Asian Shore Crab Hemigrapsus Sanguineus Research Articles

Variation in metabolic rate during low tide aerial exposure in the Asian shore crab Hemigrapsus sanguineus

Intertidal animals deal with physical gradients daily that create stressful conditions across the shore. These physical gradients influence the physiological performance of organisms, requiring responses that may differ with height on the shore. We examined the respiratory response to aerial exposure in the invasive Asian shore crab Hemigrapsus sanguineus during periods of low tide emersion using two field experiments. The first experiment simultaneously measured respiration of individuals collected from different heights on the shore, which had therefore been emersed for different lengths of time. The second experiment measured respiration of individuals collected at different times from the same tidal height. Respiration rates of crabs in both experiments increased immediately after emersion, nearly doubling by and peaking at ~ 1.5 h of aerial exposure, before decreasing again over the next 1.5 h. These results suggest that the energetic cost of low tide exposure is greatest shortly after emersion during the first half of the typical low tide period, but then decreases thereafter. These respiration patterns facilitate the broad intertidal distribution of this species on rocky shores throughout its range.

Using external morphology as a proxy for stomach size in Hemigrapsus sanguineus.

Stomach morphology can provide insights into an organism's diet. Gut size or length is typically inversely related to diet quality in most taxa, and has been used to assess diet quality in a variety of systems. However, it requires animal sacrifice and time-consuming dissections. Measures of external morphology associated with diet may be a simpler, more cost-effective solution. At the species level, external measures of the progastric region of the carapace in brachyuran crabs can predict stomach size and diet quality, with some suggestion that this approach may also work to examine individual diet preferences and specialization at the individual level; if so, the size of the progastric region could be used to predict trends in diet quality and consumption for individuals, which would streamline diet studies in crabs. Here, we tested whether external progastric region size predicts internal stomach size across latitude and time of year for individuals of the invasive Asian shore crab Hemigrapsus sanguineus. We found that the width of the progastric region increased at a faster rate with body size than stomach width. In addition, the width of the progastric region followed different trends across sites and over time compared to stomach width. Our results therefore suggest that the progastric region may not be used as a proxy for stomach size variation across individuals.

Shift from income breeding to capital breeding with latitude in the invasive Asian shore crab Hemigrapsus sanguineus

Organisms vary in the timing of energy acquisition and use for reproduction. Thus, breeding strategies exist on a continuum, from capital breeding to income breeding. Capital breeders acquire and store energy for breeding before the start of the reproductive season, while income breeders finance reproduction using energy acquired during the reproductive season. Latitude and its associated environmental drivers are expected to heavily influence breeding strategy, potentially leading to latitudinal variation in breeding strategies within a single species. We examined the breeding strategy of the Asian shore crab Hemigrapsus sanguineus at five sites spanning nearly 10° of latitude across its invaded United States range. We hypothesized that the primary breeding strategy of this species would shift from income breeding to capital breeding as latitude increases. We found that though this species’ breeding strategy is dominated by capital breeding throughout much of the range, income breeding increases in importance at lower latitudes. This latitudinal pattern is likely heavily influenced by the duration of the foraging and breeding seasons, which also vary with latitude. We also found that reproductive characteristics at the northern and southern edges of the invaded range were consistent with continued range expansion. We suggest that the reproductive flexibility of the Asian shore crab is a key facilitator of its continued invasion success. Our results highlight the influence of latitude on the breeding strategy of a species and emphasize the need for further research regarding the ecological importance and implications of flexibility in breeding strategies within species.

Investigating the impact of introduced crabs on the distribution and morphology of littorinid snails: Implications for the survival of the snail Littorina saxatilis

Introduced species can have profound impacts on communities by displacing and consuming native organisms. The intertidal communities in the Gulf of Maine have been radically altered through a suite of invasions in the early 1800s, including the herbivorous snail Littorina littorea and the omnivorous European green crab Carcinus maenas leading to morphological and distributional changes in two native gastropod grazers (Littorina saxatilis and Littorina obtusata). Green crabs and native cancrid crabs occupy the mid to low intertidal zones, leaving the high intertidal zones relatively free from crab predation pressure. However, a more recent invasion in the 1980s by the intertidal omnivorous Asian shore crab Hemigrapsus sanguineus, has eliminated this predator-free zone in the high intertidal. This invasion is of particular concern to populations of L. saxatilis, which were relegated to the high intertidal through combined competition and predation pressure by L. littorea and C. maenas. In order to quantify the potential impact of H. sanguineus on native snails, we conducted both field and lab experiments, testing the susceptibility of different sized snails to predation by introduced crabs. Additionally, we measured the distribution, abundance, and morphology of intertidal snails and crabs. Smaller snails of all species were the most susceptible to predation, although susceptibility differed among snail species and predation capabilities differed among crab species. L. saxatilis was the most vulnerable snail species to predation, with 73 and 64% of the population susceptible to large H. sanguineus and small C. maenas, respectively, while >96% of the L. littorea population was resistant to predation. L. saxatilis has been relegated to the high intertidal where there is high abiotic stress and poor-quality food, but until the invasion of H. sanguineus, there was little predation risk. This added predation pressure may further threaten populations of L. saxatilis, especially as the upper intertidal becomes more hostile with increasing temperatures and possible species introductions.

Past energy allocation overwhelms current energy stresses in determining energy allocation trade-offs.

Regeneration of lost appendages is a gradual process in many species, spreading energetic costs of regeneration through time. Energy allocated to the regeneration of lost appendages cannot be used for other purposes and, therefore, commonly elicits energetic trade-offs in biological processes. We used limb loss in the Asian shore crab Hemigrapsus sanguineus to compare the strength of energetic trade-offs resulting from historic limb losses that have been partially regenerated versus current injuries that have not yet been repaired. Consistent with previous studies, we show that limb loss and regeneration results in trade-offs that reduce reproduction, energy storage, and growth. As may be expected, we show that trade-offs in these metrics from historic limb losses far outweigh trade-offs from current limb losses, and correlate directly with the degree of historic limb loss that has been regenerated. As regenerating limbs get closer to their normal size, these historical injuries get harder to detect, despite the continued allocation of additional resources to limb development. Our results demonstrate the importance of and a method for identifying historic appendage losses and of quantifying the amount of regeneration that has already occurred, as opposed to assessing only current injury, to accurately assess the strength of energetic trade-offs in animals recovering from nonlethal injury.

Aliens eating aliens: an introduced amphipod as a potential prey of an invasive rocky shore crab in laboratory experiments

Behavioural interactions between introduced predators and introduced prey are still largely underestimated. The present work takes advantage of the co-occurrence of two introduced species, the Asian shore crab Hemigrapsus sanguineus and the amphipod Ptilohyale littoralis, respectively first recorded on rocky shores along the French coast of the eastern English Channel in 2005 and 2016. In this context, the predation by male and female H. sanguineus on P. littoralis was examined under controlled laboratory conditions, by presenting either juveniles of the blue mussel Mytilus edulis or adult P. littoralis to H. sanguineus. We subsequently assessed the potential prey preference of the Asian shore crab for P. littoralis and M. edulis by presenting the two prey items simultaneously in the same proportion. In the absence of choice, male H. sanguineus preyed significantly more on M. edulis than P. littoralis. In contrast, females preyed significantly less on M. edulis than P. littoralis; however, male and female H. sanguineus consumed similar numbers of P. littoralis. When choice was possible between P. littoralis and M. edulis, the crab did not exhibit preference strictosensu for any type of prey. These results suggest that the Asian shore crab cannot be considered as a naive predator when confronted to a newly introduced prey. Our results also suggest that the amphipod P. littoralis did not exhibit any effective antipredator response towards the crab. These observations nevertheless warrant further work on the effects of abiotic factors (e.g. temperature) as well as other biotic interactions (e.g. presence of other prey or predators for H. sanguineus) may have on the observed prey-predator interactions between H. sanguineus and M. edulis and P. littoralis.

On their way to the north: larval performance of Hemigrapsus sanguineus invasive on the European coast—a comparison with the native European population of Carcinus maenas

The Asian shore crab Hemigrapsus sanguineus has become invasive in North Europe and it co-occurs and competes with the native European shore crab Carcinus maenas. Both species develop through a feeding and dispersive larval phase characterised by several zoeal and a settling megalopa stage. Larvae of marine crabs are vulnerable to food limitation and warming has the potential to exacerbate the negative effects of food limitation on survival and growth. We quantified the combined effects of temperature and food limitation on larval performance (survival and growth) of H. sanguineus and we compared our results with those reported on performance of C. maenas larvae, under the same experimental design and methodology. Larvae from four females of H. sanguineus collected on Helgoland (North Sea) were experimentally reared from hatching to megalopa, at four temperatures (range 15–24 °C) and two food conditions (permanent vs. daily limited access to food). Larval survival of H. sanguineus was low at 15 °C and increased with temperature, in contrast to the high survival reported for C. maenas larvae in the range 15–24 °C. Food limitation reduced survival and body mass of H. sanguineus larvae at all temperatures, but without evidence of the exacerbating effect caused by high temperatures and reported for C. maenas. By contrast, high temperature (24 °C) mitigated the negative effect of food limitation on body mass on H. sanguineus larvae. Advantages of H. sanguineus over C. maenas appear especially under the increased temperatures expected from climate change.

Lack of behavioral effect of surgical mask leachate on the Asian shore crab Hemigrapsus sanguineus: Implications for invasion success in polluted coastal waters

The COVID-19 pandemic generated a new source of plastic mass pollution, i.e. surgical masks, that preferentially accumulate in intertidal environments. Made of polymers, surgical masks are likely to leach additives and impact local intertidal fauna. As typical endpoints of complex developmental and physiological functions, behavioral properties are non-invasive key variables that are particularly studied in ecotoxicological and pharmacological studies, but have, first and foremost, adaptive ecological significance. In an era of ever-growing plastic pollution, this study focused on anxiety behaviors, i.e. startle response, scototaxis (i.e. preference for dark or light areas), thigmotaxis (i.e. preference for moving toward or away from physical barriers), vigilance and level of activity, of the invasive shore crab Hemigrapsus sanguineus in response to leachate from surgical masks. We first showed that in the absence of mask leachates H. sanguineus is characterized by a short startle time, a positive scototaxis, a strong positive thigmotaxis, and an acute vigilance behavior. Specifically, a significantly higher level of activity was observed in white areas, in contrast to the lack of significant differences observed in black areas. Noticeably, the anxiety behaviors of H. sanguineus did not significantly differ after a 6-h exposure to leachate solutions of masks incubated in seawater for 6, 12, 24, 48 and 96 h. In addition, our results were consistently characterized by a high inter-individual variability. This specific feature is discussed as an adaptive behavioral trait, which – through the observed high behavioral flexibility – increases H. sanguineus resilience to contaminant exposures and ultimately contribute to its invasion success in anthropogenically-impacted environments.

Effects of season and latitude on the diet quality of the invasive Asian shore crab Hemigrapsus sanguineus

Invasive species alter invaded ecosystems via direct impacts such as consumption. In turn, an invasive species’ ability to thrive in new habitats depends on its ability to exploit available resources, which may change over time and space. Diet quality and quantity are indicators of a consumer’s consumptive effects and can be strongly influenced by season and latitude. We examined the effects of season and latitude on the diet quality and quantity of the invasive Asian shore crab Hemigrapsus sanguineus throughout a non-winter sampling year at 5 different sites spanning 8° of latitude across its invaded United States range. We found that diet quality, averaged through time, largely follows an expected latitudinal cline, being higher in the center of its range and lower toward the southern and northern edges. We also found that while some sites show similar patterns of diet quality variation with season, no pattern is consistent across all latitudes. Finally, we found that crabs at sites with low diet quality during summer reproductive months did not compensate by increasing total consumption. Because the Asian shore crab is an important consumer in its invaded ecosystems, understanding how its diet quality and quantity vary with season and latitude can help us better understand how this species influences trophic interactions and community structure, how it has been able to establish across a wide ecological and environmental range, and where future range expansion is most likely to occur.

Latitudinal and temporal variation in injury and its impacts in the invasive Asian shore crab Hemigrapsus sanguineus

Nonlethal injury is a pervasive stress on individual animals that can affect large portions of a population at any given time. Yet most studies examine snapshots of injury at a single place and time, making the implicit assumption that the impacts of nonlethal injury are constant. We sampled Asian shore crabs Hemigrapsus sanguineus throughout their invasive North American range and from the spring through fall of 2020. We then documented the prevalence of limb loss over this space and time. We further examined the impacts of limb loss and limb regeneration on food consumption, growth, reproduction, and energy storage. We show that injury differed substantially across sites and was most common towards the southern part of their invaded range on the East Coast of North America. Injury also varied idiosyncratically across sites and through time. It also had strong impacts on individuals via reduced growth and reproduction, despite increased food consumption in injured crabs. Given the high prevalence of nonlethal injury in this species, these negative impacts of injury on individual animals likely scale up to influence population level processes (e.g., population growth), and may be one factor acting against the widespread success of this invader.

Metabolic rates of the Asian shore crab Hemigrapsus sanguineus in air as a function of body size, location, and injury.

Rapid warming in the Gulf of Maine may influence the success or invasiveness of the Asian shore crab, Hemigrapsus sanguineus. To better predict the effects of climate change on this invasive species, it is necessary to measure its energy dynamics under a range of conditions. However, previous research has only focused on the metabolism of this intertidal species in water. We sampled adult crabs from three different sites and measured their metabolic rates in the air. We show that metabolic rate increases with body mass and the number of missing limbs, but decreases with the number of regenerating limbs, possibly reflecting the timing of energy allocation to limb regeneration. Importantly, metabolic rates measured here in the air are ~4× higher than metabolic rates previously measured for this species in water. Our results provide baseline measurements of aerial metabolic rates across body sizes, which may be affected by climate change. With a better understanding of respiration in H. sanguineus, we can make more informed predictions about the combined effects of climate change and invasive species on the northeast coasts of North America.

Cannibalism as a potential factor affecting recruitment of the invasive Asian shore crab Hemigrapsus sanguineus

Cannibalism affects the population dynamics of many marine species, but its potential for influencing population sizes of non-native species is not well understood. A series of laboratory experiments was conducted in 2016 to examine the frequency of cannibalism in the Asian shore crab (Hemigrapsus sanguineus) and the factors influencing its rates of occurrence. Predators and prey varied in size from planktonic megalopae (the last larval stage) and the first benthic crab stage to small juveniles (3–6 mm carapace width, CW), large juveniles (7–10 mm CW), and adult crabs (12–15 mm CW). Individual crabs were paired with groups of smaller prey in glass culture dishes with sand and rocks as potential shelter for the prey, both with and without an alternative food source (commercial crab pellets). Adult and large juvenile crabs consumed megalopae, first-stage crabs and small juveniles, regardless of the presence of food. Megalopae were cannibalized the most heavily, and the size difference between predator and prey was an important factor determining cannibalism rates. Cannibalism of settling megalopae and newly settled juveniles by larger conspecifics could affect recruitment to benthic populations of this non-native crab species.

Ontogeny of osmoregulation of the Asian shore crab Hemigrapsus sanguineus at an invaded site of Europe.

We studied the ontogeny of osmoregulation of the Asian shore crab Hemigrapsus sanguineus at an invaded area in the North Sea. H. sanguineus is native to Japan and China but has successfully invaded the Atlantic coast of North America and Europe. In the invaded areas, H. sanguineus is becoming a keystone species as driver of community structure and the adults compete with the shore crab Carcinus maenas. Strong osmoregulatory abilities may confer the potential to use and invade coastal areas already earlier in the life cycle. We reared larvae and first juveniles at 24°C in seawater from hatching to intermoult of each developmental stage (zoea I-V, megalopa, crab I). We exposed each stage to a range of salinities (0–39 ppt) for 24 h, and then we quantified haemolymph osmolality, using nano-osmometry. In addition, we quantified osmolality in field-collected adults after acclimation to the test salinities for 6 days. Larvae of H. sanguineus were able to hyper-osmoregulate at low salinities (15 and 20 ppt) over the complete larval development, although the capacity was reduced at the zoeal stage V; at higher salinities (25–39 ppt), all larval stages were osmoconformers. The capacity to slightly hypo-regulate at high salinity appeared in the first juvenile. Adults were able to hyper-osmoregulate at low salinities and hypo-regulate at concentrated seawater (39 ppt). H. sanguineus showed a strong capacity to osmoregulate as compared to its native competitor C. maenas, which only hyper-regulates at the first and last larval stages and does not hypo-regulate at the juvenile-adult stages. The capacity of H. sanguineus to osmoregulate over most of the life cycle should underpin the potential to invade empty niches in the coastal zone (characterized by low salinity and high temperatures). Osmoregulation abilities over the whole life cycle also constitute a strong competitive advantage over C. maenas.

Comparing methods for predicting the impacts of invasive species

With the accelerating spread of invasive species around the globe, there is an urgent need to predict where invaders are likely to have the largest impacts and what those impacts will be. Predicting the impacts of individual invasive species a priori has proven to be a problematic endeavor, and several possible methods for doing this have been proposed. In this study we use a model system to compare the different predictive methods that have been proposed to determine which provides the most accurate predictions. Community composition in this system was measured at two separate sites in the Gulf of Maine 20 years ago, just before the arrival of the invasive Asian shore crab Hemigrapsus sanguineus. We repeated the community assessment at each site and determined the overall impacts of this invader over the last two decades. We then predicted the likely impacts based on the historical community composition using 3 methods: the invasion history of this species at sites further south along the mid-Atlantic, scaling up per capita consumption rates that have been measured on individual species in the recipient community, and using mesocosm experiments that used a range of experimental approaches. We show that the invader has had clear, but moderate impacts on the invaded sites since its arrival. We further show that for this omnivorous consumer, predictions based on invasion history and on long term field mesocosm experiments performed much better than predictions based on scaling up from consumption on individual species or based on short term mesocosm experiments conducted in the lab or field. These results highlight potential biases that can be produced by some mesocosm approaches and by scaling up from per capita consumption on individual prey species measured in isolation to predict consumer impacts under more natural conditions.

Risk screening of the potential invasiveness of non-native marine crustacean decapods and barnacles in the Mediterranean Sea

Non-native marine crustaceans can exert detrimental impacts on native marine communities by altering habitat and ecosystem function. The Mediterranean Sea is particularly vulnerable to introductions of non-native crustaceans, as evidenced by their remarkably high establishment success. In this study, 20 species of non-native marine crustacean decapods and barnacles of which eleven extant and nine ‘horizon’ were screened for their potential invasiveness in the Mediterranean Sea. Using the Aquatic Species Invasiveness Screening Kit and including an additional nine native species to increase accuracy, calibrated risk thresholds of 3.5 for the BRA (Basic Risk Assessment) and 8.5 for the BRA + CCA (BRA + Climate Change Assessment) were obtained that distinguished reliably between invasive and non-invasive species. All 20 non-native species were classified as carrying a high risk of invasiveness for the Mediterranean Sea, both for the BRA and the BRA + CCA. Chinese mitten crab Eriocheir sinensis was by far the highest risk species, followed by Harris mud crab Rhithropanopeus harrisii, Asian shore crab Hemigrapsus sanguineus, bay barnacle, Amphibalanus improvisus, and lesser swimming crab Charybdis (Goniohellenus) longicollis. The findings of this study will provide management and control directions for non-native marine crustaceans in the Mediterranean Sea, with special emphasis on regulations regarding ballast waters, which represent one of the main introduction pathways for these aquatic organisms.

Mechanisms of possible self-limitation in the invasive Asian shore crab Hemigrapsus sanguineus

Population sizes of invasive species are commonly characterized by boom-bust dynamics, and self-limitation via resource depletion is posited as one factor leading to these boom-bust changes in population size. Yet, while this phenomenon is well-documented in plants, few studies have demonstrated that self-limitation is possible for invasive animal species, especially those that are mobile. Here we examined the invasive Asian shore crab Hemigrapsus sanguineus, a species that reached very high abundances throughout invaded regions of North America, but has recently declined in many of these same regions. We examined the relationship between diet, energy storage, reproduction, and growth in crabs collected from the New Hampshire coast. We show that energy storage and reproduction both increase with diet quality, while growth declines with diet quality. These results suggest that self-limitation may be a contributing factor to the recent declines of H. sanguineus at sites where this invader was once much more abundant. Further, these results suggest a diet-associated tradeoff in energy allocation to different vital rates, with a focus on reproduction when high quality resources are consumed, and a focus instead on growth when poor quality resources are consumed.

Startle response of the invasive Asian shore crab under different environmental conditions

In recent years, researchers have recognized the importance of behavioral factors for invasive species success. There is evidence that an organism’s ability to quickly react to stimuli may be one behavioral factor leading to invasive success. In this study, we analyzed the startle response of the invasive Asian shore crab Hemigrapsus sanguineus (De Haan, 1835), which measured here as a binary response variable of either active or inactive behavior when the rock they were under was suddenly lifted up. The behavior of H. sanguineus was compared to other crab species in the area to determine how species-dependent their responses were. This response was compared across an intertidal gradient with variation in temperature, intertidal elevation, substrate, time of day, and male-to-female ratios to determine if these affected the crabs’ responses. We found that H. sanguineus was more responsive than other species, perhaps indicative of a generally high activity level for this species. Surprisingly, we found that their behavior was not affected by any environmental factors tested. The consistently active behavior of this crab could be one factor contributing to its high level of success as an invasive species.

Megalopae of the invasive crab Hemigrapsus sanguineus metamorphose in response to marine biofilms from different potential habitats

Habitat characteristics favoring settlement and metamorphosis of larvae can affect the distribution and abundance of benthic marine invertebrates. Less selective responses to habitat cues by settling larvae can lead to broader distributions of species and potentially successful introduction to new geographic locations. This study tested whether marine biofilms, which serve as habitat cues for many invertebrates, enhance settlement and metamorphosis (molting) of megalopae of the invasive Asian shore crab Hemigrapsus sanguineus. Biofilms were allowed to develop on plastic mesh netting deployed in coastal locations with abundant rocky substrate (adult habitat) and estuarine locations with few rocks. Molting rates of laboratory-reared megalopae maintained with biofilm-covered netting were determined. Controls consisted of mesh netting without field-derived biofilms and seawater without netting. Experiments were also conducted with megalopae of a native species of estuarine mud crab, Dyspanopeus sayi, for comparison of responses. Both species molted faster in response to field biofilms regardless of habitat of biofilm development, although the response to biofilms compared to netting without biofilms was stronger for H. sanguineus than D. sayi. The broad response of megalopae of both species to biofilms suggests that positive responses to biofilms and physical structure could be important for successful colonization of crab species in general.

Invasion dynamics: interactions between the European Green Crab Carcinus maenas and the Asian Shore Crab Hemigrapsus sanguineus

As ecosystems are increasingly inhabited by multiple invasive species, interactions among exotic species have the potential to negatively impact native communities and habitats, often in counterintuitive ways. In rocky intertidal habitats along southern New England, the recently introduced Asian Shore Crab, Hemigrapsus sanguineus, has been implicated in the displacement of the more established invasive European Green Crab, Carcinus maenas. We coupled faunal sampling with two field experiments to examine whether H. sanguineus is also displacing C. maenas in the Gulf of Maine. First, we investigated natural recruitment of the two species across an elevation gradient, using cage exclusion units to manipulate predator access. Excluding H. sanguineus adults significantly increased the density of C. maenas recruits, but not conspecific recruit density. Next, to simulate the transition that occurred from pre-invasion to the establishment of H. sanguineus, we conducted a predator inclusion experiment manipulating the density of adult H. sanguineus to examine the effects on C. maenas and conspecific recruit densities. In contrast to the previous experiment, we found that the density of H. sanguineus included in cages negatively affected conspecific recruit density but had no effect on C. maenas recruit density. We hypothesize that juxtaposing results from the two experiments may be a consequence of the considerably greater abundance of C. maenas recruits relative to H. sanguineus recruits during the first experiment. Our work highlights the importance of considering the multitude of factors that influence competitive and intraguild predation interactions, including density-dependent effects, ontogeny and life history characteristics.

Seasonal lipid storage and dietary preferences of native European vs. invasive Asian shore crabs

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.