Description: The Asian shore crab H. sanguineus first appeared at the French coast in the late 1990’s. It rapidly extended its range further north to the German Wadden Sea and recently to western Sweden. In the intertidal area, it co-occurs with the European green crab C. maenas. As the ecophysiology of H. sanguineus is virtually unknown, the physiological capacities of both species and their potential for intra-guild competition were investigated. The aim of this study was to specifically compare the energy deposition and dietary preferences of ovigerous females of both species. Females of H. sanguineus and C. maenas carrying immature or mature eggs were collected in April, June, August and October 2015 in an intertidal area of the Island of Helgoland, North Sea. Total lipid levels and fatty acid compositions were determined of both midgut glands and eggs. In H. sanguineus, total lipid levels of the midgut glands were clearly higher than those of C. maenas (40% vs. 10% dry mass, DM). Immature eggs were quite lipid-rich in both species with 30% and 25%DM, respectively, whereas in mature eggs, lipid levels decreased to ~15%DM each. A Principal Component Analysis of the fatty acid compositions of midgut glands and eggs revealed separate clusters for both species with C. maenas lipids more characterized by membrane fatty acids. In C. maenas fatty acids of midgut glands and eggs clustered together largely dominated by carnivory biomarkers. Fatty acids of midgut glands and all eggs of H. sanguineus formed separate clusters and trophic markers indicate a more herbivorous diet. Higher lipid levels and thus more pronounced energy deposition in H. sanguineus midgut glands indicate higher starvation tolerance for females, a potential competitive advantage over C. maenas. Direct food competition, however, seems negligible, as H. sanguineus prefers a more herbivorous diet than C. maenas. Deviating fatty acid compositions in H. sanguineus midgut glands and eggs suggest that this species may represent an income breeder, utilizing energy from both the midgut gland but also from dietary input. Most brachyuran crabs are capital breeders, which rely exclusively on internal reserves.

Description: The intertidal Asian shore crab H. sanguineus was initially found on the French coast in the late 1990’s. It rapidly extended its range further north and is now well established in the German Wadden Sea. Recently, it was also found in western Sweden. In its intertidal habitat, it co-occurs with the European green crab C. maenas. The ecophysiology of H. sanguineus is virtually unknown. In this study, the physiological capacities of both species and their potential for intra-guild competition were investigated. We specifically compared the energy deposition and dietary preferences of ovigerous females of both species. Females of both species carrying immature or mature eggs were collected in April, June, August and October 2015 in an intertidal site on the Island of Helgoland, North Sea, Germany. Total lipid levels of midgut glands and eggs were acquired via extraction. Subsequently, fatty acid compositions were determined through gas chromatography. Total lipid levels of H. sanguineus midgut glands were clearly higher than those of C. maenas (40% vs. 10% dry mass, DM). Immature eggs of both species were quite lipid-rich with 30% and 25%DM, respectively. In mature eggs, lipid levels decreased to ~15%DM each. A Principal Component Analysis of the fatty acid compositions of midgut glands and eggs revealed separate clusters for both species. Lipids of C. maenas were characterized more by membrane fatty acids. Fatty acids of C. maenas midgut glands and eggs clustered together. They were largely dominated by carnivory biomarkers. Contrastingly, fatty acids of midgut glands and all eggs of H. sanguineus formed separate clusters and trophic markers indicated a more herbivorous diet. Higher lipid levels and thus more pronounced energy deposition in H. sanguineus midgut glands indicate higher starvation tolerance for females, a potential competitive advantage over C. maenas. Direct food competition, however, seems negligible, as H. sanguineus prefers a more herbivorous diet than C. maenas. Deviating fatty acid compositions among H. sanguineus midgut glands and eggs suggests that this species may represent an income breeder, utilizing energy from both the midgut gland and dietary input. In contrast, most brachyuran crabs are capital breeders, relying exclusively on internal reserves.

Description: The invasive Asian shore crab Hemigrapsus sanguineus and the native European green crab Carcinus maenas share intertidal habitats along European North Atlantic shores and may compete for food. We evaluated the energy-storing capacities of the 2 species and determined their dietary preferences by means of lipid analysis and fatty acid trophic marker indices. Specimens of both sexes and various sizes were sampled in the rocky intertidal of the island of Helgoland (North Sea) in April, June, August, and October 2015. Total lipids of the midgut glands were significantly higher in H. sanguineus than in C. maenas and followed a distinct seasonal cycle in both sexes (ca. 20−50% of dry mass, DM). The lower lipid contents of C. maenas (ca. 20% of DM) remained at a similar level throughout the seasons. The seasonal differences in the females of H. sanguineus may be due to higher reproductive output and, consequently, lipid turnover, but remain unexplained in males. Trophic indices for Bacillariophyceae, Chlorophyta, and especially Phaeophyceae were higher in H. sanguineus than in C. maenas, suggesting a higher degree of herbivory of the invader. In contrast, the Rhodophyta index was higher in C. maenas. Thus, competition for food between the 2 species will probably be low in habitats rich in macroalgae. The ability of H. sanguineus to utilize mainly energy-poor algae but accumulate high-energy reserves may be an advantage for successfully establishing persistent populations in new habitats.

Description: Hemigrapsus sanguineus, the Asian shore crab, is a successful invader at the U.S. east coast and the European Atlantic coast, where populations are established from France to Germany. Numerous studies on behavioral aspects and population development are available, however, physiology and energetic impact of the invader on its ecosystem are virtually unknown. We measured respiration rates of H. sanguineus and its native equivalent, the European shore crab C. maenas, from the island of Helgoland, North Sea, Germany. Measurements were conducted at 5, 10, 15 and 20 °C, using a flow-through system. Specific respiration rates were overall similar for both species. Rates increased with temperature and decreased with the mass of the crab. Mass-standardized data revealed that small H. sanguineus showed significantly higher respiration rates at higher temperatures than C. maenas. This difference vanished with increasing crab size. Statistical modelling was used to develop species-specific equations to calculate a crab´s respiration rate in relation to its mass and the ambient temperature. Taking the overall biomass and temperature into account, the population respiration, carbon uptake and the metabolic energy demand, a measure for the population´s ecosystem impact, was calculated. In the Helgoland intertidal, total biomass of H. sanguineus is now about 59% of that of the C. maenas population but the cumulative metabolic energy demand represents already 78% of the value for C. maenas. With still increasing abundances of H. sanguineus and globally rising temperatures, the ecosystem impact of this invader will increase in the future.

Description: Since 1999, the invasive Asian shore crab Hemigrapsus sanguineus successfully established populations along the Atlantic coastline of Europe, now reaching from the French Channel coast to the German Bight. While various studies on population development and behavior are available, the physiology of H. sanguineus and its energetic impact on the habitat are virtually unknown. We measured the oxygen consumption of H. sanguineus and of its native equivalent, the European shore crab Carcinus maenas, and assessed the energetic ecosystem impact of the invader versus the native species. Respiration rates were determined in a flow-through system at 5, 10, 15 and 20 °C, the crab´s natural temperature range in European habitats. We compared mass-specific respiration rates of mass-standardized crabs of 0.5, 5 and 10 g body mass. Small H. sanguineus specimens consumed significantly more oxygen at higher temperatures than small C. maenas. This difference was less distinct in medium-sized animals and absent in larger animals. We established species-specific models to calculate respiration rates for a crab using the individual´s biomass and the water temperature. These models were applied to the crab populations from the intertidal of the island of Helgoland (Germany) based on data from August 2014. While total population biomass of H. sanguineus was as high as 59% of that of C. maenas, population respiration and energy demand of the invader reached about 78% relative to the value for the native species. As H. sanguineus abundances are still rising, the food uptake and thus the energetic ecosystem impact of this invader will continuously increase in the future.

Description: The physiology of the successful intertidal invader, the Asian shore crab Hemigrapsus sanguineus, is essentially unknown. It is quite abundant from France to German coasts but information on food preferences, energy storage and breeding strategy are still lacking. Hemigrapsus sanguineus and its native ecological equivalent, the European green crab Carcinus maenas, were collected in April, June, August and October 2016 in the intertidal area of the island of Helgoland, North Sea, Germany. Midgut glands were dissected and in case of ovigerous females, additionally the egg masses were sampled. Lipids were extracted and the fatty acid composition was determined by gas chromatography. The lipid contents of H. sanguineus midgut glands was higher and more variable over the seasons than those of C. maenas (20 - 60% vs. ~20%, respectively). Trophic marker fatty acids revealed a more herbivorous diet of H. sanguineus, whereas C. maenas profiles suggested a wide range from more herbivorous to more carnivorous diets. The fatty acid compositions of eggs and midgut glands formed two separate clusters in H. sanguineus but both tissue types were similar in C. maenas. Hemigrapsus sanguineus stored much more energy than C. maenas and may cope better with longer periods of starvation. The invaders were able to build up and maintain these reserves by utilizing mostly low quality algal food. Further, H. sanguineus is an income breeder, utilizing recently eaten food for reproduction, whereas C. maenas is a capital breeder. The physiological properties suggest that the invader is competitively superior over the native crab.

Description: Die Asiatische Felsenkrabbe Hemigrapsus sanguineus wurde erstmals vor Le Havre und in der Oosterschelde entdeckt. Seitdem breitete sie sich entlang der südlichen Nordseeküste bis in das Skagerrak aus. In diesem Lebensraum konkurriert sie mit der einheimischen Strandkrabbe Carcinus maenas. Zahleiche Arbeiten befassten sich mit dem Konkurrenzverhalten und dem Invasionserfolg von H. sanguinaeus. Wir haben im Helgoländer Eulitoral die Speicherlipide der Weibchen beider Arten im Hinblick auf ihre Reproduktionsstrategien untersucht. Eiertragende C. maenas wurden im April und Juni 2015 beprobt. Sie hatten einen Lipidgehalt von ca. 10 % der Trockenmasse (%TM) in der Mitteldarmdrüse und von 15-25 % in den Eiern. Eiertragende H. sanguineus hatten im Juni, August und Oktober jeweils einen Lipidgehalt von 35-45 %TM in den Mitteldarmdrüsen und 15-30%TM in den Eiern. Bei C. maenas war die Fettsäurezusammensetzung der Mitteldarmdrüsen und der Eier sehr ähnlich, was ein Hinweis auf das „capital breeding“-Prinzip ist: Konsumierte Energie wird zunächst in der Mitteldarmdrüse gespeichert und dann einmalig in die Ovarien transferiert. Im Gegensatz dazu unterschieden sich bei H. sanguineus die Fettsäuremuster der Mitteldarmdrüsen und der Eier deutlich voneinander, was auf das „income breeding“- Prinzip hinweist: Konsumierte Lipide werden direkt in die Ovarien transferiert. Dadurch kann diese Art lipidreichere Eier und mehrere Gelege pro Jahr produzieren, was ihren Ausbreitungserfolg steigert.

Description: From the depths of the oceans to the shallow estuaries and wetlands of our coasts, organisms of the marine environment are teeming with unique adaptations to cope with a multitude of varying environmental conditions. With millions of years and a vast volume of water to call their home, they have become quite adept at developing specialized and unique techniques for survival and – given increasing human mediated transport – biological invasions. A growing world human population and a global economy drives the transportation of goods across the oceans and with them invasive species via ballast water and attached to ship hulls. In any given 24-hour period, there are about 10,000 species being transported across different biogeographic regions. If any of them manage to take hold and establish a range in an exotic habitat, the implications for local ecosystems can be costly. This review on marine invasions highlights trends among successful non-indigenous species (NIS), from vectors of transport to ecological and physiological plasticity. Apart from summarizing patterns of successful invasions, it discusses the implications of how successfully established NIS impact the local environment, economy and human health. Finally, it looks to the future and discusses what questions need to be addressed and what models can tell us about what the outlook on future marine invasions is.