Effects of Lunatia heros predation on the population dynamics of Mya arenaria and Macoma balthica in Maine,USA

The interaction between the naticid snail predator Lunatia heros and 2 iteroparous, infaunal, intertidal bivalves was investigated in Lubec, Maine, USA. The Mya arenaria population consists primarily of young, small individuals. M. arenaria survivorship is low when young (3.5% yr^-1 for the first 5 yr), then increases. M. arenaria can attain a length of 60 mm, but it is susceptible to L. heros attack only until it is 30 mm long. It delays reproduction until it is 20 mm long (3.8 yr) and diverts its resources instead into rapid early growth (4.9 mm yr^-1 for the first 5 yr). The Macoma balthica population has a larger proportion of older individuals than does that of M. arenaria. Survivorship is higher for M. balthica than for M. arenaria (76.3% yr^-1 for the first 5 yr). Unlike M. arenaria, M. balthica attains a final length of only 25 mm and all sizes are susceptible to L. heros attack. M. balthica grows slowly (2.7 mm yr^-1 for the first 5 yr) and diverts its resources into earlier reproduction at a length of 10 mm (2.9 yr). These contrasting life-history patterns and the possible relationship between bivalve resource allocation and refuges from predation are discussed.     

Macroinfaunal structure and effects of thermal discharges in a mesohaline habitat of Chesapeake Bay,near a Nuclear Power Plant

The soft-bottom macroinfauna of the Chesapeake Bay near the Calvert Cliffs Nuclear Power Plant, Maryland, USA, was studied quarterly in 1977. A total of 42,900 organisms distributed among 55 taxa was taken in 180 Smith-McIntyre grabs. Two communities and an ecotone between these two communities were identified in association with three sediment types: sand, sand-clay, and clay. Reproductive activities of 8 numerically dominant species are described. The macroinfauna at Calvert Cliffs was largely seasonal; species richness and organism abundances were lowest in late summer. The polychaetes Scolecolepides viridis and Heteromastus filiformis and the bivalves Macoma balthica and Mya arenaria were dominant in March-June. In September-December, the polychaetes Neanthes succinea, Paraprionospio pinnata, and Glycinde solitaria and the bivalve Gemma gemma became dominant. There were clear indications of differences associated with sediment types in total number of species, total number of individuals, and abundances of the dominant species. The shallower and intermediate habitats had higher sand-clay ratios, lower organic carbon contents, and larger numbers of organisms than the deeper clay habitat. Within each depth, no apparent detrimental effects caused by the thermal discharges on total number of species, total number of individuals, abundances of the dominant species, species diversity or evenness were found. Increased abundance was observed at the discharge stations for a few species, especially for Mya arenaria and N. succinea.     

The capacity of the filter-feeding bivalve Mya arenaria L. to affect water transport in sandy beds

Benthic filter feeding macrofauna organisms may be an important factor linking sediment and water column. Transport of water and concomitantly of suspended matter is directly related to the size of the benthic filter feeding population. This paper aims to determine the potential for water transport by the bivalve Mya arenaria along a coastal stretch of roughly 100 km length in the southern Baltic Sea. Quantification of population filtration rates specific to the area is based on distribution, abundance and biomass of M. arenaria and calculated according to previously published filtration rate–biomass relations. Calculated rates range up to >8 m³ m^–2 day^–1 (at 5–20 m water depth in sandy sediment) with the potential to locally process a volume of water equivalent to the water column within <1 day. Data from 1991–2002 at one site suggest that the area-specific potential population filtration rate remains remarkably constant in time despite changes in population structure of M. arenaria. The related impact on pore water exchange within the permeable sediment associated with the leakage of water from the gape of M. arenaria valves is discussed.Communicated by L. Hagerman, Helsingør     

Diet and food preferences of the adult horseshoe crab Limulus polyphemus in Delaware Bay,New Jersey,USA

Adult horseshoe crabs Limulus polyphemus (L.) feed on a wide variety of infaunal and epifaunal invertebrates during their spring spawning migration in Delaware Bay, New Jersey, USA. Comparison of the gut contents with estimates of available prey showed that the most abundant potential prey item, the bivalve Gemma gemma, was avoided. The thinner shelled but comparatively scarce clam Mulinia lateralis was a preferred prey item. In the laboratory, crabs fed on G. gemma when it was the only available item but not when M. lateralis or soft-shell clams, Mya arenaria, were offered in conjunction. Large M. lateralis (>10mm) were preferred to small M. lateralis; there was no discrimination between M. lateralis and M. arenaria of the same size. Male and female horseshoe crabs had similar gut contents and laboratory feeding preferences, despite the fact that females are larger than males. Crabs spawning later in the summer contained more food than did crabs collected at the peak of spawning activity.     

Population dynamics of the soft-shell clam Mya arenaria

A life table was constructed for Mya arenaria from Gloucester, Massachusetts, USA, based on schedules of age-specific fecundity and mortality determined under natural conditions. Mortality rates decrease with size and age in this species, with the period of maximum mortality occurring during the summer months. Mortality rates during the fall and winter were considerably lower, perhaps due to the inactivity of natural predators. The survivorship curve for M. arenaria approximates the Type 3 curve of Deevey (1947). Mean life expectancy is low in recently-settled clams, peaks when the individual reaches 30.0 to 34.9 mm (1 year of age) and remains fairly high for most of the remainder of life. The intrinsic rate of natural increase (r _max) is very high: 4.74. This enormous rate of potential increase is offset by high rates of larval mortality in the plankton. Unlike the reproductive values of most animals studied, those in M. arenaria peak late in life, well after the know age of first reproduction. This is probably the result of increased fecundity with age. The implications of this work in the area of resource management are discussed.     

Importance of predation by infaunal polychaetes in controlling the structure of a soft-bottom community in Maine,USA

Field manipulations in a Maine, USA intertidal mud flat showed that Nereis virens adults were an important factor in regulating the abundance of Corophium volutator, an amphipod which comprised 63% of the total number of individuals in the benthic community. Removal of N. virens adults resulted in an increase in C. volutator abundance, while addition of N. virens adults led to a decrease. There was a significant negative correlation between the abundance of N. virens adults and that of C. volutator. C. volutator was also negatively correlated with the density of all the other infaunal species combined. These results suggest that there is at least a 3-level interactive system (N. virens —C. volutator — other infaunal species) within the infauna in Maine; N. virens is a factor in controlling the abundance of C. volutator, and C. volutator may play a role in regulating the densities of other infaunal species. Models of generalized cropping in soft-bottom systems may be too simple if they ignore complex trophic relationships within the infauna.     

Multiple predators indirectly alter community assembly across ecological boundaries

Models of habitat selection often assume that organisms choose habitats based on their intrinsic quality, regardless of the position of these habitats relative to low-quality habitats in the landscape. We created a habitat matrix in which high-quality (predator-free) aquatic habitat patches were positioned adjacent to (predator-associated) or isolated from (control) patches with single or two species of caged predators. After 16 days of colonization, larval insect abundance was reduced by 50% on average in both the predator and predator-associated treatments relative to isolated controls. Effects were largely similar among predator treatments despite variation in number of predator species, predator biomass, and whether predators were native or nonnative. Importantly, the strength of effects did not depend on whether predators were physically present. These results demonstrate that predator cues can cascade with equal strength across ecological boundaries, indirectly altering community assembly via habitat selection in intrinsically high-quality habitats.     

Potential effects of a non-indigenous predator in its expanded range: assessing green crab, <Emphasis Type="Italic">Carcinus maenas</Emphasis>, prey preference in a productive coastal area of Atlantic Canada

The non-indigenous green crab (Carcinus maenas) is an important predator on bivalve wild beds in coastal areas worldwide. This study explored size-dependent green crab prey preference on American oysters (Crassostrea virginica), blue mussels (Mytilus edulis), and soft-shell clams (Mya arenaria) in a productive coastal system of Atlantic Canada. Using two sizes of prey and three different experimental manipulations, small, medium, and large green crabs were given a choice among these three bivalves, and their daily feeding rates were monitored over the course of 3 days. For both prey sizes, green crabs showed an early feeding preference for soft-shell clams and, only as they declined in numbers, a switch toward mussels and subsequently toward oysters. We found that such changes in the timing (order) of prey preference are related to prey differences in shell thickness, a fairly reliable indicator of prey shell strength.     

Counterintuitive effects of large-scale predator removal on a midlatitude rodent community

Historically, small mammals have been focal organisms for studying predator-prey dynamics, principally because of interest in explaining the drivers of the cyclical dynamics exhibited by northern vole, lemming, and hare populations. However, many small-mammal species occur at relatively low and fairly stable densities at temperate latitudes, and our understanding of how complex predator assemblages influence the abundance and dynamics of these species is surprisingly limited. In an intact grassland ecosystem in western Montana, USA, we examined the abundance and dynamics of Columbian ground squirrels (Spermophilus columbianus), deer mice (Peromyscus maniculatus), and montane voles (Microtus montanus) on 1-ha plots where we excluded mammalian and avian predators and ungulates, excluded ungulates alone, or allowed predators and ungulates full access. Our goal was to determine whether the relatively low population abundance and moderate population fluctuations of these rodents were due to population suppression by predators. Our predator-exclusion treatment was divided into two phases: a phase where we excluded all predators except weasels (Mustela spp.; 2002-2005), and a phase where all predators including weasels were excluded (2006-2009). Across the entire duration of the experiment, predator and/or ungulate exclusion had no effect on the abundance or overall dynamics of ground squirrels and deer mice. Ground squirrel survival (the only species abundant enough to accurately estimate survival) was also unaffected by our experimental treatments. Prior to weasel exclusion, predators also had no impacts on montane vole abundance or dynamics. However, after weasel exclusion, vole populations reached greater population peaks, and there was greater recruitment of young animals on predator-exclusion plots compared to plots open to predators during peak years. These results suggest that the impacts of predators cannot be generalized across all rodents in an assemblage. Furthermore, they suggest that specialist predators can play an important role in suppressing vole abundance even in lower-latitude vole populations that occur at relatively low densities.     

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.     

Local interactions between predators and prey call into question commonly used functional responses

Commonly used functional response models (Holling’s type I and type II models) assume that the encounter rate of a predator increases linearly with prey density, provided that the predator is searching for prey. In other other words, aN (a is the baseline encounter rate and N is prey density) describes the encounter rate. This study examined whether the models are adequate when predators and prey interact locally by using a spatially explicit individual based model because local interactions affect the spatial distribution of predators and prey, which also affects the encounter rate. Predators were assumed to possess a spatial perception range that influenced their foraging behavior (e.g., if a prey is in the perception range, the predator moves towards the prey). The effect of antipredator behavior by prey was also examined. The results suggest that prey and predator densities as well as handling time affect the baseline rate (i.e., parameter a) as opposed to the common assumption that the parameter is constant. The nature of model deviations depended on both the antipredator behavior and the predators’ perception range. Understanding these deviations is important as they qualitatively affect community dynamics.     

Siphon size and burying depth in deposit- and suspension-feeding benthic bivalves

This paper analyses the significance of siphon investment in the life strategy of benthic bivalves. It describes the relationships between siphon weight, burying depth and shell size in Mya arenaria, Cerastoderma edule. Scrobicularia plana and Macoma balthica. All data were collected on an intertidal flat in the Dutch Wadden Sea during seven successive winter and summer periods. The four species have in common that (1) the increase of depth in relation to size can be described with an S-curve; (2) there is a linear relationship between log siphon weight and log shell size; (3) siphon investment is maximal for the size classes with the greatest increase in their depth; (4) siphon weight, in proportion to total body weight, decreases gradually for the larger size classes whose depth does not increase; (5) burying depth increases with siphon weight if individuals within a same size class are compared, but burying depth levels off above a certain siphon weight. Macoma balthica and Scrobicularia plana live twice as deep in winter as in summer, although siphon weight for both seasons is about the same. In summer both species use a part of the siphon to graze the surface around the burrow, whereas deposit feeding does not occur in winter. This might explain the seasonal variation in burying depth. On the other hand Cerastoderma edule and Mya arenaria, which are both suspension feeders, show hardly any increase of depth in winter as compared to summer. For benthic bivalves the risk of being taken by a predator decreases with depth. The burying depth levels off where individuals reach the depth refuge (though in winter Scrobicularia plana live at greater depth). The conclusion is that siphon size is one of the main factors determining the burying depth of benthic bivalves and thus plays a critical role in their survival.     

Positive and Negative Effects of a Threatened Parrotfish on Reef Ecosystems

Species that are strong interactors play disproportionately important roles in the dynamics of natural ecosystems. It has been proposed that their presence is necessary for positively shaping the structure and functioning of ecosystems. We evaluated this hypothesis using the case of the world's largest parrotfish (Bolbometopon muricatum), a globally imperiled species. We used direct observation, animal tracking, and computer simulations to examine the diverse routes through which B. muricatum affects the diversity, dispersal, relative abundance, and survival of the corals that comprise the foundation of reef ecosystems. Our results suggest that this species can influence reef building corals in both positive and negative ways. Field observation and simulation outputs indicated that B. muricatum reduced the abundance of macroalgae that can outcompete corals, but they also feed directly on corals, decreasing coral abundance, diversity, and colony size. B. muricatum appeared to facilitate coral advancement by mechanically dispersing coral fragments and opening up bare space for coral settlement, but they also damaged adult corals and remobilized a large volume of potentially stressful carbonate sediment. The impacts this species has on reefs appears to be regulated in part by its abundance—the effects of B. muricatum were more intense in simulation scenarios populated with high densities of these fish. Observations conducted in regions with high and low predator (e.g., sharks) abundance generated results that are consistent with the hypothesis that these predators of B. muricatum may play a role in governing their abundance; thus, predation may modulate the intensity of the effects they have on reef dynamics. Overall our results illustrate that functionally unique and threatened species may not have universally positive impacts on ecosystems and that it may be necessary for environmental managers to consider the diverse effects of such species and the forces that mediate the strength of their influence. Efectos Positivos y Negativos de un Pez Loro Amenazado Sobre Ecosistemas Arrecifales     

Behavioral responses to a novel predator and competitor of invasive mosquitofish and their non-invasive relatives (Gambusia sp.)

Attributes of the recipient community may affect the invasion success of arriving non-indigenous organisms. In particular, biotic interactions may enhance the resistance of communities to invasion. Invading organisms typically encounter a novel suite of competitors and predators, and thus their invasiveness may be affected by how they cope with these interactions. Behavioral plasticity may help invaders to respond appropriately to novelty. We examined the behavioral responses of highly invasive mosquitofish to representative novel competitors and predators they might encounter as they spread through North America. We compared the behavior of invasive Gambusia holbrooki and G. affinis to that of two close relatives of lower invasive potential (G. geiseri and G. hispaniolae) in order to elucidate whether responses to novelty related to invasiveness. In short-term assays, female Gambusia were paired with a novel competitor, Pimephales promelas, and a novel predator, Micropterus dolomieu. Behavioral responses were measured in terms of foraging success and efficiency, activity, refuge use, predator inspections, and interspecific aggression. Contrary to a priori predictions, invasive and non-invasive responses to novel interactions did not differ consistently. In response to novel competition, both invasive species increased foraging efficiency, but so did G. geiseri. In response to novel predation, only G. holbrooki decreased consumption and activity and increased refuge use. No antipredator response was observed in G. affinis. We found consistent differences, however, between invasives and non-invasives in foraging behavior. Both in the presence and absence of the competitor and the predator, invasives foraged more efficiently and consumed more prey than non-invasives.Communicated by P. Bednekoff     

Comparison of sterol composition of some Baltic Sea and Black Sea mussels

Sterol composition of three Baltic Sea mussels (Mytilus edulis, Cardium glaucum, Mya arenaria) was compared with the sterol composition of three biologically closely related mussels from the Black Sea (Mytilus galloprovincialis, Cardium edule, Mya arenaria). The qualitative sterol composition of the mussels studied was similar, but some quantitative differences were found.     

Ecological and Economic Services Provided by Birds on Jamaican Blue Mountain Coffee Farms

Abstract: Coffee farms can support significant biodiversity, yet intensification of farming practices is degrading agricultural habitats and compromising ecosystem services such as biological pest control. The coffee berry borer (Hypothenemus hampei) is the world's primary coffee pest. Researchers have demonstrated that birds reduce insect abundance on coffee farms but have not documented avian control of the berry borer or quantified avian benefits to crop yield or farm income. We conducted a bird‐exclosure experiment on coffee farms in the Blue Mountains, Jamaica, to measure avian pest control of berry borers, identify potential predator species, associate predator abundance and borer reductions with vegetation complexity, and quantify resulting increases in coffee yield. Coffee plants excluded from foraging birds had significantly higher borer infestation, more borer broods, and greater berry damage than control plants. We identified 17 potential predator species (73% were wintering Neotropical migrants), and 3 primary species composed 67% of migrant detections. Average relative bird abundance and diversity and relative resident predator abundance increased with greater shade‐tree cover. Although migrant predators overall did not respond to vegetation complexity variables, the 3 primary species increased with proximity to noncoffee habitat patches. Lower infestation on control plants was correlated with higher total bird abundance, but not with predator abundance or vegetation complexity. Infestation of fruit was 1–14% lower on control plants, resulting in a greater quantity of saleable fruits that had a market value of US$44–$105/ha in 2005/2006. Landscape heterogeneity in this region may allow mobile predators to provide pest control broadly, despite localized farming intensities. These results provide the first evidence that birds control coffee berry borers and thus increase coffee yield and farm income, a potentially important conservation incentive for producers.     

Effects of different management strategies on long-term trends of Australian threatened and near-threatened mammals

Monitoring is critical to assess management effectiveness, but broadscale systematic assessments of monitoring to evaluate and improve recovery efforts are lacking. We compiled 1808 time series from 71 threatened and near-threatened terrestrial and volant mammal species and subspecies in Australia (48% of all threatened mammal taxa) to compare relative trends of populations subject to different management strategies. We adapted the Living Planet Index to develop the Threatened Species Index for Australian Mammals and track aggregate trends for all sampled threatened mammal populations and for small (<35 g), medium (35–5500 g), and large mammals (>5500 g) from 2000 to 2017. Unmanaged populations (42 taxa) declined by 63% on average; unmanaged small mammals exhibited the greatest declines (96%). Populations of 17 taxa in havens (islands and fenced areas that excluded or eliminated introduced red foxes [Vulpes vulpes] and domestic cats [Felis catus]) increased by 680%. Outside havens, populations undergoing sustained predator baiting initially declined by 75% but subsequently increased to 47% of their abundance in 2000. At sites where predators were not excluded or baited but other actions (e.g., fire management, introduced herbivore control) occurred, populations of small and medium mammals declined faster, but large mammals declined more slowly, than unmanaged populations. Only 13% of taxa had data for both unmanaged and managed populations; index comparisons for this subset showed that taxa with populations increasing inside havens declined outside havens but taxa with populations subject to predator baiting outside havens declined more slowly than populations with no management and then increased, whereas unmanaged populations continued to decline. More comprehensive and improved monitoring (particularly encompassing poorly represented management actions and taxonomic groups like bats and small mammals) is required to understand whether and where management has worked. Improved implementation of management for threats other than predation is critical to recover Australia's threatened mammals.     

The interplay between foraging mode, habitat structure, and predator presence in antlions

Antlion larvae are sand-dwelling insect predators, which ambush small arthropod prey while buried in the sand. In some species, the larvae construct conical pits and are considered as sit-and-wait predators which seldom relocate while in other species, they ambush prey without a pit but change their ambush site much more frequently (i.e., sit-and-pursue predators). The ability of antlion larvae to evade some of their predators which hunt them on the sand surface is strongly constrained by the degree of sand stabilization or by sand depth. We studied the effect of predator presence, predator type (active predatory beetle vs. sit-and-pursue wolf spider), and sand depth (shallow vs. deep sand) on the behavioral response of the pit building Myrmeleon hyalinus larvae and the sit-and-pursue Lopezus fedtschenkoi larvae. Predator presence had a negative effect on both antlion species activity. The sit-and-wait M. hyalinus larvae showed reduced pit-building activity, whereas the sit-and-pursue L. fedtschenkoi larvae decreased relocation activity. The proportion of relocating M. hyalinus was negatively affected by sand depth, whereas L. fedtschenkoi was negatively affected also by the predator type. Specifically, the proportion of individual L. fedtschenkoi that relocated in deeper sand was lower when facing the active predator rather than the sit-and-pursue predator. The proportion of M. hyalinus which constructed pits decreased in the presence of a predator, but this pattern was stronger when exposed to the active predator. We suggest that these differences between the two antlion species are strongly linked to their distinct foraging modes and to the foraging mode of their predators. Reut Loria and Inon Scharf contributed equally to the paper.     

Variation and persistence of the middle rocky intertidal community of central Chile,with and without human harvesting

Humans are an important intertidal predator in central Chile. Following a five-year study we report on the effects of the exclusion of humans from the rocky intertidal at Las Cruces, central Chile. The middle intertidal of harvested and non-harvested areas diverged in species diversity and composition during the experiment. In harvested areas the middle intertidal rocky shore was dominated throughout the study by a monoculture of mussels, Perumytilus purpuratus. When humans were excluded, the middle intertidal community switched to one dominated by barnacles (predominantly Jehlius cirratus and Chthamalus scabrosus); this community has persisted for at least three years, despite the presence of forces e.g. mussel larvae, that have the potential to alter the community structure. Such changes were mediated by the muricid gastropod Concholepas concholepas, a key-stone predator. As a consequence of the above changes, the species diversity, H', (primary space occupiers) in the non-harvested area increased from H'=0 at the beginning of the study in 1983 (when the middle intertidal community was dominated by mussels), to values ca H'=2 toward the middle of the study in 1984 (which coincided with the maximum predatory impact of C. concholepas) and subsequently decreasing to ca H'=0.5 at the end of the study in 1987 (when the middle intertida community was dominated by barnacles).Please address all correspondence to J. C. Castilla, Department of Zoology, University of Washington, Seattle, Washington 98195, USA, until March 1990