Comparative feeding ecology of four sympatric skate species off central California, USA

The big (Raja binoculata), California (R. inornata), longnose (R. rhina), and sandpaper (Bathyraja kincaidii), skates are commonly found on soft-bottom regions of the central California continental shelf and upper slope. The feeding ecology of this assemblage was compared to evaluate the degree of trophic separation among species, based on the results of previous species-specific diet studies. Specimens were collected from fishery independent trawl surveys conducted during September 2002–March 2003 at depths of 9–536 m. Using single and compound measures, diet composition of small (≤60 cm TL) and large (>60 cm TL) individuals were compared within continental shelf (≤200 m) and slope (>200 m) regions using traditional, multivariate, and novel techniques. Diet compositions within size classes were similar in both regions. Diet compositions between size classes generally differed, however, with fishes more important and crustaceans (especially shrimps and euphausiids) less important in the diets of large individuals. Crabs contributed substantially to skate diet compositions on the shelf, but were uncommon prey items at deeper depths, probably because of their relative scarcity in slope waters. Conversely, cephalopods were common prey items at slope depths, but were rarely ingested at shelf depths. The studied skate assemblage appears to consist primarily of generalist crustacean and fish predators that exhibit high dietary overlap at similar sizes. It is possible that resource competition among skates and groundfish species has been reduced because of considerable recent declines in the biomass of upper trophic level groundfishes. Skates may therefore play important roles in contemporary benthic food web dynamics off central California.     

Stable isotope analysis of vertebrae reveals ontogenetic changes in habitat in an endothermic pelagic shark

Ontogenetic changes in habitat are driven by shifting life-history requirements and play an important role in population dynamics. However, large portions of the life history of many pelagic species are still poorly understood or unknown. We used a novel combination of stable isotope analysis of vertebral annuli, Bayesian mixing models, isoscapes and electronic tag data to reconstruct ontogenetic patterns of habitat and resource use in a pelagic apex predator, the salmon shark (Lamna ditropis). Results identified the North Pacific Transition Zone as the major nursery area for salmon sharks and revealed an ontogenetic shift around the age of maturity from oceanic to increased use of neritic habitats. The nursery habitat may reflect trade-offs between prey availability, predation pressure and thermal constraints on juvenile endothermic sharks. The ontogenetic shift in habitat coincided with a reduction of isotopic niche, possibly reflecting specialization upon particular prey or habitats. Using tagging data to inform Bayesian isotopic mixing models revealed that adult sharks primarily use neritic habitats of Alaska yet receive a trophic subsidy from oceanic habitats. Integrating the multiple methods used here provides a powerful approach to retrospectively study the ecology and life history of migratory species throughout their ontogeny.     

Juvenile fishes in macrophyte beds: influence of food resources, habitat structure and body size

Mean juvenile fish abundance and fish frequency in a large lowland river during low discharge largely differed among the unvegetated and three morphologically contrasted macrophyte habitats. Single separate models revealed that juvenile fish distribution was largely influenced by trophic variables. With the exception of Leuciscus cephalus , which responded mainly to physical variables (depth and substratum), multiple regression models emphasized the importance of trophic variables for fish distribution. For Blicca bjoerkna , L. cephalus and Lepomis gibbosus , habitat shifts with respect to prey size were apparent; small juvenile fishes mainly responded to small zooplankton abundance, whereas large individuals were more influenced by the abundance of large zooplankton. Whatever the species, predictions from multiple regression models were always better for large individuals. Small juvenile fishes appeared to be less affected by the habitat variables measured, and exhibited more uniform spatial distribution. The relative importance of trophic resources and habitat physical structure among macrophyte types for fish-habitat relationships is discussed, and the necessity of quantifying habitat structural complexity is emphasized.     

Prey selection by age-0 walleye pollock, Theragra chalcogramma, in nearshore waters of the Gulf of Alaska

Juvenile walleye pollock, Theragra chalcogramma, is the dominant forage fish on the continental shelf of the Gulf of Alaska, yet little is known about the feeding habits of this important interval of pollock life history. The taxonomic composition and size of prey found in the stomachs of age-0 juveniles collected at three nearshore locations in the Gulf of Alaska in September 1990 were compared to the composition and size of zooplankton collected in concurrent plankton tows. The maximum length of prey consumed increased dramatically over the length range of pollock examined (58–110 mm) from approximately 7 mm to 30 mm, due mainly to the consumption of large euphausiids and chaetognaths by the bigger individuals. The maximum width of prey changed little over this size range although there was a general increase in prey width with increasing predator size. The minimum prey length and width did not change with increasing fish size. Juvenile pollock generally selected the larger prey sizes relative to what was available. Juvenile pollock showed a marked preference for adult euphausiids and decapod larvae and an avoidance of copepods and chaetognaths relative to the numbers collected in net tows. These results are discussed relative to the feeding ecology of these juvenile fishes. This revised version was published online in July 2006 with corrections to the Cover Date.     

The diet composition of immature loggerheads: Insights on trophic niche, growth rates, and fisheries interactions

For immature animals, diet quality and composition influence expression of life history traits such as growth rates and ultimately life stage duration and age to maturity. Circumglobally distributed loggerhead turtles (Caretta caretta) exhibit a multi-decade immature stage that generally occupies neritic habitats and is characterized by slow growth and an omnivorous diet. Although adult nesting populations are geographically distinct, foraging areas for immature loggerheads show a high degree of mixing of individuals that originate from multiple nesting stocks. Furthermore, despite their generalist foraging ecology, immature loggerheads have been observed to supplement their natural diets with fish from fishery discards and/or caught in fishing gear. However, whether trophic opportunism results in variation in loggerhead growth rates within or among feeding areas has not been investigated. In Core Sound, North Carolina (NC), USA, immature loggerheads demonstrate highly variable size-specific growth rates, in contrast to other studies that report discernible somatic growth functions in immature marine turtles. To determine whether inter-individual variation in growth rates at this site was due to variation in diet composition, and specifically variation in consumption of fish, we analyzed carbon and nitrogen stable isotope ratios of loggerhead blood plasma and of tissue samples of putative loggerhead prey, as well as commercially important fish species. Our results indicated that growth rates were not related to trophic levels at which individual turtles fed, but rather probably reflected inter-individual variation in overwintering or foraging behavior (i.e. nearshore vs. offshore). Furthermore, loggerhead diets were highly diverse, and comprised mainly blue crabs and/or whelks, as well as small proportions of cannonball jellies. Fish were unimportant dietary components for loggerheads. Although loggerheads in NC do not appear to feed on fish catch or discards, immature turtles showed dietary preferences for prey items that are also valuable to or are commonly taken as bycatch in commercial fisheries (e.g. blue crabs and whelks, respectively) in the region. Thus, the status of these prey items/fishery stocks as well as trends in loggerhead populations should be monitored to mitigate potential competitive interactions between fisheries activities and loggerhead turtles.     

Interactive effects of episodic hypoxia and cannibalism on juvenile blue crab mortality

We hypothesized that as the spatial extent of hypoxic bottom water increased, (1) adult blue crab predator densities would increase in shallow habitats as they avoided hypoxia, and that (2) juvenile blue crabs, which use shallow unvegetated habitat as a predation refuge from adult conspecifics, would experience increased mortality rates during crowding by cannibalistic adult blue crabs. These hypotheses were tested along a depth gradient of sandy-mud shoreline in the Neuse River Estuary (NRE), North Carolina, USA using a combination of (1) hydrographic measurements to characterize the spatial extent of hypoxia, (2) beach seines to quantify the density of adult blue crab predators in relatively shallow water as a function of 1, and (3) tethering experiments to quantify relative rates of predation on juvenile blue crabs as a function of 1 and 2. During our seven tethering experiments, the NRE study site experienced a range of DO scenarios including normoxia, chronic hypoxia, and hypoxic upwelling. No known predators of juvenile blue crabs, other than adult conspecifics, were collected in any of our shallow-water seines. During the transition from normoxia to chronic hypoxia, blue crab predator densities in shallow refuge habitats increased 4-fold, and relative mortality rates of juvenile blue crabs in shallow habitats increased exponentially with the density of adult conspecifics. Conversely, during hypoxic upwelling events, the density of adult blue crabs in shallow water declined, which may explain why the relative mortality of juvenile crabs did not increase significantly with the increasing spatial extent of hypoxia. Thus, juvenile blue crabs may be relatively safe from adult conspecifics during hypoxic upwelling events, but not during chronic hypoxia. These experimental results highlight the need to consider the effects of dynamic water quality on mobile consumers emigrating from degraded habitats when considering indirect trophic impacts beyond the immediate area of impact.     

The Mangrove Nursery Paradigm Revisited: Otolith Stable Isotopes Support Nursery-to-Reef Movements by Indo-Pacific Fishes

Mangroves and seagrass beds have long been perceived as important nurseries for many fish species. While there is growing evidence from the Western Atlantic that mangrove habitats are intricately connected to coral reefs through ontogenetic fish migrations, there is an ongoing debate of the value of these coastal ecosystems in the Indo-Pacific. The present study used natural tags, viz. otolith stable carbon and oxygen isotopes, to investigate for the first time the degree to which multiple tropical juvenile habitats subsidize coral reef fish populations in the Indo Pacific (Tanzania). Otoliths of three reef fish species (Lethrinus harak, L. lentjan and Lutjanus fulviflamma) were collected in mangrove, seagrass and coral reef habitats and analyzed for stable isotope ratios in the juvenile and adult otolith zones. δ¹³C signatures were significantly depleted in the juvenile compared to the adult zones, indicative of different habitat use through ontogeny. Maximum likelihood analysis identified that 82% of adult reef L. harak had resided in either mangrove (29%) or seagrass (53%) or reef (18%) habitats as juveniles. Of adult L. fulviflamma caught from offshore reefs, 99% had passed through mangroves habitats as juveniles. In contrast, L. lentjan adults originated predominantly from coral reefs (65–72%) as opposed to inshore vegetated habitats (28–35%). This study presents conclusive evidence for a nursery role of Indo-Pacific mangrove habitats for reef fish populations. It shows that intertidal habitats that are only temporarily available can form an important juvenile habitat for some species, and that reef fish populations are often replenished by multiple coastal habitats. Maintaining connectivity between inshore vegetated habitats and coral reefs, and conserving habitat mosaics rather than single nursery habitats, is a major priority for the sustainability of various Indo Pacific fish populations.     

Energy flow to two abundant consumers in a subtropical oyster reef food web

Oyster reefs are among the most threatened coastal habitat types, but still provide critical habitat and food resources for many estuarine species. The structure of oyster reef food webs is an important framework from which to examine the role of these reefs in supporting high densities of associated fishes. We identified major trophic pathways to two abundant consumers, gray snapper (Lutjanus griseus) and crested goby (Lophogobius cyprinoides), from a subtropical oyster reef using stomach content and stable isotope analysis. The diet of gray snapper was dominated by crabs, with shrimp and fishes also important. Juvenile gray snapper fed almost entirely on oyster reef-associated prey items, while subadults fed on both oyster reef- and mangrove-associated prey. Based on trophic guilds of the gray snapper prey, as well as relative δ¹³C values, microphytobenthos is the most likely basal resource pool supporting gray snapper production on oyster reefs. Crested goby had omnivorous diets dominated by bivalves, small crabs, detritus, and algae, and thus were able to take advantage of prey relying on production from sestonic, as well as microphytobenthos, source pools. In this way, crested goby represent a critical link of sestonic production to higher trophic levels. These results highlight major trophic pathways supporting secondary production in oyster reef habitat, thereby elucidating the feeding relationships that render oyster reef critical habitat for many ecologically and economically important fish species.     

The effects of seafloor habitat complexity on survival of juvenile fishes: Species-specific interactions with structural refuge

Marine fishes are often associated with structurally complex microhabitats that are believed to provide a refuge from predation. However, the effects of habitat complexity on predator foraging success can be strongly modified by predator and prey behaviors. We conducted a series of laboratory experiments to evaluate the effects of sea floor habitat complexity on juvenile fish survivorship using multiple predator (striped searobin and summer flounder) and prey (winter flounder, scup, and black sea bass) species to identify potentially important species-habitat interactions. Three habitats of varying complexity (bare sand, shell, and sponge) common to coastal marine environments were simulated in large aquaria (2.4 m diameter, 2400 L volume). Prey survivorship increased significantly with greater habitat complexity for each species combination tested. However, examination of multiple prey and predator species across habitats revealed important effects of predator × habitat and prey × habitat interactions on prey survival, which appeared to be related to species-specific predator and prey behavior in complex habitats. Significant species × habitat interactions imply that the impact of reduced seafloor habitat complexity may be more severe for some species than others. Our results indicate that the general effects of seafloor habitat complexity on juvenile fish survivorship may be broadly applicable, but that the interaction of particular habitats with search tactics of predators as well as habitat affinities and avoidance responses of prey can produce differences among species that contribute to variable mortality.     

Preference Alters Consumptive Effects of Predators: Top-Down Effects of a Native Crab on a System of Native and Introduced Prey

Top-down effects of predators in systems depend on the rate at which predators consume prey, and on predator preferences among available prey. In invaded communities, these parameters might be difficult to predict because ecological relationships are typically evolutionarily novel. We examined feeding rates and preferences of a crab native to the Pacific Northwest, Cancer productus, among four prey items: two invasive species of oyster drill (the marine whelks Urosalpinx cinerea and Ocenebra inornata) and two species of oyster (Crassostrea gigas and Ostrea lurida) that are also consumed by U. cinerea and O. inornata. This system is also characterized by intraguild predation because crabs are predators of drills and compete with them for prey (oysters). When only the oysters were offered, crabs did not express a preference and consumed approximately 9 juvenile oysters crab⁻¹ day⁻¹. We then tested whether crabs preferred adult drills of either U. cinerea or O. inornata, or juvenile oysters (C. gigas). While crabs consumed drills and oysters at approximately the same rate when only one type of prey was offered, they expressed a strong preference for juvenile oysters over drills when they were allowed to choose among the three prey items. This preference for oysters might negate the positive indirect effects that crabs have on oysters by crabs consuming drills (trophic cascade) because crabs have a large negative direct effect on oysters when crabs, oysters, and drills co-occur.     

Habitat-based submersible assessment of macro-invertebrate and groundfish assemblages at Heceta Bank, Oregon, from 1988 to 1990

Exploitation of groundfish off the U.S. Pacific coast reached maximum levels during the 1990s, resulting in severe declines in at least nine species of groundfish. From 1988 to 1990, we used the 2-man submersible Delta to make 42 dives and run replicate visual belt transects at six stations ranging from 67-360 m in depth at Heceta Bank on the outer continental shelf of Oregon. We identified four major habitats and associated benthic macroinvertebrate and groundfish assemblages: (1) shallow rock ridges and large boulders (< 100 m deep) dominated by basket stars, juvenile rockfishes, yelloweye rockfish, and lingcod; (2) mid-depth small boulder-cobbles (100-150 m) dominated by crinoids, brittle stars, rosethorn, pygmy/Puget sound, and canary rockfishes; (3) deep cobble (150-200 m) dominated by crinoids, brittle stars and various small rockfish species, and (4) deep mud slope (> 200 m) dominated by fragile urchins, sea cucumbers, shortspine thornyhead, and flatfishes. Although substantial interannual variation in groundfish abundance among seafloor types was evident in the 12 most abundant and/or commercially important fish taxa sampled, high variance resulted in statistically significant differences among years only in juvenile rockfishes. These data provide a baseline for future comparisons exploring long-term change this continental-shelf ecosystem.     

The relative importance of habitat-specific settlement, predation and juvenile dispersal for distribution and abundance of young juvenile shore crabs Carcinus maenas L.

Young juveniles of many motile benthic species are concentrated in structurally complex habitats, but the proximate causes of this distribution are usually not clear. In the present study, I assessed three potentially important processes affecting distribution and abundance of early benthic stages in the shore crab (Carcinus maenas): (1) selection of habitat by megalopae (postlarvae); (2) habitat-specific predation; and (3) post-settlement movements by juveniles. These processes were assessed concurrently over 3-9 days at two spatial scales: at the scale of square meters using cage techniques within nursery areas, and at the scale of hectares using isolated populations of juvenile shore crabs in small nursery areas as mesocosms. The results were compared to habitat-specific distribution in the field.Shore crab megalopae and first instar juveniles (settlers) were distributed non-randomly among micro-habitats in the assessed nursery areas, with great densities in both mussel beds, eelgrass and filamentous algal patches (on average 114-232 settlers m⁻²), and significantly smaller densities on open sand habitats at all times (on average 4 settlers m⁻²). The same habitat-specific settlement pattern was found in cages where predators were excluded, suggesting that active habitat selection at settlement was responsible for the initial distribution. Older juveniles (second to ninth instar crabs) were also sparse on sand, but in contrast to settlers, were concentrated in mussel beds, which showed significantly greater densities than eelgrass and algal habitats. The cage experiment demonstrated a dynamic distribution of juvenile crabs. Young juveniles constantly migrated over open sand habitats (20 m or further) and colonized the experimental plots in a habitat-specific pattern that reflected the distribution in the field. This pattern was also found for very small crabs colonizing predator-exclusion cages, suggesting that selection of habitat by migrating juveniles caused the ontogenetic change in habitat use. Although post-settlement movements were great within nursery areas, juvenile dispersal at a regional scale appeared to be small, and the recruitment of juvenile shore crabs to the shallow bays occurred mainly through pelagic megalopae.Conservative estimates at the scale of whole nursery areas, based on migration trap data and field samples, indicated great mortality of settlers and early benthic stages of shore crabs. Results from the cage experiment suggest that predation by crabs and shrimp were responsible for the high settlement mortality. Both enclosed cannibalistic juvenile crabs and local predators on uncaged habitat plots caused significant losses of settlers in all habitats (on average 22% and 64% 3 day⁻¹, respectively). The effect of predators was highly variable between trials, but differed little between habitat types, and predation had no detectable proximate effect on juvenile distribution, despite the great losses. Small settlement densities on sand habitats in combination with a refuge at low prey numbers, and an aggregation of cannibalistic juvenile crabs in nursery habitats appear to decrease the effect of habitat-specific predation rates on the distribution of juvenile shore crabs. This study demonstrates that active habitat selection at settlement followed by a dynamic redistribution of young juveniles can be the proximate processes responsible for habitat-specific distribution of epibenthic juveniles, and indicate that predation represents a major evolutionary process reinforcing this behavior.     

Differences in predator composition alter the direction of structure‐mediated predation risk in macrophyte communities

Structural complexity strongly influences the outcome of predator–prey interactions in benthic marine communities affecting both prey concealment and predator hunting efficacy. How habitat structure interacts with species‐specific differences in predatory style and antipredatory strategies may therefore be critical in determining higher trophic functions. We examined the role of structural complexity in mediating predator–prey interactions across several macrophyte habitats along a gradient of structural complexity in three different bioregions: western Mediterranean Sea (WMS), eastern Indian Ocean (EIO) and northern Gulf of Mexico (NGM). Using sea urchins as model prey, we measured survival rates of small (juveniles) and medium (young adults) size classes in different habitat zones: within the macrophyte habitat, along the edge and in bare sandy spaces. At each site we also measured structural variables and predator abundance. Generalised linear models identified biomass and predatory fish abundance as the main determinants of predation intensity but the efficiency of predation was also influenced by urchin size class. Interestingly though, the direction of structure‐mediated effects on predation risk was markedly different between habitats and bioregions. In WMS and NGM, where predation by roving fish was relatively high, structure served as a critical prey refuge, particularly for juvenile urchins. In contrast, in EIO, where roving fish predation was low, predation was generally higher inside structurally complex environments where sea stars were responsible for much of the predation. Larger prey were generally less affected by predation in all habitats, probably due to the absence of large predators. Overall, our results indicate that, while the structural complexity of habitats is critical in mediating predator–prey interactions, the direction of this mediation is strongly influenced by differences in predator composition. Whether the regional pool of predators is dominated by visual roving species or chemotactic benthic predators may determine if structure dampens or enhances the influence of top–down control in marine macrophyte communities.     

Seasonal variation in the trophic structure of a spatial prey subsidy linking aquatic and terrestrial food webs: adult aquatic insects

Research over the past decade has established spatial resource subsidies as important determinants of food web dynamics. However, most empirical studies have considered the role of subsidies only in terms of magnitude, ignoring an important property of subsidies that may affect their impact in recipient food webs: the trophic structure of the subsidy relative to in situ resources. This may be especially important when subsidies are composed of organisms, as opposed to nutrient subsidies, because the trophic position of subsidy organisms may differ from in situ prey. I explored the relative magnitude and trophic structure of a cross-habitat prey subsidy, adult aquatic insects, in terrestrial habitats along three streams in the south–central United States. Overall, adult aquatic insects contributed more than one-third of potential insect prey abundance and biomass to the terrestrial habitat. This contribution peaked along a permanent spring stream, reaching as high as 94% of abundance and 86% of biomass in winter. Trophic structure of adult aquatic and terrestrial insects differed. Nearly all adult aquatic insects were non-consumers as adults, whereas all but one taxon of terrestrial insects were consumers. Such a difference created a strong relationship between the relative contribution of the prey subsidy and the trophic structure of the prey assemblage: as the proportion of adult aquatic insects increased, the proportion of consumers in the prey assemblage declined. Specific effects varied seasonally and with distance from the stream as the taxonomic composition of the subsidy changed, but general patterns were consistent. These findings show that adult aquatic insect subsidies to riparian food webs not only elevate prey availability, but also alter the trophic structure of the entire winged insect prey assemblage.     

Trophic relationships of juvenile blue crabs (Callinectes sapidus) in estuarine habitats

Salt marshes and shallow-water macroalgal beds are known to provide nursery habitat for many species of fish and invertebrates. The role of these habitats as refuge from predation is well established, but the degree to which indigenous primary production within the nursery provides food for growth and development of estuarine species remains unresolved. In this study, we tested the hypothesis that juvenile blue crabs depend on indigenous primary production, directly or indirectly, during their entire stay within the nursery. To test this hypothesis, we conducted isotopic studies and stomach content analyses of juveniles from habitats near the mouth of Delaware Bay and from an adjacent lagoonal estuary (ca. 39.5° N, 75.1° W). Primary producers, marsh detritus, various life-history stages of blue crabs and potential prey species were sampled in the main estuary and in an adjacent marsh during the summer and early fall of two consecutive years. Newly settled juveniles (<15 mm carapace width) from the marsh were about 1.8‰ lighter in carbon (−17.2‰) relative to larger juveniles from the marsh (15–30 mm carapace width) and appeared to have retained a carbon isotopic signature indicative of the phytoplankton-based food web associated with larval stages. However, the signature of juveniles changed as a function of size. Large juveniles and crabs >60 mm were enriched in δ¹³C (−14.7 ± 0.1‰) compared to small crabs, suggesting a gradual shift in diet from a planktonic to a detritus-based food web with increasing size. As with crabs from Delaware Bay, the δ¹³C signature of juvenile crabs sampled from macroalgal beds in the lagoonal estuary (Rehoboth Bay) changed as a function of size. Also, δ¹³C ratios of crabs varied among the various species of macroalgae. The δ¹⁵N composition of primary producers in the marsh and main estuary also was reflected in the δ¹⁵N values of crabs and other benthic consumers in the respective habitats. Results of stomach-content analysis in this study were consistent with isotope data. Observed changes in prey preferences were related to changes in size of juvenile crabs and also differed among habitats. Gut content analyses of the three size classes of juveniles in macroalgal beds from Rehoboth Bay indicated that the crabs depend heavily on various amphipod species that occur on the seaweeds. These amphipods graze directly on the macroalgae and are among the most abundant invertebrates in the macroalgal beds. This implies a direct trophic relationship between the juvenile crabs and the macroalgae. In summary, our study provides strong evidence that the value of nursery areas such as salt marshes and macroalgal beds goes beyond that of providing refuge from predation, and that species using these nurseries (e.g. juvenile blue crabs) are ultimately dependent on primary production originating in benthic plants indigenous to the nursery.     

Invasive species cause large-scale loss of native California oyster habitat by disrupting trophic cascades

Although invasive species often resemble their native counterparts, differences in their foraging and anti-predator strategies may disrupt native food webs. In a California estuary, we showed that regions dominated by native crabs and native whelks have low mortality of native oysters (the basal prey), while regions dominated by invasive crabs and invasive whelks have high oyster mortality and are consequently losing a biologically diverse habitat. Using field experiments, we demonstrated that the invasive whelk’s distribution is causally related to a large-scale pattern of oyster mortality. To determine whether predator–prey interactions between crabs (top predators) and whelks (intermediate consumers) indirectly control the pattern of oyster mortality, we manipulated the presence and invasion status of the intermediate and top trophic levels in laboratory mesocosms. Our results show that native crabs indirectly maintain a portion of the estuary’s oyster habitat by both consuming native whelks (density-mediated trophic cascade) and altering their foraging behavior (trait-mediated trophic cascade). In contrast, invasive whelks are naive to crab predators and fail to avoid them, thereby inhibiting trait-mediated cascades and their invasion into areas with native crabs. Similarly, when native crabs are replaced with invasive crabs, the naive foraging strategy and smaller size of invasive crabs prevents them from efficiently consuming adult whelks, thereby inhibiting strong density-mediated cascades. Thus, while trophic cascades allow native crabs, whelks, and oysters to locally co-exist, the replacement of native crabs and whelks by functionally similar invasive species results in severe depletion of native oysters. As coastal systems become increasingly invaded, the mismatch of evolutionarily based strategies among predators and prey may lead to further losses of critical habitat that support marine biodiversity and ecosystem function. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The underrated importance of predation in transmission ecology of direct lifecycle parasites

Most parasites with complex life cycles exploit trophic webs to pass from host to host in order to develop and, eventually, reproduce. Thus predation constitutes the necessary route for transmission. Conversely, the transmission of parasites that use a single host to develop and reproduce should be, in principle, not particularly affected by host trophic ecology. Here I challenge this view, showing that predation may be relevant also for direct lifecycle parasites. I used a large dataset of fish trophic interactions to investigate if the degree of monogenean species overlap in predators and prey deviated from randomness. I demonstrated that predators and prey often share more monogenean parasite genera than explained by host habitat ecology, geographical distribution and phylogeny. This suggests that predation may play an important role in promoting monogenean host range expansion. In addition, a non‐negligible proportion of considered prey–predator pairs showed a significantly high overlap in their monogenean parasites at the species level. This may indicate a tendency of some monogenean parasites to evolve transmission strategies targeted towards host interactions. If this hypothesis is true, these monogenean parasites would be much more vulnerable to co‐extinction than previously thought. Synthesis Predation is not expected to play an important role in the ecology and evolution of simple life cycle parasites. Yet, several predator fish tend to share with their prey more monogenean parasites than one would expect predicted from their geographical distribution, habitat preference, and or phylogenetic relationships. This suggests that some monogenean parasites have evolved transmission strategies more targeted towards host interactions than towards species‐specific traits. If this hypothesis is supported, it would have strong implications on host–parasite evolutionary ecology, primarily, suggesting the existence of peculiar situations where some parasites have evolved high specialized host finding behaviors to expand their host range.     

Fish stocking creates an ecological trap for an avian predator via effects on prey availability

In anthropogenic landscapes animals may be lured into low‐quality habitats where they survive or reproduce poorly (‘ecological traps’). I investigated breeding habitat selection in relation to intra‐seasonal changes in food availability and reproductive output in red‐necked grebes Podiceps grisegena, a size‐limited predator, of common carp Cyprinus carpio ponds. Carp farms constitute highly heterogeneous habitat mosaics due to separate stocking of different age/size fish. Pond features significant for grebe settling decisions, i.e. hydroperiod and emergent vegetation cover, had no obvious effects on prey abundance for chicks and on fledging success. Breeding grebes avoided ponds containing fish too large for them to ingest but exhibited little preference between ponds with medium‐sized one‐year‐old carp that could be exploited by pre‐laying birds, and ponds designated for young‐of‐the‐year carp, where only invertebrates and amphibians were available as prey in early spring. Red‐necked grebes settling on ponds with medium‐sized fish failed to predict future shifts in interactions with carp stocks; carp exceeded the prey‐size threshold of chicks and adversely affected their non‐fish prey levels. The resulting food shortage led to severe egg‐to‐fledging mortality rates compared to fishless ponds or those containing young‐of‐the‐year fish. This study shows that waterbirds vulnerable to competition from fish can risk maladaptive habitat selection due to unrecognised spatial and temporal variation in food resources caused by fish stocking practices. Ecological traps created by perturbations to trophic interactions may be common but difficult to detect because altered dynamics of trophic resources can affect wildlife indirectly. As with other types of ecological traps, manipulation of habitat features identified as attractive cues for settling animals, but not related to critical food resources, may help to reduce perceptual pitfalls. For example, wetland management to mitigate trap effects driven by commercially stocked fish should preserve abundant emergent vegetation in habitats with weak fish impact and extend their hydroperiod.     

Variation in fish density,assemblage composition and relative rates of predation among mangrove,seagrass and coral reef habitats

We tested the hypothesis for several Caribbean reef fish species that there is no difference in nursery function among mangrove, seagrass and shallow reef habitat as measured by: (a) patterns of juvenile and adult density, (b) assemblage composition, and (c) relative predation rates. Results indicated that although some mangrove and seagrass sites showed characteristics of nursery habitats, this pattern was weak. While almost half of our mangrove and seagrass sites appeared to hold higher proportions of juvenile fish (all species pooled) than did reef sites, this pattern was significant in only two cases. In addition, only four of the six most abundant and commercially important species (Haemulon flavolineatum, Haemulon sciurus, Lutjanus apodus, Lutjanus mahogoni, Scarus iserti, and Sparisoma aurofrenatum) showed patterns of higher proportions of juvenile fish in mangrove and/or seagrass habitat(s) relative to coral reefs, and were limited to four of nine sites. Faunal similarity between reef and either mangrove or seagrass habitats was low, suggesting little, if any exchange between them. Finally, although relative risk of predation was lower in mangrove/seagrass than in reef habitats, variance in rates was substantial suggesting that not all mangrove/seagrass habitats function equivalently. Specifically, relative risk varied between morning and afternoon, and between sites of similar habitat, yet varied little, in some cases, between habitats (mangrove/seagrass vs. coral reefs). Consequently, our results caution against generalizations that all mangrove and seagrass habitats have nursery function.     

Species–area relationship within benthic habitat patches of a tropical floodplain river: An experimental test

The curvilinear relationship between species richness and habitat area (species–area relationship (SAR)) is a fundamental ecological pattern. The relationship is often viewed from a long‐term perspective across relatively large spatial scales, reflecting a balance between immigration and extinction dynamics. We explored whether predictions of SAR also manifest over short time periods (days) in benthic habitat patches of a dynamic floodplain river where littoral faunal assemblages are continuously assembled and disassembled with changing water levels. We examined the relationship of patch size with faunal abundance (i.e. fish and aquatic invertebrates), taxonomic richness, trophic group richness and overall assemblage composition. Strong taxa–area relationships emerged despite the relatively short experimental time period (21 days); larger patches had more taxa and trophic groups. For the smallest patches, taxonomic richness was especially sensitive to abundance of individuals; abundance of individuals was a less important predictor of taxonomic and trophic group richness for the largest patches. Despite the relatively short time frame for study within this temporally dynamic ecosystem, our findings indicate a strong SAR for fishes and macroinvertebrates inhabiting patchy habitats in the littoral zone of this tropical river.