Prey preferences of invasive (Hemigrapsus sanguineus, H. takanoi) and native (Carcinus maenas) intertidal crabs in the European Wadden Sea

Invasive predators can have wide-ranging effects on invaded ecosystems and identifying the prey spectra and preferences of invaders are important steps in assessing their potential impacts on native biota. In this study, we investigated prey preferences of two invasive crab species (Hemigrapsus sanguineus and Hemigrapsus takanoi) that recently invaded Europe's shores and compared them with preferences of native shore crabs (Carcinus maenas) of similar size. In laboratory experiments, all three crab species preferred animal over algal prey. In general, sessile mussels (Mytilus edulis) were preferred over motile amphipods (Gammarus locusta) by all three crab species but amphipod predation was lower in the invasive compared with the native crabs. For the two invasive crab species, this pattern was the same in treatments where prey was offered separately (no-choice treatments) or simultaneously (choice treatments), while for the native crabs, mussel preference disappeared in choice treatments. The general preference of mussels by all three crab species suggests that local invasions of crabs most likely lead to increased competition among crabs. In addition, given that local densities of invasive crabs are often much higher than those of native crabs, predation pressure on native mussels can be expected to strongly increase at invaded sites. In contrast, local predation pressure on amphipods may be less affected by the crab invasions. Further field studies are needed to establish the magnitude of competition and predation pressure exerted by the invaders under natural conditions

Invasive oysters as new hosts for native shell-boring polychaetes: Using historical shell collections and recent field data to investigate parasite spillback in native mussels in the Dutch Wadden Sea

Biological invasions can have manifold effects on native biota, including impacts on species interactions in invaded ecosystems. Among those are effects on parasite-host interactions, for example in cases where invaders serve as a new host for native parasites, leading to an amplification of the parasite population which may ultimately result in increased infection levels in the original native hosts (parasite spillback). In this study, we investigated the potential spillback of the native shell-boring polychaete Polydora ciliata from invasive Pacific oysters (Crassostrea (Magallana) gigas) to native mussels (Mytilus edulis) in the Dutch Wadden Sea. A field survey in the intertidal revealed that the majority of P. ciliata can nowadays be found in the shells of live but also dead Pacific oysters and much less in shells of other potential host species: mussels and periwinkles (Littorina littorea). Using a unique historical shell collection based on long-term sampling programmes in the intertidal and subtidal of the western Dutch Wadden Sea, we compared P. ciliata infections in mussels before and after the invasion of the Pacific oyster by means of x-ray scans. Both for the intertidal and subtidal, we did not detect differences in prevalence of P. ciliata in mussels between pre- and post-invasion periods. This suggests that the invasion of the Pacific oyster may not have caused a strong spillback to mussels regarding P. ciliata and thus the invasion probably had little indirect infection-mediated effects on the condition and fitness of native mussels. Instead, the acquisition of native P. ciliata by the invasive oysters suggests that they may themselves be affected by the new infections and this may warrant further research

Large-scale patterns in trematode richness and infection levels in marine crustacean hosts

Little is known about the patterns of variation in parasitism in marine hosts. Trematodes, the dominant parasites in intertidal systems, are transmitted from their first intermediate hosts (snails) to a range of second intermediate hosts, including crustaceans. Using published studies of trematode infections in crustacean hosts, we investigated general patterns of variation in trematode species richness and infection levels (i.e. percentage of hosts infected and mean number of individual parasites per host). Since the production and release of infective stages in snails is strongly temperature dependent, we also investigated a potential decrease in trematode infection levels with increasing latitude (as a proxy for decreasing temperature). Trematode species richness in the crustacean hosts was generally low (mostly 1 or 2), and infection levels were moderate. However, there were differences among taxa in some groups, particularly among brachyuran crabs, which showed significantly higher values than in other groups. For amphipods, which were the only well-studied group across a large range of latitudes, we found negative correlations between latitude and the trematode species richness or measures of infection level considered here. These relationships persisted after correction of the potentially confounding effects of sampling effort, host body size and host generic identity (as a control for phylogenetic influences). We discuss these findings in light of environmental mediation of parasite transmission, in particular with respect to the probably fundamental role of temperature in driving the output of trematode infective stages in marine systems

Freshening rather than warming drives trematode transmission from periwinkles to mussels

In the Western Baltic Sea, climate change is happening at much faster rate than in most other seas and organisms are additionally exposed to a steep and variable salinity gradient. Climate change has previously been shown to affect parasite transmission in other marine ecosystems, yet little is known about potential effects of warming and desalination on parasite–host interactions. In laboratory experiments, we determined the combined effects of projected seawater warming and freshening on the emergence, activity, survival, and infectivity of cercariae (free-swimming infectious stage) of the trematode Himasthla elongata (Mehlis 1831), shed from its first intermediate host, the periwinkle Littorina littorea (Linnaeus 1758), in the Baltic Sea. We also assessed the susceptibility of the second intermediate host, the mussel Mytilus edulis Linnaeus, 1758, to cercarial infections. Generally, salinity was the main driver, particularly of cercarial activity, infectivity, and mussel susceptibility to infection. At the lowest salinity (13), cercariae were 50% less active compared to the highest salinity (19). Infection success and host susceptibility followed a similar pattern, with 47% and 43% less metacercariae (encysted stage) present at salinity 13 than at salinity 19, respectively. In contrast, effects of simulated warming were found only for cercarial survival, with cercarial longevity being higher at 19 than at 23 °C. No significant interactions between temperature and salinity were found. In contrast to the literature, the results suggest that a climate change-driven freshening (partly also warming) may lead to a general decline of marine trematodes, with possible beneficial effects for the involved hosts

Über die Beurteilung des Gütegrades von Mischungen bei beliebigen Verteilungsgesetzen für die Korngewichte der einzelnen Mischungskomponenten

Two granular materials (P) and (Q) are mixed. The distribution functions of particle weight are supposed to be known for both components (P) und (Q). To judge the efficiency of the mixing process a statistical theory is developed to calculate the variations of the mixture's composition when samples are taken at random from the container. It is important to distinguish the notions of "volume frequency", "weight frequency", and "particle frequency" of the different components within the mixture. Single particles are supposed to obey statistical laws. The results which are stictly valid only for samples with equal numbers of particles are shown to be approximately valid for samples of equal weight (or volume), too. - The theory is generalised for mixtures consisting of three or more components

Biological invasions and host–parasite coevolution: different coevolutionary trajectories along separate parasite invasion fronts

Host–parasite coevolution has rarely been observed in natural systems. Its study often relies on microparasitic infections introducing a potential bias in the estimation of the evolutionary change of host and parasite traits. Using biological invasions as a tool to study host–parasite coevolution in nature can overcome these biases. We demonstrate this with a cross-infection experiment in the invasive macroparasite <i>Mytilicola intestinalis</i> and its bivalve host, the blue mussel <i>Mytilus edulis</i>. The invasion history of the parasite is well known for the southeastern North Sea and is characterised by two separate invasion fronts that reached opposite ends of the Wadden Sea (i.e. Texel, The Netherlands and Sylt, Germany) in a similar time frame. The species’ natural history thus makes this invasion an ideal natural experiment to study host–parasite coevolution in nature. We infected hosts from Texel, Sylt and Kiel (Baltic Sea, where the parasite is absent) with parasites from Texel and Sylt, to form sympatric, allopatric and naïve infestation combinations, respectively. We measured infection rate, host condition and parasite growth to show that sympatric host–parasite combinations diverged in terms of pre- and post-infection traits within <100 generations since their introduction. Texel parasites were more infective and more efficient at exploiting the host’s resources. Hosts on Texel, on the other hand, evolved resistance to infection, whereas hosts on Sylt may have evolved tolerance. This illustrates that different coevolutionary trajectories can evolve along separate invasion fronts of the parasite, highlighting the use of biological invasions in studies of host–parasite coevolution in nature

Effects of first intermediate host density, host size and salinity on trematode infections in mussels of the south-western Baltic Sea

Trematode prevalence and abundance in hosts are known to be affected by biotic drivers as well as by abiotic drivers. In this study, we used the unique salinity gradient found in the south-western Baltic Sea to: (i) investigate patterns of trematode infections in the first intermediate host, the periwinkle Littorina littorea and in the downstream host, the mussel Mytilus edulis, along a regional salinity gradient (from 13 to 22) and (ii) evaluate the effects of first intermediate host (periwinkle) density, host size and salinity on trematode infections in mussels. Two species dominated the trematode community, Renicola roscovita and Himasthla elongata. Salinity, mussel size and density of infected periwinkles were significantly correlated with R. roscovita, and salinity and density correlated with H. elongata abundance. These results suggest that salinity, first intermediate host density and host size play an important role in determining infection levels in mussels, with salinity being the main major driver. Under expected global change scenarios, the predicted freshening of the Baltic Sea might lead to reduced trematode transmission, which may be further enhanced by a potential decrease in periwinkle density and mussel size

Spread of the invasive shell-boring annelid Polydora websteri (Polychaeta, Spionidae) into naturalised oyster reefs in the European Wadden Sea

With globally growing aquaculture activities, the co-introduction of parasites alongside large-scale movements of commercial species poses an increasing risk for marine ecosystems. Here, we present the first record of the shell-boring polychaete Polydora websteri Hartman in Loosanoff and Engle, 1943 in invasive Pacific oysters Crassostrea (Magallana) gigas (Thunberg, 1793) in the European Atlantic Ocean. In October 2014, mud blisters in the shells of wild Pacific oysters and specimens of a spionid polychaete were observed in close proximity to a commercial oyster farm at the island of Sylt (Germany) in the European Wadden Sea. Subsequent investigations indicated that these blisters only occurred near the farm and that no other mollusc species were affected. Morphological and molecular analysis identified the polychaete as Polydora websteri, a species that nowadays widely occurs around the globe, but likely is native to the Asian Pacific. Later sampling activities detected P. websteri also at other locations around Sylt as well as in the Dutch part of the Wadden Sea at the island of Texel. The number of polychaetes in the oysters was, however, relatively low and mostly below 10 individuals per oyster. Together, this evidence suggests that P. websteri is currently extending its range. As the introduction of P. websteri may have severe ecological and economic implications, this study aims to alert others to look for P. websteri at Western European coasts within farmed or wild Pacific oysters to further document its spread

Cryptic invasion of a parasitic copepod: Compromised identification when morphologically similar invaders co-occur in invaded ecosystems

Despite their frequent occurrence and strong impacts on native biota, biological invasions can long remain undetected. One reason for this is that an invasive species can be morphologically similar to either native species or introduced species previously established in the same region, and thus be subject to mistaken identification. One recent case involves congeneric invasive parasites, copepods that now infect bivalve hosts along European Atlantic coasts, after having been introduced independently first from the Mediterranean Sea (Mytilicola intestinalis Steuer, 1902) and later from Japan (Mytilicola orientalis Mori, 1935). At least one report on M. intestinalis may have actually concerned M. orientalis, and M. orientalis thus qualifies as a "cryptic invader". Because these two parasitic copepods are morphologically similar, knowledge about their distribution, impact and interactions depends crucially on reliable species identification. In this study, we evaluated the reliability of morphological identification of these two species in parts of their invasive range in Europe (Dutch Delta and Wadden Sea) in comparison with molecular methods of well-established accuracy based on COI gene sequences and ITS1 restriction fragment length polymorphism. Based on seven easily measured or scored macro-morphological variables that were recorded for 182 individual copepods isolated from blue mussels (Mytilus edulis Linnaeus, 1758), principal component analysis showed two relatively distinct but overlapping morphological species groups for females, but no clear separation in males. Discriminant function analysis showed that the females can be discriminated reasonably well based on some of the morphological characteristics (identification error rate of 7%) while males cannot (error rate of 25%). The direction of the dorsolateral thoracic protuberances was identified as the most important trait for species discrimination, but among the morphological features checked, none could flawlessly discriminate between both species. We recommend the use of molecular techniques in future studies of invasive Mytilicola to reliably discriminate between the species. The morphological similarity of these two invaders suggests a more general problem of cryptic invasions and compromised identification of parasites in invaded ecosystems. This problem should be borne in mind whenever invasive parasites are investigated

Production of marine trematode cercariae: a potentially overlooked path of energy flow in benthic systems

Parasites, in particular trematodes, are unseen but ubiquitous components of marine intertidal ecosystems. Although parasites are known to affect population dynamics and food web structure, their potential function as an unrecognized path of energy flow in these ecosystems is yet to be quantified. We use published data on rates at which trematodes produce free-swimming infective larvae (cercariae) that are released from their gastropod intermediate hosts to investigate patterns in cercarial output as a function of different variables, and to calculate the annual production of cercariae in different marine benthic systems. Across 18 trematode species, cercarial output (no. cercariae shed snail–1 d–1) ranged over 4 orders of magnitude and was positively correlated with snail host species size. While cercarial output did not correlate with latitude, it did correlate negatively with the size of cercariae, and was influenced by the type of downstream host sought by cercariae, being highest when this host was a vertebrate. Our estimates of annual cercarial production (kJ m–2 yr–1), which take into account the density of infected snails in the habitat, were within the range of production values reported for free-living invertebrates inhabiting benthic ecosystems. These estimates would be much higher if they included all trematode species in an ecosystem, and not just single-species values. Overall, results suggest that trematode cercariae represent potentially important paths of energy flow in benthic systems as well as a potentially important food supply to benthic organisms