Inducible defenses in Olympia oysters in response to an invasive predator

The prey naiveté hypothesis suggests that native prey may be vulnerable to introduced predators because they have not evolved appropriate defenses. However, recent evidence suggests that native prey sometimes exhibit induced defenses to introduced predators, as a result of rapid evolution or other processes. We examined whether Olympia oysters (Ostrea lurida) display inducible defenses in the presence of an invasive predator, the Atlantic oyster drill (Urosalpinx cinerea), and whether these responses vary among oyster populations from estuaries with and without this predator. We spawned oysters from six populations distributed among three estuaries in northern California, USA, and raised their offspring through two generations under common conditions to minimize effects of environmental history. We exposed second-generation oysters to cue treatments: drills eating oysters, drills eating barnacles, or control seawater. Oysters from all populations grew smaller shells when exposed to drill cues, and grew thicker and harder shells when those drills were eating oysters. Oysters exposed to drills eating other oysters were subsequently preyed upon at a slower rate. Although all oyster populations exhibited inducible defenses, oysters from the estuary with the greatest exposure to drills grew the smallest shells suggesting that oyster populations have evolved adaptive differences in the strength of their responses to predators. Our findings add to a growing body of literature that suggests that marine prey may be less likely to exhibit naiveté in the face of invasive predators than prey in communities that are more isolated from native predators, such as many freshwater and terrestrial island ecosystems.     

Ocean acidification impairs crab foraging behaviour

Anthropogenic elevation of atmospheric CO₂ is driving global-scale ocean acidification, which consequently influences calcification rates of many marine invertebrates and potentially alters their susceptibility to predation. Ocean acidification may also impair an organism''s ability to process environmental and biological cues. These counteracting impacts make it challenging to predict how acidification will alter species interactions and community structure. To examine effects of acidification on consumptive and behavioural interactions between mud crabs (Panopeus herbstii) and oysters (Crassostrea virginica), oysters were reared with and without caged crabs for 71 days at three pCO₂ levels. During subsequent predation trials, acidification reduced prey consumption, handling time and duration of unsuccessful predation attempt. These negative effects of ocean acidification on crab foraging behaviour more than offset any benefit to crabs resulting from a reduction in the net rate of oyster calcification. These findings reveal that efforts to evaluate how acidification will alter marine food webs should include quantifying impacts on both calcification rates and animal behaviour.     

Effect of Elevated pCO2 on Metabolic Responses of Porcelain Crab (Petrolisthes cinctipes) Larvae Exposed to Subsequent Salinity Stress

Future climate change is predicted to alter the physical characteristics of oceans and estuaries, including pH, temperature, oxygen, and salinity. Investigating how species react to the influence of such multiple stressors is crucial for assessing how future environmental change will alter marine ecosystems. The timing of multiple stressors can also be important, since in some cases stressors arise simultaneously, while in others they occur in rapid succession. In this study, we investigated the effects of elevated pCO₂ on oxygen consumption by larvae of the intertidal porcelain crab Petrolisthes cinctipes when exposed to subsequent salinity stress. Such an exposure mimics how larvae under future acidified conditions will likely experience sudden runoff events such as those that occur seasonally along portions of the west coast of the U.S. and in other temperate systems, or how larvae encounter hypersaline waters when crossing density gradients via directed swimming. We raised larvae in the laboratory under ambient and predicted future pCO₂ levels (385 and 1000 µatm) for 10 days, and then moved them to seawater at ambient pCO₂ but with decreased, ambient, or elevated salinity, to monitor their respiration. While larvae raised under elevated pCO₂ or exposed to stressful salinity conditions alone did not exhibit higher respiration rates than larvae held in ambient conditions, larvae exposed to elevated pCO₂ followed by stressful salinity conditions consumed more oxygen. These results show that even when multiple stressors act sequentially rather than simultaneously, they can retain their capacity to detrimentally affect organisms.     

Mixed Effects of Elevated pCO2 on Fertilisation,Larval and Juvenile Development and Adult Responses in the Mobile Subtidal Scallop Mimachlamys asperrima (Lamarck, 1819)

Ocean acidification is predicted to have severe consequences for calcifying marine organisms especially molluscs. Recent studies, however, have found that molluscs in marine environments with naturally elevated or fluctuating CO₂ or with an active, high metabolic rate lifestyle may have a capacity to acclimate and be resilient to exposures of elevated environmental pCO₂. The aim of this study was to determine the effects of near future concentrations of elevated pCO₂ on the larval and adult stages of the mobile doughboy scallop, Mimachlamys asperrima from a subtidal and stable physio-chemical environment. It was found that fertilisation and the shell length of early larval stages of M. asperrima decreased as pCO₂ increased, however, there were less pronounced effects of elevated pCO₂ on the shell length of later larval stages, with high pCO₂ enhancing growth in some instances. Byssal attachment and condition index of adult M. asperrima decreased with elevated pCO₂, while in contrast there was no effect on standard metabolic rate or pH_e. The responses of larval and adult M. asperrima to elevated pCO₂ measured in this study were more moderate than responses previously reported for intertidal oysters and mussels. Even this more moderate set of responses are still likely to reduce the abundance of M. asperrima and potentially other scallop species in the world’s oceans at predicted future pCO₂ levels.     

Different responses of invasive and native species to elevated CO2 concentration

Increasing atmospheric CO₂ concentration is regarded as an important factor facilitating invasion. However, the mechanisms by which invasive plants spread at the expense of existing native plants are poorly understood. In this study, three invasive species (Mikania micrantha, Wedelia trilobata and Ipomoea cairica) and their indigenous co-occurring species or congeners (Paederia scandens, Wedelia chinensis and Ipomoea pescaprae) in South China were exposed to elevated CO₂ concentration (700 μmol mol^?1). The invasive species showed an average increase of 67.1% in photosynthetic rate, significantly different from the native species (24.8%). On average the increase of total biomass at elevated CO₂ was greater for invasive species (70.3%) than for the natives (30.5%). Elevated CO₂ also resulted in significant changes in biomass allocation and morphology of invasive M. micrantha and W. trilobata. These results indicate a substantial variation in response to elevated CO₂ between these invasive and native plant species, which might be a potential mechanism partially explaining the success of invasion with ongoing increase in atmospheric CO₂.     

Elevated CO2 affects shell dissolution rate but not calcification rate in a marine snail

As CO₂ levels increase in the atmosphere, so too do they in the sea. Although direct effects of moderately elevated CO₂ in sea water may be of little consequence, indirect effects may be profound. For example, lowered pH and calcium carbonate saturation states may influence both deposition and dissolution rates of mineralized skeletons in many marine organisms. The relative impact of elevated CO₂ on deposition and dissolution rates are not known for many large-bodied organisms. We therefore tested the effects of increased CO₂ levels—those forecast to occur in roughly 100 and 200 years—on both shell deposition rate and shell dissolution rate in a rocky intertidal snail, Nucella lamellosa. Shell weight gain per day in live snails decreased linearly with increasing CO₂ levels. However, this trend was paralleled by shell weight loss per day in empty shells, suggesting that these declines in shell weight gain observed in live snails were due to increased dissolution of existing shell material, rather than reduced production of new shell material. Ocean acidification may therefore have a greater effect on shell dissolution than on shell deposition, at least in temperate marine molluscs.     

Feeding behavior of black carp Mylopharyngodon piceus (Pisces: Cyprinidae) on fry of other fish species and trematode transmitting snail species

Fish raised in aquaculture ponds may get infected with fishborne zoonotic trematodes (FZT) during the nursing stage. Freshwater snails serve as intermediate hosts for FZT and we wanted to explore the possibility of controlling snails by stocking nursery ponds with a few juvenile specimens of the mollusc-eating fish, black carp (Mylopharyngodon piceus). Obviously, the risk that black carp might also prey on the juvenile fishes in nursery ponds should first be assessed. Laboratory trials showed that all size classes of juvenile black carp consumed fry of common carp (Cyprinus carpio) even when offered snails as food; the odds of survival of fry from tanks with medium sized and large black carp was 5.6% and 39.9%, respectively of that of fry in tanks with small sized black carp. Since the large black carp also consumed fewer snails than medium sized fish, we believe that large specimens were stressed in the experimental aquaria. Under semi-field conditions, presence of the black carp had no effect on survival of fry of Oreochromis niloticus and C. carpio both in the absence and presence of snails as alternative food. The black carp consumed most snails offered with the exception of some of the large snails. We conclude that under field conditions, predation by black carp on fish fry is minimal and field trials in nursery ponds are warranted. Due to the risks that black carp pose to native imperiled snails and other molluscs, trials should be restricted to ponds within the fish’s native or existing range.     

Differential proteomic responses of selectively bred and wild‐type Sydney rock oyster populations exposed to elevated CO2

Previous work suggests that larvae from Sydney rock oysters that have been selectively bred for fast growth and disease resistance are more resilient to the impacts of ocean acidification than nonselected, wild‐type oysters. In this study, we used proteomics to investigate the molecular differences between oyster populations in adult Sydney rock oysters and to identify whether these form the basis for observations seen in larvae. Adult oysters from a selective breeding line (B2) and nonselected wild types (WT) were exposed for 4 weeks to elevated pCO₂ (856 μatm) before their proteomes were compared to those of oysters held under ambient conditions (375 μatm pCO₂). Exposure to elevated pCO₂ resulted in substantial changes in the proteomes of oysters from both the selectively bred and wild‐type populations. When biological functions were assigned, these differential proteins fell into five broad, potentially interrelated categories of subcellular functions, in both oyster populations. These functional categories were energy production, cellular stress responses, the cytoskeleton, protein synthesis and cell signalling. In the wild‐type population, proteins were predominantly upregulated. However, unexpectedly, these cellular systems were downregulated in the selectively bred oyster population, indicating cellular dysfunction. We argue that this reflects a trade‐off, whereby an adaptive capacity for enhanced mitochondrial energy production in the selectively bred population may help to protect larvae from the effects of elevated CO₂, whilst being deleterious to adult oysters.     

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.     

Competitive interactions between native and invasive exotic plant species are altered under elevated carbon dioxide

We hypothesized that the greater competitive ability of invasive exotic plants relative to native plants would increase under elevated CO₂ because they typically have traits that confer the ability for fast growth when resources are not limiting and thus are likely to be more responsive to elevated CO₂. A series of competition experiments under ambient and elevated CO₂ glasshouse conditions were conducted to determine an index of relative competition intensity for 14 native-invasive exotic species-pairs. Traits including specific leaf area, leaf mass ratio, leaf area ratio, relative growth rate, net assimilation rate and root weight ratio were measured. Competitive rankings within species-pairs were not affected by CO₂ concentration: invasive exotic species were more competitive in 9 of the 14 species-pairs and native species were more competitive in the remaining 5 species-pairs, regardless of CO₂ concentration. However, there was a significant interaction between plant type and CO₂ treatment due to reduced competitive response of native species under elevated compared with ambient CO₂ conditions. Native species had significantly lower specific leaf area and leaf area ratio under elevated compared with ambient CO₂. We also compared traits of more-competitive with less-competitive species, regardless of plant type, under both CO₂ treatments. More-competitive species had smaller leaf weight ratio and leaf area ratio, and larger relative growth rate and net assimilation rate under both ambient and elevated CO₂ conditions. These results suggest that growth and allocation traits can be useful predictors of the outcome of competitive interactions under both ambient and elevated CO₂ conditions. Under predicted future atmospheric CO₂ conditions, competitive rankings among species may not change substantially, but the relative success of invasive exotic species may be increased. Thus, under future atmospheric CO₂ conditions, the ecological and economic impact of some invasive exotic plants may be even greater than under current conditions.     

Predation on invasive zebra mussel, Dreissena polymorpha, by pumpkinseed sunfish, rusty crayfish, and round goby

The enemy release hypothesis states that invasive species are successful in their new environment because native species are not adapted to utilize the invasive. If true for predators, native predators should have lower feeding rates on the invasive species than a predator from the native range of the invasive species. We tested this hypothesis for zebra mussel (Dreissena polymorpha) by comparing handling time and predation rate on zebra mussels in the laboratory by two North American species (pumpkinseed, Lepomis gibbosus, and rusty crayfish, Orconectes rusticus) and one predator with a long evolutionary history with zebra mussels (round goby, Neogobius melanostomus). Handling time per mussel (7 mm shell length) ranged from 25 to >70 s for the three predator species. Feeding rates on attached zebra mussels were higher for round goby than the two native predators. Medium and large gobies consumed 50–67 zebra mussels attached to stones in 24 h, whereas pumpkinseed and rusty crayfish consumed <11. This supports the hypothesis that the rapid spread of zebra mussels in North America was facilitated by low predation rates from the existing native predators. At these predation rates and realistic goby abundance estimates, round goby could affect zebra mussel abundance in some lakes.     

Adaptation and acclimatization to ocean acidification in marine ectotherms: an in situ transplant experiment with polychaetes at a shallow CO2 vent system

Metabolic rate determines the physiological and life-history performances of ectotherms. Thus, the extent to which such rates are sensitive and plastic to environmental perturbation is central to an organism''s ability to function in a changing environment. Little is known of long-term metabolic plasticity and potential for metabolic adaptation in marine ectotherms exposed to elevated pCO₂. Consequently, we carried out a series of in situ transplant experiments using a number of tolerant and sensitive polychaete species living around a natural CO₂ vent system. Here, we show that a marine metazoan (i.e. Platynereis dumerilii) was able to adapt to chronic and elevated levels of pCO₂. The vent population of P. dumerilii was physiologically and genetically different from nearby populations that experience low pCO₂, as well as smaller in body size. By contrast, different populations of Amphiglena mediterranea showed marked physiological plasticity indicating that adaptation or acclimatization are both viable strategies for the successful colonization of elevated pCO₂ environments. In addition, sensitive species showed either a reduced or increased metabolism when exposed acutely to elevated pCO₂. Our findings may help explain, from a metabolic perspective, the occurrence of past mass extinction, as well as shed light on alternative pathways of resilience in species facing ongoing ocean acidification.     

Home-field advantage: native signal crayfish (Pacifastacus leniusculus) out consume newly introduced crayfishes for invasive Chinese mystery snail (Bellamya chinensis)

The introduction of non-indigenous plants, animals and pathogens is one of today’s most pressing environmental challenges. Freshwater ecologists are challenged to predict the potential consequences of species invasions because many ecosystems increasingly support novel assemblages of native and non-native species that are likely to interact in complex ways. In this study we evaluated how native signal crayfish (Pacifastacus leniusculus) and non-native red swamp crayfish (Procambarus clarkii) and northern crayfish (Orconectes virilis) utilize a novel prey resource: the non-native Chinese mystery snail (Bellamya chinensis). All species are widespread in the United States, as well as globally, and recent surveys have discovered them co-occurring in lakes of Washington State. A series of mesocosm experiments revealed that crayfish are able to consume B. chinensis, despite the snail’s large size, thick outer shell and trapdoor defense behaviour. Crayfish exhibited size-selective predation whereby consumption levels decreased with increasing snail size; a common pattern among decapod predators. Comparison of prey profitability curves—defined as the yield of food (weight of snail tissue) per second of feeding time (the time taken to crack the shell and consume the contents)—suggests that small and very large snails may represent the most profitable prey choice. By contrast, previous studies have reported the opposite pattern for crayfish consumption on thin-shelled snails. For all snail size classes, we found that native P. leniusculus and invasive O. virilis consumed greater numbers of snails than invasive P. clarkii. Moreover, P. leniusculus consistently handled and consumed snails at a faster pace compared to both invasive crayfishes across the range of snail sizes examined in our study. These results suggest not only that B. chinensis is a suitable food source for crayfish, but also that native P. leniusculus may ultimately out-consume invasive crayfishes for this new prey resource.     

Phenotypic Plasticity of the Introduced New Zealand Mud Snail,Potamopyrgus antipodarum,Compared to Sympatric Native Snails

Phenotypic plasticity is likely to be important in determining the invasive potential of a species, especially if invasive species show greater plasticity or tolerance compared to sympatric native species. Here in two separate experiments we compare reaction norms in response to two environmental variables of two clones of the New Zealand mud snail, Potamopyrgus antipodarum, isolated from the United States, (one invasive and one not yet invasive) with those of two species of native snails that are sympatric with the invader, Fossaria bulimoides group and Physella gyrina group. We placed juvenile snails in environments with high and low conductivity (300 and 800 mS) in one experiment, and raised them at two different temperatures (16°C and 22°C) in a second experiment. Growth rate and mortality were measured over the course of 8 weeks. Mortality rates were higher in the native snails compared to P. antipodarum across all treatments, and variation in conductivity influenced mortality. In both experiments, reaction norms did not vary significantly between species. There was little evidence that the success of the introduced species is a result of greater phenotypic plasticity to these variables compared to the sympatric native species.     

Effects of elevated CO2 on the vasculature and phenolic secondary metabolism of Plantago maritima

We have examined the effect of elevated CO₂ on the vasculature and phenolic secondary metabolism on clones of the maritime plant Plantago maritima (L.). Plants were exposed to either ambient (360 μmol CO₂ mol⁻¹) or elevated (600 μmol CO₂ mol⁻¹) atmospheric CO₂ within a Solardome facility and harvested after 12 months' growth. Histochemical analysis of the leaves identified increases in the diameter of the minor leaf vein and associated lignified vessels in plants exposed to elevated CO₂. In the roots the number of lignified root vessels and stele width were also increased, but overall the lignified vessel-wall thickness was reduced in plants exposed to elevated CO₂, compared to those grown under ambient CO₂. To investigate whether or not these subtle changes in lignification were associated with perturbations in phenolic metabolism, aromatic natural products were analysed by HPLC-MS after treatment with cellulase to hydrolyse the respective glycosidic conjugates. The phenylpropanoids p-coumaric acid, caffeic acid, ferulic acid and the flavone luteolin were identified, together with the caffeoyl phenylethanoid glycosides, verbascoside and plantamajoside which were resistant to enzymatic digestion. Exposure to enhanced CO₂ resulted in subtle changes in the levels of individual metabolites. In the foliage a one-year exposure to enhanced CO₂ resulted in an increased accumulation of caffeic acid, whilst in the roots p-coumaric acid and verbascoside were enhanced. Our results suggest that significant changes in the vasculature of P. maritima on exposure to increased CO₂ are associated with only minor changes in the leaves of specific lignin-related metabolites.     

Elevated CO<Subscript>2</Subscript> increases energy-use efficiency of invasive <Emphasis Type="Italic">Wedelia trilobata</Emphasis> over its indigenous congener

Increasing atmospheric CO₂ concentration is regarded as an important factor facilitating plants invasions by stimulating invasive species growth. However, the physiological mechanisms by which invasive plants increase at the expense of existing native plants are poorly understood. Plant growth is always related to energy-use process including energy assimilation and expenditure, and thus examination of energetic properties could provide mechanistic insight into growth responses to increased CO₂. The aims of this study were to examine the effect of rising CO₂ on the growth and energetic properties of alien invasive species (Wedelia trilobata (L.) Hitchc.) and its native congener (Wedelia chinensis (Osbeck.) Merr.) in South China, and to determine if the specific energetic properties of invasive species at elevated CO₂ favoring its growth. Elevated CO₂ stimulated a greater increase in biomass production for invasive W. trilobata (58.9%) than for its indigenous congener (48.1%). Meanwhile, elevated CO₂ altered the energetic properties differently upon species. For invasive W. trilobata, elevated CO₂ significantly increased total energetic gain via photosynthetic activity (A _total), but decreased energetic cost of biomass construction (CC), and thus enhanced photosynthetic energy-use efficiency (PEUE) by 85.3%. In contrast, the indigenous W. chinensis showed a slight increase in PEUE by 43.8%. Additionally, W. trilobata individuals grown in elevated CO₂ increased energy allocation towards stems. Statistic analysis revealed significant associations between growth characteristics (relative growth rate and biomass) and energetic properties (CC and PEUE), suggesting the greater growth stimulation in invasive species could be partly explained by its specific energetic properties in elevated CO₂ concentration. The invasive species showed a greater increase in energy-use efficiency under elevated CO₂, which consequently facilitated its growth. It might be a physiological mechanism promoting success of invasion with ongoing increase in atmospheric CO₂ concentration.     

Silent assassins: predation of native New Zealand trichopteran eggs by non-native freshwater gastropods

Aquatic insects that oviposit on rocks may evolve strategies to mitigate loss of eggs to opportunistic predation by common native grazers, but such strategies may be ineffective against non-native grazers. We tested whether the eggs of common New Zealand caddisfly families (Hydrobiosidae and Hydropsychidae) that oviposit on rocks were more susceptible to predation by native or non-native snails. The native snail Potamopyrgus antipodarum (Grey, 1843) and non-native snails Physa acuta Draparnaud, 1805 and Pseudosuccinea columella (Say, 1817) were presented with caddisfly egg masses and the number of individual eggs consumed was recorded after 24 and 48 hours. The larger Pseudosuccinea readily consumed the eggs, especially spumaline-encapsulated Hydrobiosidae eggs. Physa consumed very few Hydrobiosidae eggs and no Hydropsychidae eggs, whereas Potamopyrgus did not consume eggs of either family. The trichopteran egg masses tested did not succumb to predation by the native Potamopyrgus but are highly vulnerable to predation by a larger non-native snail.     

Growth of three cattail (<Emphasis Type="Italic">Typha</Emphasis>) taxa in response to elevated CO<Subscript>2</Subscript>

The establishment of non-native species and the increase in atmospheric CO₂, in combination, have the ability to alter current ecosystems. Previous studies have shown that invasive species tend to respond more strongly to CO₂ than natives, but these comparisons have been of different and unrelated species. To assess how response to CO₂ might be related to invasiveness per se, we compared a native (Typha latifolia) with a congeneric invasive (Typha angustifolia), as well as their hybrid (T. × glauca). All three taxa are common components of wetland vegetation, often occurring in near monocultures. An open-top chamber experiment was used to examine the effects of elevated and ambient CO₂ concentrations on the three taxa. All three increased rhizome biomass by 40% in elevated CO₂. Although the absolute increase did not differ among taxa, the invasive T. angustifolia had a much higher proportional response in biomass and photosynthetic rate (45 and 40% respectively). The weaker response of the two larger taxa native T. latifolia (16 and 2%) and hybrid T. × glauca (−4% and −1%) was possibly driven by soil nutrient deficiency, such that they were not able to benefit from increased CO₂. However, under low nutrients the smaller species T. angustifolia may become more a problematic invader in the future.