametes only occurred at concentrations >10 mg l⁻¹, which is 5 orders of magnitude greater than levels of phlorotannins in the water column around beds of E. radiata, and 1–3 orders of magnitude greater than estimated levels in the boundary layer at the surface of the plant. We concluded that phlorotannins have a negligible impact on patterns of epiphytism on E. radiata, and proceeded to investigate other factors influencing the distribution and abundance of epiphytes. In our samples the relative age of different parts of the thallus was strongly correlated with epiphyte abundance, with epiphyte densities greatest on the oldest tissue and least on the youngest. Distal parts of the thalli also had greater epiphyte loads than basal parts. Field experiments in which kelp tissue was suspended at two heights in an E. radiata forest for varying lengths of time confirmed the importance of the length of time that the tissue was in the water, and its height in the water column, to the development of an epiphyte community. Comparison of epiphyte loads on tissue from primary (smooth) and secondary (rough) laminae in these experiments indicated that surface rugosity also affected fouling. Macroherbivores were rare on E. radiata, and abundances of mesofauna and epiphytes were positively related, suggesting that grazers were not important determinants of patterns of epiphyte abundance. Although phlorotannins have been previously suggested to play an important role as defences against epiphytes, we suggest that water-soluble compounds such as phlorotannins are less likely to be effective defences against epiphytes than non-polar metabolites, which can adhere to the surface of the producing organism.     

Are vascular epiphytes nitrogen or phosphorus limited? A study of plant (15) N fractionation and foliar N : P stoichiometry with the tank bromeliad Vriesea sanguinolenta

Although there is unambiguous evidence for vascular epiphytic plants to be limited by insufficient water and nutrient supply under natural conditions, it is an open debate whether they are primarily phosphorus (P) or nitrogen (N) limited. Plant (15) N fractionation and foliar N : P stoichiometry of a tank epiphyte (Vriesea sanguinolenta), and its response to combined N-P fertilization, were studied under semi-natural conditions over 334 d to clarify the type of nutrient limitation. Plants collected in the field and experimental plants with limited nutrient supply showed significant plant (15) N fractionation (mean 5‰) and plant N : P ratios of c. 13.5. Higher relative growth rates and declines in plant (15) N fractionation (0.5‰) and in foliar N : P ratios to 8.5 in the high N-P treatment indicated that these epiphytes were P limited in situ. The critical foliar N : P ratio was 10.4, as derived from the breakpoint in the relationship between plant (15) N fractionation and foliar N : P. We interpret the widespread (15) N depletion of vascular epiphytes relative to their host trees as deriving from (15) N fractionation of epiphytes as a result of P limitation. High foliar N : P ratios (> 12) corroborate widespread P limitation (or co-limitation by N and P) of epiphytic bromeliads and, possibly, other epiphyte species.