The surf zone: a semi-permeable barrier to onshore recruitment of invertebrate larvae?

The supply of larvae to the shore is important for population replenishment and intertidal community dynamics but its variability at most scales is not well understood. We tested the relationship between nearshore mussel larval abundance and intertidal settlement rates over several years at multiple spatiotemporal scales in Oregon and New Zealand. Abundance of competent larvae nearshore and intertidal recruitment rates were simultaneously quantified using collectors mounted at different depths on moorings 50-1100 m from shore, and at adjacent rocky intertidal sites. Total mussel larval abundance and oceanographic conditions were also measured in some locations. At all scales, abundance of nearshore mussel larvae was unrelated to intertidal recruitment rates. In the intertidal, patterns of mussel recruitment were persistent in space, with sites of consistently high or low recruitment. In contrast, nearshore competent larval abundance showed generally similar abundances among sites except for a high, and spatially-inconsistent, variability in Oregon during 1998 only. On moorings, recruitment tended to be greater on midwater collectors than shallower or deeper. Finally, on moorings larval abundance in traps and recruitment on collectors was unrelated. These results suggest that (1) among sites, the size of the nearshore larval pool is relatively uniform while onshore recruitment varies and is unrelated to larval abundance, (2) temporal variability in nearshore larval availability is not strongly expressed onshore, (3) vertical stratification of competent larvae nearshore is strong and may influence transport and recruitment, and (4) within-coast variability in onshore recruitment is strongly driven by processes occurring locally within the surf zone that need to be studied to understand coastal recruitment dynamics.     

Microbial communities across nearshore to offshore coastal transects are primarily shaped by distance and temperature

Recent studies have focused on linking marine microbial communities with environmental factors, yet, relatively little is known about the drivers of microbial community patterns across the complex gradients from the nearshore to open ocean. Here, we examine microbial dynamics in 15 five-station transects beginning at the estuarine Piver's Island Coastal Observatory (PICO) time-series site and continuing 87 km across the continental shelf to the oligotrophic waters of the Sargasso Sea. 16S rRNA gene libraries reveal strong clustering by sampling site with distinct nearshore, continental shelf and offshore oceanic communities. Water temperature and distance from shore (which serves as a proxy for gradients in factors such as productivity, terrestrial input and nutrients) both most influence community composition. However, at the phylotype level, modelling shows the distribution of some taxa is linked to temperature, others to distance from shore and some by both factors, highlighting that taxa with distinct environmental preferences underlie apparent clustering by station. Thus, continental margins contain microbial communities that are distinct from those of either the nearshore or the offshore environments and contain mixtures of phylotypes with nearshore or offshore preferences rather than those unique to the shelf environment.     

Does life history predict past and current connectivity for rocky intertidal invertebrates across a marine biogeographic barrier?

The southeast Australian coast potentially includes a complex biogeographic barrier, largely lacking exposed rocky shore that may limit the dispersal of rocky intertidal taxa and contribute to the maintenance of two biogeographic regions. Surprisingly, within the 300-km barrier region, several species considered exposed rocky shore specialists occurred within sheltered sites. We analysed COI sequence variation for 10 rocky intertidal invertebrate species, with a range of life histories, to test the hypotheses that larval type and habitat specificity are strong predictors of gene flow between biogeographic regions. Our data revealed that the southeast corner of Australia includes a strong barrier to gene flow for six of eight species with planktonic larvae, and a coalescence analysis of sequence differentiation (IM model) suggests that a barrier has existed since the Pleistocene. In contrast, two direct developers were not affected by the barrier. Our comparative approach and data from earlier studies (reviewed here) do not support the hypothesis that larval type predicts gene flow across this barrier, instead we found that the ability to utilize sheltered habitat provides a clearer explanation of the phylogeographic break. Indeed, the species that displayed little or no evidence of a phylogeographic break across the barrier each displayed unexpectedly relaxed habitat specificity.     

A metabolic approach to study algal–bacterial interactions in changing environments

Increasing evidence exists that bacterial communities interact with and shape the biology of algae and that their evolutionary histories are connected. Despite these findings, physiological studies were and still are generally carried out with axenic or at least antibiotic‐treated cultures. Here, we argue that considering interactions between algae and associated bacteria is key to understanding their biology and evolution. To deal with the complexity of the resulting ‘holobiont’ system, a metabolism‐centred approach that uses combined metabolic models for algae and associated bacteria is proposed. We believe that these models will be valuable tools both to study algal–bacterial interactions and to elucidate processes important for the acclimation of the holobiont to environmental changes.     

Nearshore benthic community structure at the Bounty and Antipodes Islands, Subantarctic New Zealand

Management decisions aimed at protecting biodiversity ideally should be based on biological information, but for remote and logistically difficult sites, such as are found at high latitudes, these data may be lacking. During March 2009, surveys were completed of the nearshore rocky reef communities around the Bounty and Antipodes Islands, in New Zealand??s subantarctic region. Previously considered to support the same habitat types (which used physical variables as surrogates for biological communities), analysis of photoquadrats taken at both island groups showed that the rocky reef communities were significantly different, both in terms of their species composition and in terms of their potential ecological function. While Antipodes Island supported fairly typical subantarctic shallow subtidal marine communities dominated by nongeniculate coralline algae, the rocky reefs at the Bounty Islands were dominated by filter- and suspension-feeding invertebrates, in particular encrusting sponges, barnacles and mussels. The mobile invertebrate fauna associated with these communities were also significantly different between the two island groups. Contrasting geology, oceanographic conditions and nutrient input from seabird and pinniped colonies may all contribute to the observed nearshore community structures at the Bounty and Antipodes Islands. Our research provides a baseline for assessing change in the subantarctic region and highlights the importance of using biological community data where available, to inform conservation management decisions.     

Ciguatera Fish Poisoning in La Habana,Cuba: A Study of Local Social–Ecological Resilience

Following the collapse of the Cuban economy in the early 1990s, epidemiologists in the Cuban Ministry of Health noticed dramatic increases in reported outbreaks of ciguatera fish poisoning (CFP) in some coastal communities. This article summarizes the results of a comparative case study which applied an ecosystem approach to human health to investigate this issue. Situated learning and complexity theories were used to interpret the results of the investigation. CFP outbreaks are influenced by a complex set of interactions between ecological and socioeconomic processes. This study found that the level of organization of the local sports fishing community and the degree of degradation of the local nearshore marine ecosystem appear to be key factors influencing the diverging levels of CFP outbreaks recorded in the 1990s in the communities studied.     

Intertidal zonation and latitudinal gradients on macroalgal assemblages: Species,functional groups and thallus morphology approaches

Macroalgae are unavoidable biological elements when monitoring and assessing costal environments. However, these tasks can be difficult to address because macroalgae a) present a high natural variability across a range of spatial and temporal scales, b) they imply a high sampling and laboratory processing effort and good taxonomical expertise (as they are a very diverse group of species), and c) there is insufficient knowledge about their structural and functional characteristics. This work addressed how the vertical (intertidal zonation) and horizontal (latitudinal gradient) variability of macroalgae assemblages are structured across continental Portugal, as well as how some surrogates for species-level biodiversity measures (namely functional groups and thallus morphology approaches) respond to such large-scale variability. Particularly, it was tested if intertidal zonation patterns are higher than fine-scale horizontal variation, and however, if vertical variation decreases along broad-scale horizontal variation. To do so, cover per species was taken (using a photographical and GIS methodological approach) from five sites located along the shoreline and along respective upper- mid- and lower-intertidal zones. The work findings include that both intertidal and latitudinal gradients impose deep structural changes on assemblages patterns. That is, broad-scale processes along Portuguese latitudes act as strongly as vertical stress gradients on assemblages patterns. Functional groups and thallus morphology approaches were useful to generalize the latitudinal assemblages patterns, where some groups emerge at the expense of others, and may improve biodiversity understanding and ecological synthesis. Because these surrogates decrease taxonomical expertise needs and can provide insight into the functional structure of macroalgal communities, their patterns founded may be particularly useful as reference data for further monitoring, so that shifts in such patterns might represent early warning surrogate approaches to detect environmental impact changes. Ultimately, to generate broader databases on rocky shore assemblages diversity (from species-level to functional groups and thallus morphologies approaches) can be useful for large-scale comparisons and for establishing ecological reference conditions, including for monitoring programs and environmental impact studies.     

Habitat and scale shape the demographic fate of the keystone sea urchin Paracentrotus lividus in Mediterranean macrophyte communities

Demographic processes exert different degrees of control as individuals grow, and in species that span several habitats and spatial scales, this can influence our ability to predict their population at a particular life-history stage given the previous life stage. In particular, when keystone species are involved, this relative coupling between demographic stages can have significant implications for the functioning of ecosystems. We examined benthic and pelagic abundances of the sea urchin Paracentrotus lividus in order to: 1) understand the main life-history bottlenecks by observing the degree of coupling between demographic stages; and 2) explore the processes driving these linkages. P. lividus is the dominant invertebrate herbivore in the Mediterranean Sea, and has been repeatedly observed to overgraze shallow beds of the seagrass Posidonia oceanica and rocky macroalgal communities. We used a hierarchical sampling design at different spatial scales (100 s, 10 s and <1 km) and habitats (seagrass and rocky macroalgae) to describe the spatial patterns in the abundance of different demographic stages (larvae, settlers, recruits and adults). Our results indicate that large-scale factors (potentially currents, nutrients, temperature, etc.) determine larval availability and settlement in the pelagic stages of urchin life history. In rocky macroalgal habitats, benthic processes (like predation) acting at large or medium scales drive adult abundances. In contrast, adult numbers in seagrass meadows are most likely influenced by factors like local migration (from adjoining rocky habitats) functioning at much smaller scales. The complexity of spatial and habitat-dependent processes shaping urchin populations demands a multiplicity of approaches when addressing habitat conservation actions, yet such actions are currently mostly aimed at managing predation processes and fish numbers. We argue that a more holistic ecosystem management also needs to incorporate the landscape and habitat-quality level processes (eutrophication, fragmentation, etc.) that together regulate the populations of this keystone herbivore.     

Larval history influences post-metamorphic condition in a coral-reef fish

Upon settlement, many fishes undergo an energetically costly metamorphic period that requires substantial nutritional reserves. Larval growth and the accumulation of lipids prior to metamorphosis are likely to influence growth and survival following this critical period. On the Caribbean island of St. Croix, I investigated relationships between larval growth, early life-history characteristics, and post-metamorphic lipid content in the bluehead wrasse Thalassoma bifasciatum. Lipid reserves remaining after metamorphosis were positively related (r2 = 0.62) to the width of the metamorphic band; thus, this otolith-derived trait may be used to estimate the condition at emergence of survivors collected at some later time. In contrast, pelagic larval duration, average larval growth, and otolith size at settlement were negatively related to post-metamorphic lipid content. Interestingly, the trend for slower growth among fish in good condition was not consistent over the entire pelagic larval duration. Analyses of daily larval growth histories indicated that fish with high lipid reserves grew rapidly in the last week prior to settlement, but relatively slowly during the early phases of larval life; those emerging with low lipid concentrations, however, displayed strikingly opposite patterns. These contrasting patterns of growth and energy storage were consistent at two sites and over three recruitment events. Otolith chemistry data suggested that differences in growth histories and body condition were consistent with the hypothesis of larval development in distinct oceanic environments (characterized by Pb concentration); but, within a water mass, differences reflected life-history trade-offs between growth and energy storage. The results have implications for understanding the processes driving juvenile survival, which may be condition dependent.     

Latitudinal trends in reproduction,recruitment and population characteristics of some rocky littoral molluscs and cirripedes

This paper reviews progress in the COST 647 rocky littoral programme involving three patellids, two trochids and two cirripedes on European Atlantic coasts. Northern geographical limits are set primarily by repopulation failure, and northern populations are characterised by short, mid-summer breeding periods, high frequency of failed or poor recruitments and low density of large, long-lived individuals. Towards the south molluscan breeding periods lengthen and become later, extending in some cases throughout much of the year but with least or no activity in mid-summer. There are probably fewer recruitment failures and thus higher densities, but individuals are of smaller maximum size and shorter life-span.The cirripedes show similar latitudinal trends in recruitment timing but Semibalanus is restricted to a single annual brood throughout its range.The two species reaching their southern limits show progressive restriction to the lowest tidal levels.Recruitment failures do not result from inadequate gonad activity. They arise in cirripedes during the planktonic phase and in the molluscs during settlement and early shore life. The temperature sensitivity of molluscan spat is primarily responsible for the north/south gradient in recruitment times from summer towards winter.Formerly University of Leeds, England     

The Response of Experimental Rocky Shore Communities to Nutrient Additions

The aim of this study was to determine whether the experimental nutrient enrichment of littoral rocky shore communities would be followed by a predicted accumulation of fast-growing opportunistic algae and a subsequent loss of perennial benthic vegetation. Inorganic nitrogen (N) and potassium (P) was added to eight concrete mesocosms inhabited by established littoral communities dominated by fucoids. The response to nutrient enrichment was followed for almost 2 1/2 years. Fast-growing opportunistic algae (periphyton and ephemeral green algae) grew significantly faster in response to nutrient enrichment, but the growth of red filamentous algae and large perennial brown algae was unaffected. However, these changes were not followed by comparable changes in the biomass and composition of the macroalgae. The biomass of opportunistic algae was stimulated only marginally by the nutrient enrichment, and perennial brown algae (fucoids) remained dominant in the mesocosm regardless of nutrient treatment level. Established rocky shore communities thus seem able to resist the effects of heavy nutrient loading. We found that the combined effects of the heavy competition for space and light imposed by canopy-forming algae, preferential grazing on opportunistic algae by herbivores, and physical disturbance, succeeded by a marked export of detached opportunistic algae, prevented the fast-growing algae from becoming dominant. However, recruitment studies showed that the opportunistic algae would become dominant when free space was available under conditions of high nutrient loading and low grazing pressure. These results show that established communities of perennial algae and associated fauna in rocky shore environments can prevent or delay the accumulation of bloom-forming opportunistic algae and that the replacement of long-lived macroalgae by opportunistic species at high nutrient loading may be a slow process. Nutrient enrichment may not, in itself, be enough to stimulate structural changes in rocky shore communities.     

Variability in the settlement of non-indigenous species in benthic communities from an oceanic island

The introduction of non-indigenous species (NIS) in new environments represents a major threat for coastal ecosystems. A good understanding of the mechanisms and magnitude of the impact of NIS colonisation on native ecosystems is becoming increasingly crucial to develop mitigation measures and prevent new invasions. In this present study, we asked if distinct coastal benthic communities from an oceanic island can have different vulnerability to NIS colonisation process. First, PVC settlement plates were deployed for 1 year on the rocky shore of two different locations of Madeira Island (North versus South coast). Then, we implemented a mesocosm experiment where recruited plate communities were maintained under different levels of NIS propagule pressure in order to assess their vulnerability to NIS colonisation process. Results showed that NIS colonisation success was not influenced by the level of propagule pressure, but however, final colonisation patterns varied depending on the origin of the communities. This variability can be attributed to major structural differences between the preponderant species of each community and therefore to the biotic substrate they offer to colonisers. This study highlights how biotic features can alter the NIS colonisation process and importantly, shows that in an urbanisation context, the nature of the resident communities facing invasions risks needs to be closely assessed.     

Population structure of an introduced species (Dreissena polymorpha) along a wave-swept disturbance gradient

The zebra mussel Dreissena polymorpha was introduced to North America during the mid-1980s, and is now a dominant member of many benthic communities in the lower Great Lakes. In this study, I explored the abundance, biomass, size structure and settlement of Dreissena inhabiting rocks along a wave-swept disturbance gradient near Middle Sister Island in western Lake Erie. Ten rocks were collected from quadrats at six sites along each of three transect lines oriented perpendicular to shore. Occurrence, abundance and biomass of Dreissena on smaller, movable rocks were positively associated with rock distance from shore (lake depth) and with rock area; rocks at nearshore sites supported little, if any, Dreissena, whereas those at offshore sites were heavily colonized. Mussel size distributions also differed in relation to shore distance. Large mussels (19 mm) were underrepresented or absent on rocks collected at nearshore sites, but were overrepresented at offshore locations (37 m). Settlement of larval mussels on settling pads was positively correlated with distance offshore and with time of exposure, though settlement was substantial even at a nearshore (10 m) location. Area-adjusted mussel dry mass increased more rapidly with distance offshore on large than on small rocks. Large rocks also required more force to displace and were significantly less likely to be disturbed when transplanted at the study site. Results from this study indicate that occurrence, abundance and size structure of Dreissena in nearshore waters of Lake Erie correspond with the frequency of habitat disturbance, though other factors including food limitation and larval supply may also contribute to these patterns. These patterns complement studies that established the significance of physical disturbance in other aquatic systems.     

Effects of an alien invertebrate species and wave action on prey selection by African black oystercatchers (Haematopus moquini)

Abstract Shorebirds foraging in the intertidal have been shown to exert a significant effect on assemblage level processes; this is particularly true of the oystercatcher–limpet–algae system. The African black oystercatcher (Haematopus moquini) is endemic to the southern African coastline, where it plays a significant role in ecosystem processes as a rocky‐shore predator, especially of mussels and limpets. This understanding was based on studies of a rocky shore environment that has since been considerably modified following invasion of an alien mussel (Mytilus galloprovincialis). This invasion has not only changed the relative proportions of different food types on the shore, but has also greatly increased overall food biomass. We tested the previous model that food selection by oystercatchers reflected prey abundance and that intake by male and female oystercatchers differed owing to bill morphology. We predicted that this difference would persist despite the changed nature of the food base. We also predicted that wave action would modify prey selection as a result of both its influence on prey behaviour and its impact on searching and handling times of the birds. Overall, both sexes consumed more limpets than expected by encounter rate alone, but contrary to prediction, the relative proportions of different prey types taken post invasion did not differ between the sexes. Dietary convergence is interpreted as a result of greatly increased food biomass on the shore, which is also reflected in increased oystercatcher densities since the invasion. Also contrary to prediction there was no evidence that waves acted as indirect modifiers of the interaction between oystercatchers and their prey. The results of this study indicate that models of trophic cascades will need to be altered in the event of a significant change in a trophic level, which then effects behavioural changes in the key predator.     

Janzen–Connell effects partially supported in reef-building corals: adult presence interacts with settler density to limit establishment

In many plant and sessile marine invertebrate (SMI) taxa, population and community dynamics are heavily influenced by processes occurring during the dispersal and establishment phases. The Janzen–Connell (J–C) hypothesis predicts increased survival of early life stages with decreasing conspecific density and increased distance from conspecific adults. Evidence of J–C effects in maintaining diversity is common in plant communities, but its importance in SMI communities remains unclear. Under controlled aquarium conditions, we examined the effect of density-dependence and adult conspecific water treatments (absent/present) on propagule settlement success and settler post-settlement survival, along with associated spatial patterns, for six broadcast-spawning, reef-building coral species from three families. We also tested if settlement success was linked to increasing propagule species diversity for three coral species from two families. We found that the probability of settlement was density independent and not influenced by adult present water treatments. Yet, adult present water treatments and settler density did have a synergistic negative effect on the probability of short-term settler survival for all species examined. Settlers also showed greater spatial aggregation as their numbers increased, but were less aggregated in adult present water treatments compared to those in adult absent water treatments. We further show evidence of significant species interactions among propagules, as settlement in single-species trials was four-fold higher compared to mixed-species trials. Our findings from controlled experimental arenas indicate that the early establishment of corals was predominantly limited by density-dependent settler–adult interactions among conspecifics and propagule–propagule interactions among heterospecifics. Thus, the proximity to established conspecific adults, settler density and species diversity of propagules are relevant drivers of local coral community diversity and structure. Based on these outcomes, we suggest that the J–C hypothesis, with demonstrated importance for plants, is partially upheld for reef corals.     

Composition dynamics of epilithic intertidal bacterial communities exposed to high copper levels

Copper has a dual role for organisms, both as micronutrient and toxic element. Copper mining activities have an enormous ecological impact because of the extraction process and the consequent release of copper-containing waste materials to the environment. In northern Chile, mainly in the Cha?aral coastal area, this phenomenon is clearly evident. The released waste material has caused a strong modification of the area, and copper enrichment of beaches and rocky shores has provoked a decrease in the richness and diversity of many species of macroorganisms. However, the effects that copper enrichment has on microbial (e.g. bacterial epilithic) communities associated with the rocky shore environment are poorly understood. Using a culture-independent molecular approach, field sampling and laboratory microcosm experiments, we determined the effects of copper enrichment on bacterial communities inhabiting the rocky shore environment. Field samples showed a strong effect of copper on the structure of the natural bacterial epilithic communities, and microcosm experiments demonstrated rapid changes in bacterial community when copper is added, and reversibility of this effect within 48?h after copper is removed.     

Population Genetics of a Trochid Gastropod Broadens Picture of Caribbean Sea Connectivity

Background

Regional genetic connectivity models are critical for successful conservation and management of marine species. Even though rocky shore invertebrates have been used as model systems to understand genetic structure in some marine environments, our understanding of connectivity in Caribbean communities is based overwhelmingly on studies of tropical fishes and corals. In this study, we investigate population connectivity and diversity of Cittarium pica, an abundant rocky shore trochid gastropod that is commercially harvested across its natural range, from the Bahamas to Venezuela.

Methodology/Principal Findings

We tested for genetic structure using DNA sequence variation at the mitochondrial COI and 16S loci, AMOVA and distance-based methods. We found substantial differentiation among Caribbean sites. Yet, genetic differentiation was associated only with larger geographic scales within the Caribbean, and the pattern of differentiation only partially matched previous assessments of Caribbean connectivity, including those based on larval dispersal from hydrodynamic models. For instance, the Bahamas, considered an independent region by previous hydrodynamic studies, showed strong association with Eastern Caribbean sites in our study. Further, Bonaire (located in the east and close to the meridional division of the Caribbean basin) seems to be isolated from other Eastern sites.

Conclusions/Significance

The significant genetic structure and observed in C. pica has some commonalities in pattern with more commonly sampled taxa, but presents features, such as the differentiation of Bonaire, that appear unique. Further, the level of differentiation, together with regional patterns of diversity, has important implications for the application of conservation and management strategies in this commercially harvested species.     

Population biology and the management of fisheries

The successful management and exploitation of fisheries requires detailed knowledge of the population dynamics and life histories of commercial species, and of the factors affecting recruitment and mortality. Much progress has been made in recent years, exemplified by the report of the 1984 Dahlem Workshop on the exploitation of marine communities, and by subsequent research discussed in this article. In some fisheries at least, management decisions can now have a firm scientific base.     

Nutrient Addition to Experimental Rocky Shore Communities Revisited: Delayed Responses, Rapid Recovery

Coastal eutrophication may alter the dominance patterns of marine macroalgae, with potential consequences for the associated fauna and the entire ecosystem. Benthic macroalgae and animals in control and nutrient-enriched mesocosms were monitored to investigate eutrophication-induced changes in rocky shore communities. During a 3-year project, nutrient addition had only minor effects on the community structure, such as increased cover and biomass of green Ulva spp. and increased abundance of certain animal species at high nutrient levels. This study is a 4-year extension of a previously reported project, with 2 extra years of effect studies (altogether 5 years) and a subsequent 2 years for recovery. During the 4th year of nutrient enrichment, the cover of Fucus vesiculosus and Fucus serratus started to decline. In the 5th year, these canopy species crashed and there was an evident take-over by green algae at high nutrient addition levels. The previously observed abundance stimulation for fauna disappeared later in the time series, probably due to the loss of the macroalgal canopy. After less than 2 years on regular seawater, the algal and animal communities had returned to within the range of normal variability. The results indicate that established rocky shore communities of perennial algae with associated fauna are able to persist for several years, even at very high nutrient levels, but that community shifts may suddenly occur if eutrophication continues. They also indicate that rocky shore communities have the ability to return rapidly to natural undisturbed conditions after the termination of nutrient enhancement.     

Predator shadows: complex life histories as generators of spatially patterned indirect interactions across ecosystems

Ecological coupling by material exchanges or dispersal between spatially distinct communities has important impacts on ecological processes, such as diversity–stability relationships, ecosystem function, and food web dynamics. One important mode of coupling between ecosystems occurs via organisms with complex life histories, which often switch between distinct ecosystems during their life cycle, and so can be channels of material exchanges between these ecosystems. Some organisms with complex life histories (e.g. frogs, dragonflies) can be abundant and effective predators during one or more life stages, and so provide conduits for strong direct and indirect interactions across ecosystem boundaries, linking the dynamics of discrete and often quite dissimilar community types. We present simple models and a case study (tailored to pond ecosystems), to explore how interactions within larval habitats can indirectly impact ecological interactions in adult habitats. Using our case study as a springboard, we propose that cohorts of predators emerging from natal habitats (e.g. ponds) cast 'predation shadows' on the surrounding adult (e.g. terrestrial) landscape. Trophic interactions within ponds, and the distribution of ponds on the landscape, can thus affect the spatial pattern in the strength of these predation shadows, creating strong spatial patterning in terrestrial trophic cascades. Our findings emphasize the importance of organisms with complex life histories as generators of strong links across ecosystem boundaries, and as potential sources of spatial variation in the strength and indirect impacts of interspecific interactions.