Habitat-specific impacts of multiple consumers on plant population dynamics

Multiple consumers often attack seeds, seedlings, and adult plants, but their population-level consequences remain uncertain. We examined how insect and small mammal consumers influenced the demography and abundance of the perennial shrub, bush lupine (Lupinus arboreus). In grassland and dune habitats we established replicate experimental lupine populations in 81-m2 plots that were either protected from, or exposed to, herbivorous voles and granivorous mice (via fencing) and/or root feeding insects (via insecticide treatment). Populations were initiated with transplanted seedlings in 1999 and 2000. We followed the demography of these cohorts, subsequent generations, and the seed bank for 5.5 years. Voles and insects killed many seedlings in dune (1999 only) and grassland (1999 and 2000) habitats. After 2000, insects and voles had minimal effects on seedling or adult survival. Seed predation by granivorous mice, however, greatly depressed seedling recruitment, resulting in lower adult lupine abundance in control plots vs. those protected from rodents. In grasslands, initial effects of voles and insects on seedling survival produced large differences among treatments in adult plant density and the cumulative number of seeds produced throughout the experiment. Differences among grassland populations in seed rain, however, had little influence on the magnitude of seedling recruitment into this habitat. Instead, recruitment out of a preexisting seed bank compensated for the lack of seed production in populations exposed to consumers. Shading by dense adults in plots protected from consumers limited seedling establishment within these populations. Although differences among populations in cumulative seed rain did not influence adult establishment, populations protected from consumers accumulated substantially larger seed banks than controls. These results illustrate how density dependence, habitat-specific seed dynamics, and particular demographic impacts of consumers interact to shape plant population responses to consumers.     

The influence of size-specific indirect interactions in predator-prey systems

Nonlethal indirect interactions between predators often lead to nonadditive effects of predator number on prey survival and growth. Previous studies have focused on systems with at least two different predator species and one prey species. However, most predators undergo extreme ontological changes in phenotype such that interactions between different-sized cohorts of a predator and its prey could lead to nonadditive effects in systems with only two species. This may be important since different-sized individuals of the same species can differ more in their ecology than similar-sized individuals of different species. This study examined trait-mediated indirect effects in a two-species system including a cannibalistic predator with different-sized cohorts and its prey. I tested for these effects using larvae of two stream salamanders, Gyrinophilus porphyriticus (predator) and Eurycea cirrigera (prey), by altering the densities and combinations of predator size classes in experimental streams. Results showed that the presence of large individuals can significantly reduce the impact of density changes of smaller conspecifics on prey survival through nonlethal means. In the absence of large conspecifics, an increase in the relative frequency of small predators significantly increased predation rates, thereby reducing prey survival. However, with large conspecifics present, increasing the density of small predators did not decrease prey survival, resulting in a 14.3% lower prey mortality than predicted from the independent effects of both predator size classes. Small predators changed their microhabitat use in the presence of larger conspecifics. Prey individuals reduced activity in response to large predators but did not respond to small predators. Both predators reduced prey growth. These results demonstrate that the impact of a predator can be significantly altered by two different types of trait-mediated indirect effects in two-species systems: between different-sized cohorts and between different cohorts and prey. This study demonstrates that predictions based on simple numerical changes that assume independent effects of different size classes or ignore size structure can be strongly misleading. We need to account for the size structure within predator populations in order to predict how changes in predator abundance will affect predator-prey dynamics.     

Facilitation by Pinus flexilis during succession: a hierarchy of mechanisms benefits other plant species

Studies of facilitation have primarily been limited to single mechanisms, species, or environments. We examined interacting mechanisms governing the facilitative effects of Pinus flexilis on two later successional understory species, Pseudotsuga menziesii and Ribes cereum, in different microhabitats and seasons at the ecotone between the Rocky Mountain forests and Great Plains grasslands in Montana, USA. In field surveys, 69% of Pseudotsuga and 91% of Ribes were located beneath P. flexilis even though P. flexilis subcrowns accounted for a small proportion of available habitat. For three years, we monitored the survival of Pseudotsuga and Ribes seedlings experimentally planted beneath P. flexilis and in the open at a windward and a leeward site. Survival of both species was highest beneath P. flexilis at a site topographically protected from strong unidirectional winds (38% for Pseudotsuga and 63% for Ribes), and lowest at a windward site and in the open where tree crowns did not provide shelter from winds (2% and 6%, respectively). These results suggest that wind amelioration contributed to the facilitative effect of P. flexilis. However, even at the leeward site, where wind speed was low, survival of Pseudotsuga and Ribes was higher beneath P. flexilis, suggesting the importance of shade. To explore the relative importance of different mechanisms, we designed an experiment with six treatments: "shade," "shade + wind," "shade + drift," "wind," "drift," and a "control." After two years, we found shade to be of overwhelming importance for the survival of Pseudotsuga and Ribes. Without shade, no other treatments were significant, but once shade was provided, wind amelioration and snow pack accumulation increased survival of Pseudotsuga, suggesting that these different facilitative mechanisms functioned in a nested hierarchical manner: some mechanisms were important only when others were already functioning. Many studies have demonstrated multiple interacting mechanisms in the way that plants interact, but to our knowledge hierarchical interactive processes have not been previously documented. If the effects of positive or competitive mechanisms are often hierarchical, then studies of isolated mechanisms may not accurately assess their importance in nature.     

The Paradox of the Long‐Term Positive Effects of a North American Crayfish on a European Community of Predators

Abstract: Invasions of non‐native species are one of the major causes of losses of native species. In some cases, however, non‐natives may also have positive effects on native species. We investigated the potential facilitative effects of the North American red swamp crayfish (Procambarus clarkii) on the community of predators in southwestern Spain. To do so, we examined the diets of predators in the area and their population trends since introduction of the crayfish. Most predator species consumed red swamp crayfish, which sometimes occurred in over 50% of their diet samples. Moreover, the abundance of species preying on crayfish increased significantly in the area as opposed to the abundance of herbivores and to predator populations in other areas of Europe, where those predators are even considered threatened. Thus, we report the first case in which one non‐native species is both beneficial because it provides prey for threatened species and detrimental because it can drive species at lower trophic levels to extinction. Increases in predator numbers that are associated with non‐native species of prey, especially when some of these predators are also invasive non‐natives, may increase levels of predation on other species and produce cascading effects that threaten native biota at longer temporal and larger spatial scales. Future management plans should include the complexity of interactions between invasive non‐natives and the entire native community, the feasibility of successful removal of non‐native species, and the potential social and economic interests in the area.     

Variable flight initiation distance in incubating Eurasian curlew

Flight initiation distances (FIDs) of nesting birds approached by a predator likely reflect evolutionary stable strategies in which birds make trade-offs between adult survival and reproductive success. Here, we test if FID (a) had an impact on hatching success, (b) was adjusted to current conditions, and (c) was consistent for individual nests. All experiments were performed with a human approaching incubating Eurasian curlews Numenius arquata, a ground-nesting wader species under high egg predation pressure. Our results show that hatching success was higher in nests where the incubating parent left at intermediate FIDs compared to short and long ones, and that FID decreased with date and time of the evening. Further, FIDs from repeated approaches were not consistent within nests. We suggest that incubating Eurasian curlews follow a “surprise” strategy, where an element of randomness is superimposed on a context-adjusted norm to prevent predators from predicting their FID behaviour.     

Connected fragmented habitats facilitate stable coexistence dynamics

In this paper we endeavor to test the controversial ideas that exist about the role of fragmentation in a conservation context. In line with earlier understanding, we find that habitat fragmentation alone results in a strong detrimental effect (especially for the predator population). Connecting the fragmented habitats facilitates predator survival and hence prey survival as compared to the unconnected fragmented case. Our main result is counterintuitive: in the presence of a high quality predator, connected fragmented habitats ensure a better chance for coexistence than does even the unfragmented case. We explain why a connected fragmented habitat might thus be beneficial for the stabilization of the system, and how connections between sub-habitats are able to protect prey population from over-exploitation. In the model, habitat fragmentation is separated from the effects of habitat destruction, in order to better understand how populations react to habitat transformation.     

Lupine calls to Texas poison control centers, 1998–2005

Many lupine species (Genus Lupinus) contain toxic compounds, although the amount of these substances varies by the plant part and species. This investigation described the epidemiology of 138 lupine exposures reported to Texas poison centers during 1998–2005. Almost 88% of the lupine exposure calls occurred in March and April, and 90% of the calls came from the central portions of the state. Sixty-one percent of the calls involved female patients, and 91% of the patients were age <6 years. The lupine exposure occurred at the patient's own residence in 79% of the cases. Of the reported lupine exposures, 93% were handled outside of health care facilities, and, of those cases with a known medical outcome, few moderate or major effects, and no deaths, were reported. The most frequently noted clinical effect among cases during 2000–2005 was vomiting, affecting 8% of the cases. In conclusion, most reported lupine exposures in Texas came from the central parts of the state, occurred in spring, and involved young children. Typical reported lupine exposures in Texas were not likely to have adverse outcomes.     

Demographic heterogeneity, cohort selection, and population growth

Demographic heterogeneity--variation among individuals in survival and reproduction--is ubiquitous in natural populations. Structured population models address heterogeneity due to age, size, or major developmental stages. However, other important sources of demographic heterogeneity, such as genetic variation, spatial heterogeneity in the environment, maternal effects, and differential exposure to stressors, are often not easily measured and hence are modeled as stochasticity. Recent research has elucidated the role of demographic heterogeneity in changing the magnitude of demographic stochasticity in small populations. Here we demonstrate a previously unrecognized effect: heterogeneous survival in long-lived species can increase the long-term growth rate in populations of any size. We illustrate this result using simple models in which each individual's annual survival rate is independent of age but survival may differ among individuals within a cohort. Similar models, but with nonoverlapping generations, have been extensively studied by demographers, who showed that, because the more "frail" individuals are more likely to die at a young age, the average survival rate of the cohort increases with age. Within ecology and evolution, this phenomenon of "cohort selection" is increasingly appreciated as a confounding factor in studies of senescence. We show that, when placed in a population model with overlapping generations, this heterogeneity also causes the asymptotic population growth rate lambda to increase, relative to a homogeneous population with the same mean survival rate at birth. The increase occurs because, even integrating over all the cohorts in the population, the population becomes increasingly dominated by the more robust individuals. The growth rate increases monotonically with the variance in survival rates, and the effect can be substantial, easily doubling the growth rate of slow-growing populations. Correlations between parent and offspring phenotype change the magnitude of the increase in lambda, but the increase occurs even for negative parent-offspring correlations. The effect of heterogeneity in reproductive rate on lambda is quite different: growth rate increases with reproductive heterogeneity for positive parent-offspring correlation but decreases for negative parent-offspring correlation. These effects of demographic heterogeneity on lambda have important implications for population dynamics, population viability analysis, and evolution.     

Antipredator defenses along a latitudinal gradient in Rana temporaria

Antipredator defenses are expected to decrease toward higher latitudes because predation rates are predicted to decrease with latitude. However, latitudinal variation in predator avoidance and defense mechanisms has seldom been studied. We studied tadpole antipredator defenses in seven Rana temporaria populations collected along a 1500-km latitudinal gradient across Sweden, along which previous studies have found increasing tadpole growth and development rates. In a laboratory common garden experiment, we measured behavioral and morphological defenses by raising tadpoles in the presence and absence of a predator (Aeshna dragonfly larva) in two temperature treatments. We also estimated tadpole survival in the presence of free-ranging predators and compared predator densities between R. temporaria breeding ponds situated at low and high latitudes. Activity and foraging were generally positively correlated with latitude in the common garden experiment. While all populations responded to predator presence by decreasing activity and foraging, high-latitude populations maintained higher activity levels in the presence of the predator. All populations exhibited defensive morphology in body and tail shape. However, whereas tail depth tended to increase with latitude in the presence of predator, it did not change with latitude in the absence of the predator. Predator presence generally increased larval period and decreased growth rate. In the southern populations, predator presence tended to have a negative effect on metamorphic size, whereas in the northern populations predators had little or a positive effect on size. Latitude of origin had a strong effect on survival in the presence of a free-ranging predator, with high-latitude tadpoles experiencing higher mortality than those from the low latitudes. In the wild, predator densities were significantly lower in high-latitude than in mid-latitude breeding ponds. Although the higher activity level in the northern populations seems to confer a significant survival disadvantage under predation risk, it is probably needed to maintain the high growth and development rates. However, the occurrence of R. temporaria at high latitudes may be facilitated by the lower predator densities in the north.     

Prey–predator dynamics with predator switching regulated by a catabolic repression control mode

Since many predators can live under certain circumstances as saprophytes or consume more than one prey, and different enzymes are generally required for each prey or nutrient digestion, the predator must be sufficiently adaptive for effective utilization of the prey mass. Control modes as induction and repression, however, act at the level of genes and cause changes in the biosynthesis rate of these enzymes. In this work, an extension of the catabolic repression control mode from the level of genes to the level of the behavior of the predator is proposed, in order to model the balanced attack of the predator on the prey. It is demonstrated that, when the prey population has the competitive advantage over the predator (in using the common substrate), the catabolic repression mechanism favors the prey population, which dominates over the predator even at low specific dilution rate values, whereas, the stable steady or periodic coexistence state is not favored. When the predator has the competitive advantage at low substrate concentrations and the prey at high substrate concentrations, the introduction of the catabolic repression mechanism in the model again favors the stable steady state of the prey, while the coexistence region is dramatically reduced. Conversely, when the prey population has the competitive advantage at low and the predator at high substrate concentrations, dominance of prey and coexistence steady state could be favored by the catabolic repression mechanism. It is concluded that the catabolic repression control favors dominance of the prey population and, under certain circumstances, coexistence of both prey and predator populations.     

Riding the storm: the response of Plantago lanceolata to simulated tidal flooding

Supra-tidal plant communities fulfil a vital role in coastal protection and conservation but despite an increased likelihood of salt-water inundation from storm surges, we understand remarkably little on how salinity affects habitats like coastal grasslands or their component species. We quantified the survival and growth of a common coastal grassland plant, Plantago lanceolata when exposed to short-duration (1-, 2-, 4-, or 8-h) immersion in sea water. We also calculated root:shoot ratios (R:SR) and specific leaf area (SLA) to examine how salinity stress affects above- and below-ground resource allocation patterns and likely interactions with other trophic levels. Immersion in sea water reduced Plantago survival particularly at longer durations of 4- and 8 h, and for surviving plants, growth was also much reduced. Contrary to studies with crop plants however, we found reduced allocation to root biomass (R:SR) and increased SLA. The fact that Plantago displayed opposite ecophysiological responses to those consistently reported in the literature highlights that the response of coastal grassland plants to storm surge events cannot be assumed from conventional wisdom. In order to better protect and conserve these internationally important ecosystems from the effects of anthropogenically induced sea-level rise, a systematic exploration of the effects of sea water flooding on coastal grasslands is required.     

Nephtys hombergii, a free-living predator in marine sediments: energy production under environmental stress

Nephtys hombergii is a free-living, burrowing predator in marine sediments. The worm is, therefore, exposed to various environmental conditions which tube-dwelling polychaetes of the same habitat most likely do not encounter. The worms have to survive periods of severe hypoxia and sulphide exposure, while at the same time, they have to maintain agility in order to feed on other invertebrates. N. hombergii is adapted to these conditions by utilising several strategies. The species has a remarkably high content of phosphagen (phosphoglycocyamine), which is the primary energy source during periods of environmental stress. With increasing hypoxia, energy is also provided via anaerobic glycolysis (pO₂<7 kPa), with strombine as the main end-product. Energy production via the succinate pathway becomes important only under severe hypoxia (<2 kPa), suggesting a biphasic response to low oxygen conditions which probably is related to the worm's mode of life. The presence of sulphide resulted in a higher anaerobic energy flux and a more pronounced energy production via glycolysis than in anoxia alone. Nevertheless, after sulphide exposure under anaerobic conditions of <24 h, N. hombergii is able to recover completely. Although N. hombergii appears to be well adapted to a habitat with short-term fluctuations in oxygen and appearance of hydrogen sulphide, its high energy demand as a predator renders it likely to limit its survival in an environment with longer lasting anoxia and concomitant sulphide exposure. Received: 28 May 1997 / Accepted: 21 June 1997     

Connell and Slatyer's models of succession in the biodiversity era

Understanding how species interactions drive succession is a key issue in ecology. In this study we show the utility of combining the concepts and methodologies developed within the biodiversity-ecosystem functioning research program with J. H. Connell and R. O. Slatyer's classic framework to understand succession in assemblages where multiple interactions between early and late colonists may include both inhibitory and facilitative effects. We assessed the net effect of multiple species interactions on successional changes by manipulating the richness, composition, and abundance of early colonists in a low-shore assemblage of algae and invertebrates of the northwestern Mediterranean. Results revealed how concomitant changes in species richness and abundance can strongly alter the net effect of inhibitory vs. facilitative interactions on succession. Increasing richness of early colonists inhibited succession, but only under high levels of initial abundance, probably reflecting the formation of a highly intricate matrix that prevented further colonization. In contrast, increasing initial abundance of early colonists tended to facilitate succession under low richness. Thus, changes in abundance of early colonists mediated the effects of richness on succession.     

Ecomorphology and disease: cryptic effects of parasitism on host habitat use, thermoregulation, and predator avoidance

Parasites can cause dramatic changes in the phenotypes of their hosts, sometimes leading to a higher probability of predation and parasite transmission. Because an organism's morphology directly affects its locomotion, even subtle changes in key morphological traits may affect survival and behavior. However, despite the ubiquity of parasites in natural communities, few studies have incorporated parasites into ecomorphological research. Here, we evaluated the effects of parasite-induced changes in host phenotype on the habitat use, thermal biology, and simulated predator-escape ability of Pacific chorus frogs (Pseudacris regilla) in natural environments. Frogs with parasite-induced limb malformations were more likely to use ground microhabitats relative to vertical refugia and selected less-angled perches closer to the ground in comparison with normal frogs. Although both groups had similar levels of infection, malformed frogs used warmer microhabitats, which resulted in higher body temperatures. Likely as a result of their morphological abnormalities, malformed frogs allowed a simulated predator to approach closer before escaping and escaped shorter distances relative to normal frogs. These data indicate that parasite-induced morphological changes can significantly alter host behavior and habitat use, highlighting the importance of incorporating the ubiquitous, albeit cryptic, role of parasites into ecomorphological research.     

Habitat Change and Plant Demography: Assessing the Extinction Risk of a Formerly Common Grassland Perennial

Abstract:  An important aim of conservation biology is to understand how habitat change affects the dynamics and extinction risk of populations. We used matrix models to analyze the effect of habitat degradation on the demography of the declining perennial plant Trifolium montanum in 9 calcareous grasslands in Germany over 4 years and experimentally tested the effect of grassland management. Finite population growth rates (λ) decreased with light competition, measured as leaf-area index above T. montanum plants. At unmanaged sites λ was <1 due to lower recruitment and lower survival and flowering probability of large plants. Nevertheless, in stochastic simulations, extinction of unmanaged populations of 100 flowering plants was delayed for several decades. Clipping as a management technique rapidly increased population growth because of higher survival and flowering probability of large plants in managed than in unmanaged plots. Transition-matrix simulations from these plots indicated grazing or mowing every second year would be sufficient to ensure a growth rate ≥1 if conditions stayed the same. At frequently grazed sites, the finite growth rate was approximately 1 in most populations of T. montanum . In stochastic simulations, the extinction risk of even relatively small grazed populations was low, but about half the extant populations of T. montanum in central Germany are smaller than would be sufficient for a probability of survival of >95% over 100 years. We conclude that habitat change after cessation of management strongly reduces recruitment and survival of established individuals of this perennial plant. Nevertheless, our results suggest extinction processes may take a long time in perennial plants, resulting in an extinction debt. Even if management is frequent, many remnant populations of T. montanum may be at risk because of their small size, but even a slight increase in size could considerably reduce their extinction risk.     

Consequences of stage-structured predators: cannibalism, behavioral effects, and trophic cascades

Cannibalistic and asymmetrical behavioral interactions between stages are common within stage-structured predator populations. Such direct interactions between predator stages can result in density- and trait-mediated indirect interactions between a predator and its prey. A set of structured predator-prey models is used to explore how such indirect interactions affect the dynamics and structure of communities. Analyses of the separate and combined effects of stage-structured cannibalism and behavior-mediated avoidance of cannibals under different ecological scenarios show that both cannibalism and behavioral avoidance of cannibalism can result in short- and long-term positive indirect connections between predator stages and the prey, including "apparent mutualism." These positive interactions alter the strength of trophic cascades such that the system's dynamics are determined by the interaction between bottom-up and top-down effects. Contrary to the expectation of simpler models, enrichment increases both predator and prey abundance in systems with cannibalism or behavioral avoidance of cannibalism. The effect of behavioral avoidance of cannibalism, however, depends on how strongly it affects the maturation rate of the predator. Behavioral interactions between predator stages reduce the short-term positive effect of cannibalism on the prey density, but can enhance its positive long-term effects. Both interaction types reduce the destabilizing effect of enrichment. These results suggest that inconsistencies between data and simple models can be resolved by accounting for stage-structured interactions within and among species.     

Interactions of an Insecticide with Larval Density and Predation in Experimental Amphibian Communities

Abstract: This study examines the effects of the short-lived insecticide carbaryl, a neurotoxin, on amphibian communities experiencing natural stresses of competition and predation. Tadpoles of three species (  Woodhouse's toad [ Bufo woodhousii ], gray treefrog [ Hyla versicolor ], and green frog [ Rana clamitans ]), representing a commonly encountered assemblage in Missouri, were reared in outdoor polyethylene pond mesocosms. We determined the effects of initial tadpole density ( low or high), predation (newts [  Notophthalmus viridescens ] absent or present), chemical exposure (0, 3.5, or 7.0 mg /L carbaryl), and their interactions on body mass, larval period, and survival to metamorphosis. Green frogs in high-density ponds did not reach metamorphosis, but metamorphs in low-density ponds and tadpoles in high-density ponds were not significantly affected by treatments or their interaction. Carbaryl reduced survival to metamorphosis in toads and treefrogs and increased mass at metamorphosis in treefrogs. Effects of carbaryl varied with predator environment and initial larval density. Interactions of carbaryl with predator and with density may result in an indirect effect of carbaryl causing increased food resources through the elimination of zooplankton populations that may compete for similar resources. Our results indicate that differences in biotic conditions influenced the potency of carbaryl and that even low concentrations induce changes that may alter community dynamics in ways not predicted from single-factor, laboratory-based studies.     

Counterintuitive density-dependent growth in a long-lived vertebrate after removal of nest predators

Examining the phenotypic and genetic underpinnings of life-history variation in long-lived organisms is central to the study of life-history evolution. Juvenile growth and survival are often density dependent in reptiles, and theory predicts the evolution of slow growth in response to low resources (resource-limiting hypothesis), such as under densely populated conditions. However, rapid growth is predicted when exceeding some critical body size reduces the risk of mortality (mortality hypothesis). Here we present results of paired, large-scale, five-year field experiments to identify causes of variation in individual growth and survival rates of an Australian turtle (Emydura macquarii) prior to maturity. To distinguish between these competing hypotheses, we reduced nest predators in two populations and retained a control population to create variation in juvenile density by altering recruitment levels. We also conducted a complementary split-clutch field-transplant experiment to explore the impact of incubation temperature (25 degrees or 30 degrees C), nest predator level (low or high), and clutch size on juvenile growth and survival. Juveniles in high-recruitment (predator removal) populations were not resource limited, growing more rapidly than young turtles in the control populations. Our experiments also revealed a remarkably long-term impact of the thermal conditions experienced during embryonic development on growth of turtles prior to maturity. Moreover, this thermal effect was manifested in turtles approaching maturity, rather than in turtles closer to hatching, and was dependent on population density in the post-hatching rearing environment. This apparent phenotypic plasticity in growth complements our observation of a strong, positive genetic correlation between individual body size in the experimental and control populations over the first five years of life (rG - +0.77). Thus, these Australian pleurodiran turtles have the impressive capacity to acclimate plastically to major demographic perturbations and enjoy the longer-term potential to evolve adaptively to maintain viability.     

Induction of anti-predator responses in the green-lipped mussel Perna viridis under hypoxia

Hypoxia due to the over enrichment of waters by nutrients is becoming a global problem. In mussels, enhanced byssus thread production is an important adaptation to the presence of crustacean predators and to energetic hydrodynamic regimes. Thread production is an energy-consuming process, so this study used the green mussel Perna viridis (L.) to examine the response to predator exposure combined with hypoxia. Hypoxia is common in sheltered bays in Hong Kong, and the mussels were collected in one such bay, Lok Wo Sha (latitude/longitude: 22o18′ N/114o10′ E) in January, 2009. The predator used in the experiments was the swimming crab Thalamita danae. Oxygen concentrations used in the 48-h experiments ranged from hypoxic to normoxic (1.5 ± 0.3 mg l⁻¹, 3.0 ± 0.3 mg l⁻¹ and 6.0 ± 0.3 mg l⁻¹). Fewer byssus threads which were also shorter and thinner were produced at reduced oxygen levels, no matter if the predator was present or not; the frequency the mussels shed stalks was also lower. Mussels exposed to the predator, however, have enhanced byssus thread production at all oxygen levels when compared with the control. This has highlighted the significance of anti-predator responses for the survival of individuals even under a stressful environment in which energy supply is limited by aerobic metabolism. Interactive effects between oxygen level and predator exposure were observed for the byssus thread production (frequency of shed stalks, mean byssus thread length, cumulative byssus thread volume), with values obtained at 1.5 and 3.0 mg O₂ l⁻¹ being statistically indistinguishable for the control group without predator but not for the predator group. The lack of differences in the byssus thread production at lower oxygen levels in the absence of predator may indicate the minimum amount of byssus that is required for settlement on a substrate.