Effect of immersion at low tide on distribution and movement of the mud snail, Ilyanassa obsoleta (Say), in the upper Bay of Fundy, eastern Canada

Habitat heterogeneity often affects movement behaviours of animals, and consequently their spatial distribution. We evaluated the effect of immersion at low tide on the distribution, fine-scale movement patterns and daily movement patterns of the mud snail Ilyanassa obsoleta on a mudflat in the upper Bay of Fundy, Canada. Mud snails migrate onto intertidal mudflats in the summer, and our field survey showed that their density was higher inside tide pools relative to adjacent areas that are exposed at low tide. Using time-lapse videography, we evaluated the effect of snail size, snail density, and immersion at low tide on fine-scale movement patterns of I. obsoleta. Time until snails stopped moving and burrowed was unaffected by snail size, but snails at low and high densities burrowed somewhat faster than those at intermediate densities. Snail size and snail density had no detectable effect on displacement speed or linearity of displacement. Immersion affected snail movement: snails within tide pools delayed burrowing and traveled in more convoluted paths compared to those on exposed mud. Snails increased their turning angles within tide pools, which is probably the mechanism by which aggregations are formed. We also performed a mark-recapture experiment to compare daily movement patterns of snails released inside and outside tide pools. Snails released in tide pools moved shorter distances, but did not orient themselves differently than snails released outside tide pools. Both groups exhibited significant directionality, moving against the mean water current direction over 24 h. In sum, immersion at low tide affected the behaviour and spatial distribution of snails, resulting in snail aggregations within tide pools. These snail aggregations, in turn, may be a major factor influencing spatial dynamics on mudflats, including causing changes in distribution patterns of the burrowing amphipod Corophium volutator, a dominant inhabitant and key species in the food web of mudflats.     

Effect of the snail Ilyanassa obsoleta (Say) on dynamics of the amphipod Corophium volutator (Pallas) on an intertidal mudflat

We investigated interactions between two dominant invertebrate species of intertidal soft-sediment environments of the northwest Atlantic, the mud snail Ilyanassa obsoleta and the burrowing amphipod Corophium volutator, on a mudflat of the upper Bay of Fundy, Canada. Distribution of I. obsoleta on the mudflat was highly patchy and negatively correlated with density of C. volutator. Manipulation of snail density in cages showed that I. obsoleta influences C. volutator; specifically, increasing density of snails reduced density, increased patchiness in distribution, decreased recapture rates and decreased immigration of C. volutator. Ilyanassa obsoleta seems to be affecting C. volutator through an influence on survival rate and emigration rate, although temporal variation in these effects was observed. Given that both I. obsoleta and C. volutator show a preference for tide pools, an important microhabitat on mudflats, snails might have a profound impact on C. volutator population dynamics.     

Behavioral Response of Corophium volutator to Shorebird Predation in the Upper Bay of Fundy,Canada

Predator avoidance is an important component of predator-prey relationships and can affect prey availability for foraging animals. Each summer, the burrow-dwelling amphipod Corophium volutator is heavily preyed upon by Semipalmated Sandpipers (Calidris pusilla) on mudflats in the upper Bay of Fundy, Canada. We conducted three complementary studies to determine if adult C. volutator exhibit predator avoidance behavior in the presence of sandpipers. In a field experiment, we monitored vertical distribution of C. volutator adults in bird exclosures and adjacent control plots before sandpipers arrived and during their stopover. We also made polymer resin casts of C. volutator burrows in the field throughout the summer. Finally, we simulated shorebird pecking in a lab experiment and observed C. volutator behavior in their burrows. C. volutator adults were generally distributed deeper in the sediment later in the summer (after sandpipers arrived). In August, this response was detectably stronger in areas exposed to bird predation than in bird exclosures. During peak predator abundance, many C. volutator adults were beyond the reach of feeding sandpipers (>1.5 cm deep). However, burrow depth did not change significantly throughout the summer. Detailed behavioral observations indicated that C. volutator spent more time at the bottom of their burrow when exposed to a simulated predator compared to controls. This observed redistribution suggests that C. volutator adults move deeper into their burrows as an anti-predator response to the presence of sandpipers. This work has implications for predators that feed on burrow-dwelling invertebrates in soft-sediment ecosystems, as density may not accurately estimate prey availability.     

Effects of Ilyanassa obsoleta (Say) on the abundance and vertical distribution of Corophium volutator (Pallas) on mudflats of the upper Bay of Fundy

Invertebrates in soft-bottom marine communities exhibit a range of responses to predators and competitors, including both emigration and avoidance. Corophium volutator (Pallas), a burrowing amphipod abundant in the upper Bay of Fundy, Canada, frequently interacts with the eastern mudsnail, Ilyanassa obsoleta (Say), as both species utilize similar habitats and food resources. Typically these interactions result in negative effects on C. volutator. There is evidence that Corophium species exhibit avoidance behaviour when interacting with other species, but whether such a response occurs in the presence of I. obsoleta is unknown. To investigate C. volutator response to I. obsoleta, total C. volutator abundance and vertical distribution within the sediment was examined in a field experiment involving I. obsoleta enclosures and exclosures at two mudflats in the upper Bay of Fundy. I. obsoleta densities were manipulated between June and August 2008 and their effects on C. volutator abundance and vertical distribution observed. Results varied between mudflats and C. volutator size classes. At Peck's Cove, total C. volutator abundance was largely unaffected by I. obsoleta. No size class exhibited a distribution shift in response to I. obsoleta, though both mid-sized individuals and adults occupied deeper layers of sediment late in the summer. Abundance of C. volutator in all size classes at Grande Anse was reduced in July in direct proportion to exposure to I. obsoleta, and adults occupied higher sediment layers in July when I. obsoleta were excluded, suggesting that I. obsoleta curtailed use of these upper layers. Juvenile abundance rebounded in August, possibly due to the mass dispersal of a newly produced juvenile cohort. Adults were reduced in all treatments in August, probably as a result of normal seasonal mortality. Differences in C. volutator responses between mudflats highlight the site-specific nature of such interactions.     

Preying at the edge of the sea: the nemertine Tetrastemma melanocephalum and its amphipod prey on high intertidal sandflats

In the European Wadden Sea, the nemertine Tetrastemma melanocephalumoccurs together with its prey, the amphipod Corophium arenarium, in the upper intertidal zone. T. melanocephalumleaves the sediment when the tide has receded and captures C. arenarium in its U-shaped burrow. Highest abundances of T. melanocephalumon the sediment surface were found on summer evenings, 2–4 h after high tide, when just a thin film of water was left on the flats. Laboratory Y-maze experiments indicated that gradients of substances produced by C. arenarium in this film of water play a role in tracking the prey. In the field, T. melanocephalum appeared in significantly higher numbers on experimental high density patches of C. arenarium. The amphipod in turn is able to recognize the nemertine. In aquarium experiments, significantly more amphipods escaped from the sediment into the water column when the predator was present. In the field, both predator and prey showed a high mobility by drifting in tidal waters. Benthic abundance maxima of T. melanocephalum and C. arenariumusually did not coincide spatially. It is assumed that the nemertines avoid tidal flats that dry out quickly leaving too little time for prey capture. T. melanocephalum is not able to dig into the sediment, but lives in burrows of Nereis diversicolor. The abundance of this polychaete was inversely related to C. arenarium, presenting a dilemma for T. melanocephalum: the spatial overlap of food and accommodation was rather small.     

Effect of predatory fish presence on habitat use and diel movement of the stream amphipod, Gammarus minus

SUMMARY.

• 1 We examined how the presence of predatory fish affected macro—and microhabitat use and movement patterns of the amphipod, Gammarus minus Say, in a second order stream.
• 2 Among macrohabitats, amphipods were consistently more abundant in runs than in pools. Densities in pools were low regardless of fish presence, whereas low densities in runs were correlated with fish presence. Siltation and the lack of coarse substrate particles probably accounted for the scarcity of G. minus in pools. Among microhabitats, G. minus was more abundant in leaf litter and gravel than in silt/sand substrates. Fish presence did not affect microhabitat use in the field. Laboratory manipulations showed, however, that predation rates by green sunfish (Lepomis cyanellus Rafinesque) were significantly lower on amphipods in leaf litter than in other microhabitats.
• 3 Most movement by G. minus in runs occurred at night and was directed downstream. In the laboratory, amphipods significantly reduced swimming activity at night in response to water conditioned with green sunfish. In the field, however, amphipods were only slightly less active in runs at night when exposed to chemical cues of confined green sunfish. The distribution of amphipods in streams is influenced by the presence of fish, but chemical cues alone may not be important in triggering predator avoidance behaviours in nature.

     

Zur Nahrungsaufnahme vonTetrastemma melanocephalum (Nemertini)

The proboscis of hoplonemerteans is armed with a stylet apparatus, which is used for capturing prey.Tetrastemma melanocephalum (Johnston) is a common hoplonemertean in the littoral of the Baltic Sea and the North Sea. It lives in the phytal and on sand- and mudflats and occurs in large numbers in theCorophium belts in the waddenseas.T. melanocephalum feeds on arthropods, mainly on copepods and amphipods. The prey is caught with the proboscis, penetrated by the stylet and grows weak (reduction of movements) within a few seconds; it is then sucked out. During the summer, a 15–35 mm long nemertean captures about 3 specimens ofCorophium volutator of 3.5–6 mm length per day. In theCorophium belt at Sahlenburg, German Bight, North Sea, 29 nemerteans/0.25m² were found; they feed mainly onC. volutator, so that on average more than 10 000 specimens ofC. volutator/m²/month are sucked out by these nemerteans. Hence,T. melanocephalum is an important consumer ofCorophium.     

<Emphasis Type="Italic">Spartina maritima</Emphasis> influence on the dynamics of the phosphorus sedimentary cycle in a warm temperate estuary (Mondego estuary,Portugal)

During the last decades the Mondego estuary has been under severe ecological stress mainly caused by eutrophication. In this salt march system, Spartina maritima covers about 10.5 ha of the intertidal areas. The objective of the present study was to evaluate the effect of Spartina maritima marshes on the dynamics of phosphorus (P) binding in the surface sediment. We compare phosphate and oxygen fluxes, P-adsorption capacity, phosphate concentrations and total amount, and the extractable P forms in the upper 20 cm of sediment in vegetated sediment with adjacent mudflats without vegetation. Sediment pore-water profiles followed a clear trend, with lower P concentrations in more superficial layers, and increasing with depth. The vegetated mudflats presented lower concentrations of dissolved inorganic phosphorus than adjacent bare bottom mudflats, lower phosphate total amount, as well as higher P-adsorption capacity. Results from the extraction procedure show that the superficial layers are the most important for estuarine phosphorus dynamics, since maximum concentrations of labile P pools are present here. In contrast, higher proportions of refractory P pool are found in deeper layers. Spartina marsh sediments had less total P, less iron bound P, and less exchangeable P than adjacent bare bottom mudflats. Also the pool of loosely sorbed P is lower in the Spartina marsh. Phosphate regeneration from the sediment to the overlying water was only 11.8 kg ha⁻¹ year⁻¹ in vegetated sediment while 25.8 kg ha⁻¹ year⁻¹ in the bare mud flat. Plant uptake for growth combined with an enhanced P-adsorption capacity of the sediment, may explain these differences. Therefore, Spartina marshes are very important agents in the sedimentary P cycle worldwide, and can be considered a useful management tool in estuarine ecosystem recovery efforts.     

Lifestyle of Korean mudskipper <Emphasis Type="Italic">Periophthalmus magnuspinnatus</Emphasis> with reference to a congeneric species <Emphasis Type="Italic">Periophthalmus modestus</Emphasis>

To investigate the life history and ecology of the mudskipper Periophthalmus magnuspinnatus, observations and collection were made on coastal mudflats in southern Korea. Periophthalmus magnuspinnatus was active from May to September on the mudflats, exclusively occupying rough and elevated or sloped mudflats of the seashore or the stream mouth, usually vegetated with halophilous grasses. The congeneric species, P. modestus, mainly occurred on extensive low-elevation and level mudflats with no visible vegetation. An apparent alternation of habitat use by P. modestus took place on mudflats at the stream mouth in mid-October, when P. magnuspinnatus began wintering in its burrow and P. modestus came onto the vacated mudflats to construct burrows for wintering. The active season for P. magnuspinnatus at 17°C or higher air temperature was a little shorter than that of P. modestus. Periophthalmus magnuspinnatus constructed a burrow for the entire season in the highest area of the intertidal mudflat, where they hid themselves during high tide or when frightened, whereas P. modestus were likely to use any burrow constructed by other animals or sunken places to hide. The main stomach contents of P. magnuspinnatus were crabs and gammarids. During the reproductive season from May to July, P. magnuspinnatus performed mating behaviors and constructed a spawning burrow similar to the ones known for P. modestus, except their body color turned dark and quivering body movements were observed in the mature male instead of a pink or orange body color and wiggling body movements as in P. modestus. Eggs, measuring 1.56–1.69 mm in major axis and 0.94–1.0 mm in minor axis, were laid on the ceiling and the side wall of the “J”-shaped spawning room generally known for Periophthalmus species. Young of both species started to occur on the mudflat in June.     

Functional response of staging semipalmated sandpipers feeding on burrowing amphipods

Despite its fundamental relevance to many ecological processes in predator–prey relationships, the functional response, which relates predator intake rate to prey density, remains difficult to document in the field. Here, I document the functional response of semipalmated sandpipers (Calidris pusilla) foraging on a burrowing amphipod Corophium volutator during three field seasons at the peak of fall migration in the upper Bay of Fundy (New Brunswick, Canada). I gathered data during the ebbing tide when all sandpipers are highly motivated to feed after a lengthy hide-tide fast. As birds follow the receding tideline, foragers encounter prey at different densities and do not aggregate in the richest food patches. Results show that intake rate increased at a decreasing rate with Corophium density, yielding a type II functional response typical of many shorebird species. Intake rate decreased in the later stages of migration stopover at a time where preferred prey items have been shown to occur at lower densities due to prior depletion. At this period of lower prey availability, intake rate also decreased with sandpiper density providing evidence for interference at low prey density. The results illustrate the fact that the functional response may not be unique but instead vary as a function of the type of competitive relationship among foragers.     

Mudflat biota since the 1930s: change beyond return?

Where, since the 1980s, patchy and variable green algal mats are prevailing, distinct belts of an amphipod (Corophium volutator) and seagrass (Zostera spp.) had dominated in the 1930s. The zonation between tide marks has been mapped in a sheltered sedimentary bay in the Wadden Sea near the island of Sylt (coastal eastern North Sea). Maps on vegetation from 1924 and on selected macrobenthos from 1932 and 1934 are compared with biannual surveys conducted from 1988 to 2006. Rising high water levels and eutrophication are suggested to be major causes of the observed long-term changes. In front of a saltmarsh, a sandy beach developed and partly displaced former cyanobacterial mats. Advancing sandiness may have inhibited C. volutator and facilitated lugworms, Arenicola marina, in the upper tidal zone. A variable occurrence of green algal mats arising in the 1980s affected infauna and seagrass by smothering the biota underneath. This dissolved a coherent belt of Zostera noltii. In the lower tidal zone, natural disturbances had lasting effects on the occurrence of mussels with attached fucoid algae. The spectrum of species became enriched by alien species (13% of macrobenthic taxa). A reversal to habitat structure and biotic zonation of the 1920–1930s does not seem possible. Aliens, in combination with climate change, are expected to further divert the ecological pattern to new configurations.     

Anthropocene invasion of an ecosystem engineer: resolving the history of Corophium volutator (Amphipoda: Corophiidae) in the North Atlantic

Resolving the natural histories of species is important for the interpretation of ecological patterns, as it provides evolutionary context for the interactions between organisms and their environment. Despite playing an integral role on the intertidal mudflats of the North Atlantic as an abundant food source for predators and as an ecosystem engineer that alters the soft sediment environment, no previous studies have provided empirical evidence to determine the biogeographical origin of the amphipod Corophium volutator. To resolve its status as introduced or indigenous in Europe and North America, we analyzed sequence data for two mitochondrial loci and two nuclear markers, aiming to determine whether the present range of C. volutator is the result of unresolved taxonomy, persistence in glacial refugia, natural trans‐Atlantic dispersal, or human‐mediated introduction. Our results demonstrate a reduced genetic diversity in North American populations that is a subsample of diversity in European populations, with coalescent analyses of mitochondrial and nuclear DNA supporting different models of multiple introductions from Europe to the Bay of Fundy and Gulf of Maine in North America. These results suggest that C. volutator was introduced to North America prior to the first surveys of local biota in the 20th Century, which has broad implications for interpretations of community and ecosystem interactions in the North Atlantic intertidal. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 115 , 288–304.     

Populations and production of bacteria in the plankton of two shallow lakes

The troglobitic amphipod crustacean Crangonyx antennatus populates two distinct habitats in southern Appalachian caves, mud-bottom pools and gravel-bottom streams. An investigation was conducted to determine if stream populations possess behavioral adaptations to lotic conditions. Observations on the thigmotactic behavior of C. antennatus were made seasonally in six Lee Co., Virginia caves, three containing pool habitats and three with stream habitats. Stream-dwelling amphipods were found to exhibit more cryptic behavior than those inhabiting pools under natural conditions and when disturbed. Artificial stream experiments indicate that stream-dwelling C. antennatus are better adapted behaviorally to lotic conditions than pool-dwelling amphipods.     

The feeding strategy of the dunlin (Calidris alpina L.) in artificial and non-artificial habitats at Ria de Aveiro,Portugal

Usually waders (Aves, Charadrii) feed in the inter-tidal zone over the low tide period and roost over high tide. But occasionally when their energetic needs increase, due to adverse weather conditions or to a need to store fat to fuel migration, the birds also feed during high tide, sometimes in agricultural land. At the Ria de Aveiro, about 44% (corresponding to 4500 birds) of the wintering dunlin (Calidris alpina) use artificial salt pan habitats throughout the winter whilst the others utilize natural intertidal habitats, mainly mudflats. The dunlin that feed intertidally in natural areas, such as the Canal de Mira, do not feed over high tide. In contrast, the birds that use the salt pan area feed both during low tide, in abandoned salt pans whose walls are breached and thus allow access to the tide, and high tide, in active salt pans, and keep on doing so throughout the winter. The energetic needs of both groups of birds are thought to be similar. However, the benthos of the abandoned salt pans seem to be different, and poorer in energetic terms, from those of the natural mudflats. It is argued that dunlin wintering in the salt pan area must compensate for this by feeding over high tide in active salt pans. Rather than just being a means of obtaining more energy at certain times of high energy demand, it would seem that feeding in active salt pans over high tide should be considered as a normal component of the feeding strategy of these birds, a strategy that is different from those wintering in the natural mudflats. These findings highlight the importance of the salt exploitation industry to waders and also show that artificial habitats can support large numbers of wintering waders.     

Predation rates of nemertean predators: the case of a rocky shore hoplonemertean feeding on amphipods

Estimates of the predation rates of benthic nemerteans are often based on observations of single individuals, and consequently they may not be representative for all members of a population of these predators. Herein we conducted controlled and repeatable laboratory experiments on the predation rate of the hoplonemertean Amphiporus nelsoni Sánchez 1973, which is common at exposed rocky shores along the central Chilean coast. During the austral fall (April, May 2000), nemerteans were observed in relatively high numbers crawling in the intertidal zone during early morning or late-afternoon low tides. When these nemerteans were offered living amphipods held by a forceps, they immediately attacked the amphipods and fed on them. In the laboratory experiments, nemerteans preferred the amphipod Hyale maroubrae Stebbing, 1899, which is also very common in the natural habitat of A. nelsoni. The nemerteans preyed to a higher extent on small males and non-ovigerous females than would have been expected from their abundance. We suggest that these (non-reproductive) stages of H. maroubrae are very mobile and therefore have a high likelihood of encounters with nemerteans. Predation rates reached maxima when nemerteans were provided prey densities of four or more of their preferred prey species, H. maroubrae, furthermore indicating that encounter rates with prey may affect predation rates. In long-term laboratory experiments, A. nelsoni consumed more amphipods during low tide conditions than during high tide conditions. Many nemerteans in the field prefer particular environmental conditions (e.g. nocturnal low tides), which restricts the time available for successful feeding. In the long-term experiment, predation rates of A. nelsoni never exceeded 0.5 amphipods nemertean^–1 d^–1. Maximum feeding events were 3 or 4 amphipods nemertean^–1 d^–1, but this only occurred during 10 out of a possible 2634 occasions. Nemerteans that had consumed 3 or 4 amphipods during 1 day, consumed substantially less prey during the following days. Towards the end of the long-term experiment, average predation rates decreased to 0.2 amphipods nemertean^–1 d^–1, corresponding to predation rates reported for other nemertean species (0.1–0.3 prey items nemertean^–1 d^–1). We suggest that predation rates from laboratory experiments represent maximum estimates that may not be directly transferable to field populations. Additionally, low predator–prey encounter rates with preferred prey in the field may further limit the predation impact of nemertean predators in natural habitats.     

The contrasting diet of Harpagifer antarcticus (Notothenioidei, Harpagiferidae) at two localities of the South Shetland Islands, Antarctica

The diet of Harpagifer antarcticus was studied at two localities of the South Shetland Islands, Antarctica. The analysis of the stomach contents of specimens collected in tide pools at Potter Cove, King George Island, indicated that gammarid amphipods (mainly Gondogeneia sp.) were the main prey of this fish, followed by polychaetes, gastropods and isopods. By contrast, the specimens from Harmony Point, Nelson Island, which were recovered from stomach contents of Antarctic shags Phalacrocorax bransfieldensis feeding at depths of 46–110 m, preyed almost exclusively on the Antarctic krill Euphausia superba. These results are discussed and compared with previous studies. Received: 27 August 1997 / Accepted: 17 November 1997     

Laboratory Studies of Parasite Transmission Aspects between Hydrobia spp. (Gastropoda) and Corophium volutator (Amphipoda)

Aspects of parasite transmission between Hydrobia spp. and Corophium volutator, first and second intermediate host of digenetic trematodes, were investigated under laboratory conditions. H. ventrosa is used as an intermediate host by several trematode species. Under laboratory conditions the most frequently observed emergence from H. ventrosa was of cercariae of Maritrema subdolum. The number of cercariae shed per day varied considerably. It was observed that 30 cercariae on average and up to 450 cercariae at maximum can emerge from a single H. ventrosa per day. Cercarial production continued until the death of the snails. The life-span of cercariae of the species M. subdolum decreased as the water temperature increased. It can be concluded that under natural conditions the cercariae, after emerging, have a maximum period of 1 day in which to seek out their second intermediate host C. volutator. Almost all specimens of C. volutator exposed to cercariae of the species M. subdolum died within the test period of 6 days. High average cercarial densities caused short life-spans (<50 h), while at lower densities longer survival times were possible. For cercariae of other Trematoda species, we were unable to find any equally clear evidence of a reduction in the survival rate of C. volutator within the test period. Mortality of C volutator, and other effects of infestation, as observed in our experiment, can be assumed to be a result of the penetration process of the cercariae but they also can be attributed to the mesocercariae.     

Competition in foraging flocks of migrating semipalmated sandpipers

I examined the effect of competitor density on foraging success in staging semipalmated sandpipers (Calidris pusilla) foraging on a burrowing amphipod (Corophium volutator) in each of two study years. Little is known about the effect of competitor density when predation attempts disturb prey, causing a temporary decrease in food availability. Controlling for Corophium density and other potentially confounding factors such as temperature, pecking rate and capture rate increased linearly with sandpiper density. Success rate, the ratio of captures to pecks, was not influenced by sandpiper density. The effect of sandpiper density was similar in each of the two study years and was documented early and late in the low tide period. The increase in foraging rate is argued to be a response to increased competition for rapidly depleting prey at the temporal scale of exploitation by a flock. Potential fitness costs associated with higher foraging costs may include decreased ability to distinguish between the profitability of different prey and reduced vigilance against predators.     

INVERTEBRATE‐MEDIATED NUTRIENT LOADING INCREASES GROWTH OF AN INTERTIDAL MACROALGA1

Even in nitrogen‐replete ecosystems, microhabitats exist where local‐scale nutrient limitation occurs. For example, coastal waters of the northeastern Pacific Ocean are characterized by high nitrate concentrations associated with upwelling. However, macroalgae living in high‐zone tide pools on adjacent rocky shores are isolated from this upwelled nitrate for extended periods of time, leading to nutrient limitation. When high‐intertidal pools are isolated during low tide, invertebrate‐excreted ammonium accumulates, providing a potential nitrogen source for macroalgae. I quantified the influence of mussels (Mytilus californianus Conrad) on ammonium accumulation rates in tide pools. I then evaluated the effects of ammonium loading by mussels on nitrogen assimilation and growth rates of Odonthalia floccosa (Esp.) Falkenb., a common red algal inhabitant of pools on northeastern Pacific rocky shores. Odonthalia was grown in artificial tide pool mesocosms in the presence and absence of mussels. Mesocosms were subjected to a simulated tidal cycle mimicking emersion and immersion patterns of high‐intertidal pools on the central Oregon coast. In the presence of mussels, ammonium accumulated more quickly in the mesocosms, resulting in increased rates of nitrogen assimilation into algal tissues. These increased nitrogen assimilation rates were primarily associated with higher growth rates. In mesocosms containing mussels, Odonthalia individuals added 41% more biomass than in mesocosms without mussels. This direct positive effect of mussels on macroalgal biomass represents an often overlooked interaction between macroalgae and invertebrates. In nutrient‐limited microhabitats, such as high‐intertidal pools, invertebrate‐excreted ammonium is likely an important local‐scale contributor to macroalgal productivity.     

Simulating control strategies for a spatially structured weed invasion: <Emphasis Type="Italic">Spartina alterniflora</Emphasis> (Loisel) in Pacific Coast estuaries

Patterns in the spatial arrangement of invasive plant populations can provide opportunity for strategic placement of control efforts. Smooth cordgrass (Spartina alterniflora) is rapidly invading the intertidal mudflats of Pacific Coast estuaries. Its pattern of spread is distinctive. Seedlings establish in open mud and then grow vegetatively to form expanding circular patches, which dot the mudflats and eventually coalesce into a contiguous monospecific meadow. The invasion typically begins in the upper tide zone and then moves down the tidal gradient. A spatially explicit model was used to simulate the spread of S. alterniflora and compare various strategies for control in a situation where only a fraction of the total infestation could be controlled each year. A strategy of killing outlying patches first and then attacking the dense meadows (moving up the tidal gradient) led to eradication with up to 44% less time and effort than a strategy of attacking the dense meadows first and outlying patches second (moving down the tidal gradient). In the control of contiguous meadows located adjacent to the shoreline, the best strategy was to approach one end of the infestation, moving across the meadow to the other end. Suppression of seeds was not an effective control strategy. In general, effective control strategies were those that first eliminate the plant in areas where current or future vegetative growth is greatest. Application of these results in control programs for S. alterniflora and similar invasive species could greatly reduce the costs of control work and improve the likelihood of local or complete eradication.