Fine‐scale diet of the Australian sea lion (Neophoca cinerea) using DNA‐based analysis of faeces

We applied DNA‐based faecal analysis to determine the diet of female Australian sea lions (n = 12) from two breeding colonies in South Australia. DNA dietary components of fish and cephalopods were amplified using the polymerase chain reaction and mitochondrial DNA primers targeting the short (~100 base pair) section of the 16S gene region. Prey diversity was determined by sequencing ~50 amplicons generated from clone libraries developed for each individual. Faecal DNA was also combined and cloned from multiple individuals at each colony and fish diversity determined. Diets varied between individuals and sites. Overall, DNA analysis identified a broad diversity of prey comprising 23 fish and five cephalopod taxa, including many species not previously described as prey of the Australian sea lion. Labridae (wrasse), Monacanthidae (leatherjackets) and Mullidae (goat fish) were important fish prey taxa. Commonly identified cephalopods were Octopodidae (octopus), Loliginidae (calamary squid) and Sepiidae (cuttlefish). Comparisons of fish prey diversity determined by pooling faecal DNA from several samples provided a reasonable but incomplete resemblance (55–71%) to the total fish diversity identified across individual diets at each site. Interpretation of diet based on the recovery of prey hard‐parts identified one cephalopod beak (Octopus sp.) and one fish otolith (Parapriacanthus elongatus). The present study highlights the value of DNA‐based analyses and their capabilities to enhance information of trophic interactions.     

Diets of Steller sea lions off the coast of Hokkaido,Japan: An inter‐decadal and geographic comparison

Inter‐decadal and geographic variations in the diets of Steller sea lion, Eumetopias jubatus, were examined based on the contents of 408 stomachs collected from coastal areas around Hokkaido Island during the periods 1994–1998 and 2005–2012. The most important prey species in the 1990s were gadid fishes (walleye pollock [Gadus chalcogrammus], Pacific cod [Gadus microcephalus] and saffron cod [Eleginus gracilis]). The frequency of occurrence and gravimetric contribution of gadids decreased in the 2000s latter period at three study sites (Rausu, Shakotan and Rebun) and were replaced by Okhotsk Atka mackerel (Pleurogrammus azonus) and smooth lumpsucker (Aptocyclus ventricosus). However, analysis based on gravimetric composition indicated that the dietary diversity of prey showed only a slight inter‐decadal difference, reflecting the wide diversity of prey ingested during both study periods. These results indicate that Steller sea lions along the Hokkaido coast are opportunistic feeders that utilize a wide variety of prey, and appear to feed mainly upon prey that is easily obtained.     

Resource partitioning among air‐breathing marine predators: are body size and mouth diameter the major determinants?

Although the body size of consumers may be a determinant factor in structuring food webs, recent evidence indicates that body size may fail to fully explain differences in the resource use patterns of predators in some situations. Here we compared the trophic niche of three sympatric and sexually dimorphic air‐breathing marine predators (the South American sea lion, Otaria flavescens, the South American fur seal, Arctocephalus australis, and the Magellanic penguin, Spheniscus magellanicus) in three areas of the Southwestern Atlantic Ocean (Río de la Plata and adjoining areas, Northern Patagonia and Southern Patagonia), in order to assess the importance of body size and mouth diameter in determining resource partitioning. Body weight and palate/bill breadth were used to characterize the morphology of each sex and species, whereas the trophic niche was assessed through the use of stable isotope ratios of carbon and nitrogen. The quantitative method Stable Isotope Bayesian Ellipses in R (SIBER) was used to compute the area of the Bayesian ellipses and the overlap of the isotopic niches. The results showed that morphological similarity was significantly correlated with isotopic distance between groups within the δ¹³C–δ¹⁵N bi‐plot space in the Río de la Plata area, but not in Northern and Southern Patagonia. Furthermore, resource partitioning between groups changed regionally, and some morphologically distinct groups exhibited a large trophic overlap in certain areas, such as the case of male penguins and male sea lions in Southern Patagonia. Conversely, female sea lions always overlapped with the much larger males of the same species, but never overlapped with the morphologically similar male fur seals. These results indicate that body size and mouth diameter are just two of the factors involved in resource partitioning within the guild of air‐breathing predators considered here, and for whom – under certain environmental conditions – other factors are more important than morphology.     

DNA-based identification methods of prey fish from stomach contents of 12 species of eastern North Pacific groundfish

Stomach content analysis of eastern North Pacific groundfish has been conducted routinely by researchers interested in understanding trophic interactions between key predator species and their prey. Identification of prey by traditional morphological methods has limitations however, due to the loss of identifiable characters from digestion and morphological similarities between taxa. Furthermore, some forage fish (e.g., osmerids, ammodytids, and juvenile gadids), common prey of Bering Sea and Gulf of Alaska groundfish, are difficult to distinguish because of their slender or fusiform shape, disarticulating easily during digestion. DNA-based identification methods were developed to differentiate among 18 fish species, some that are found at depths greater than 200 m, from four taxonomic families: Ammodytidae and Osmeridae (forage fish), Pleuronectidae (flatfish), and Gadidae (gadid fish). Polymerase chain reaction (PCR) amplification of a 739 basepair section of mitochondrial DNA cytochrome c oxidase I and an 862 basepair section of mitochondrial DNA cytochrome b was followed by restriction digest assays and resulted in species level resolution for 16 of 18 species of interest. PCR restriction digest assays applied to fish prey from stomach contents of groundfish indicated the presence of several target species, eulachon (Thaleichthys pacificus), walleye pollock (Gadus chalcogrammus), searcher (Bathymaster signatus), rock sole (Lepidopsetta bilineata), yellowfin sole (Limanda aspera) and either Bering flounder (Hippoglossoides robustus) or flathead sole (H. elassodon). The PCR restriction digest protocols improved the identification rate of predated fish from stomach contents compared to identification by conventional taxonomic methods alone, and DNA sequence analysis further resolved identification of unknown prey fish samples.     

Hooker's sea lions in southern New Zealand

The discovery at Stewart Island of a hauling‐out ground regularly used by Hooker's sea lions (Phocarctos hooked) is described, and recent sightings of sea lions in southern New Zealand are reported. An attempted copulation between a male Hooker's sea lion and a dead female fur seal (Arctocephalus forsteri) is also described.     

Variations in crab claw morphology and diet across contrasting inter‐tidal habitats

Crabs are important predators of inter‐tidal ecosystems, controlling the abundance and distribution of their prey populations. Often the same crab species occupies several habitats and, although their effects on prey have been quantified across habitats, crabs’ dietary and morphological responses to differing environmental influences have been overlooked. Here, we used the crabs Eriphia verrucosa and Pachygrapsus marmoratus as model species to examine differences in claw morphometry – size and wear – and diet between rocky shore and heterogeneous sand flat habitats. We predicted that, intra‐specifically, crabs from rocky shores would consume more hard‐shelled prey owing to their high availability and consequently, would display chelipeds with the following claw characteristics: a higher degree of claw damage, stronger musculature (higher propel height) and increased mechanical advantage (defined as the ratio of input lever length to output lever length) than crabs in the heterogeneous sand flat habitats. Sampling was performed in heterogeneous sand flat habitats and rocky shores of the Central Portuguese coast. For each crab species, carapace width, diet composition and several claw morphometric measures were recorded, revealing significant intra‐specific differences (using multivariate analysis) between shore types. We found that E. verrucosa and P. marmoratus consumed more hard prey on rocky shore than on sand flat habitats, which resulted in rocky shore crabs having more accentuated dentition wear and larger musculature than their sand flat habitat counterparts. We suggest that the strong response of crab claw morphometry to environmentally induced diet variations is an important mechanism in the successful adaptation of crab species to inhabit differing habitats. A major implication is that the impact of the same species on prey may vary largely with habitat type as a result of predation efficiency varying with claw condition.     

Angler perceptions of California sea lion (Zalophus californianus) depredation and marine policy in Southern California

California sea lions (Zalophus californianus) off the coast of Southern California are known to damage both commercial and recreational fishing activities, causing decreases to fish catch and damage to gear. Their increasing population has intensified the potential for conflict between sea lions and anglers, likely requiring changes to current legislation. The recreational fishing community in Southern California is a valuable and largely underutilized source for information and potential solutions to management and legislative problems. This recreational fishing survey-based study conducted in 2013 utilized personal interviews, conducted in the field with recreational anglers and commercial passenger fishing vessel (CPFV) crews in Southern California, to gather data on: (a) the occurrence and impact of sea lion depredation on the local fishing, (b) angler awareness and opinions on current legislation, and (c) the conflict between fishing activities and conservation efforts. Results show that surveyed CPFV operators and private boaters had the most conflict with sea lions and perceive them as more of a problem than anglers on piers, jetties or kayaks. The conflict was also reportedly more prevalent in San Diego County compared to the other counties surveyed (Orange, Los Angeles and Ventura). Participating CPFV operators were overwhelmingly in support of a government culling program for sea lions, while recreational angler respondents did not feel that a control program was necessary. These CPFV operators reported more money lost, and were willing to pay more for an effective deterrent device. There was also a consensus among respondents that fish catch is declining, yet anglers were unsatisfied with the effectiveness of current legislation designed to increase fish stocks. These data will provide a better understanding of California sea lion depredation in Southern California and its effect on recreational anglers in order to aid future mitigation efforts. Additionally, these results provide stakeholder feedback on local marine protected areas and other fisheries management legislation, and build a foundation for future conservation and education programs.     

The food and feeding of seven fish species from the Campbell Plateau,New Zealand

The food and feeding habits of hoki (Macruronus novaezelandiae), southern blue whiting (Micromesistius australis), javelin fish (Lepi‐dorhynchus denticulatus), ling (Genypterus blacodes), smooth rattail (Coelorinchus aspercephalus), silverside (Argentina elongata), and small‐scaled notothenid (Notothenia microlepidota) sampled from the Campbell Plateau in 1979 were examined. The importance of prey items in the diet has been assessed by an ‘index of relative importance’, which combines measurements of frequency of occurrence, number, and weight of prey. Hoki, southern blue whiting, and javelin fish are pelagic feeders. Hoki preyed largely on natant decapod crustaceans, amphipods, and myctophid and photichthyid fishes. The main prey of southern blue whiting were amphipods, natant decapods, and euphausiids. Javelin fish fed on natant decapods, amphipods, and small squid. Seasonal and regional differences in feeding, and dietary changes with length of fish were evident. Ling, smooth rattail, silverside, and small‐scaled notothenid are predominantly benthic feeders. Ling preyed on natant decapods, macrourid fishes, and small hoki. The diet of rattail comprised natant decapod crustaceans, opal fishes (Hemerocoetes spp.), and poly‐chaetes. Silverside fed almost solely on salps. Salps, amphipods, brachyuran crustaceans, and opal fishes were the main prey of small‐scaled notothenid.     

The first use of Fulton's K for assessing and comparing the conditions of inter‐tidal fish populations

Fulton's K condition factor was applied, for the first time, to inter‐tidal specimens of the shanny (Lipophrys pholis) and long‐spined scorpion fish (Taurulus bubalis) from two English rocky shore and two Welsh rocky shore sites during summer 2010 and winter 2011. As both species contribute to the diet of commercial species such as cod (Gadus morhua) and near‐threatened species such as the European otter (Lutra lutra), their condition may affect that of these predators. Fulton's K found that inter‐tidal Welsh fish maintained a ‘good’ condition between seasons, whereas the inter‐tidal English fish were in a poorer condition during winter. Although condition also changed amongst the sites on each coast, further studies are needed into fish morphologies, environmental parameters, prey availabilities and abundances, and fish specimen sex and maturities.     

Dietary analysis of harbour seals (Phoca vitulina) from faecal samples and overlap with fisheries in Erimo,Japan

The number of harbour seals in Japan has been rebounding since protection began in the mid‐1980s. With the increase in seal numbers, increased conflict with fisheries has occurred through depredation and the belief that seals compete with fisheries for prey. However, competition can only be determined if the prey species and quantities seals consume over time are known. We studied the diet of harbour seals in Erimo, site of the largest population of harbour seals in Japan, from 2011 to 2012 and assessed the degree of prey overlap with local fisheries. We used both hard parts and DNA techniques to identify prey items in seal scats, and compared these results to local fisheries data. A total of 46 prey occurrences was detected by both methods, of which 17 matched between techniques at least to the family level. Hard parts methods identified five incidences of prey undetected by DNA methods in five scats (one incidence per scat). DNA methods identified 24 additional prey occurrences in 13 scats, for which no hard part evidence for that prey had been found. This more than doubled the total number of prey occurrences across the 15 scats compared. Overall, the most frequently occurring harbour seal prey were walleye pollock (Theragra chalcogramma), sculpins and snailfishes. In contrast, the top three groups targeted by fisheries were codfishes, salmon and invertebrates. Many species common in the harbour seal's diet such as snailfishes and blennies were not targeted by fisheries. Fishes such as greenlings, sculpins, rockfishes and Japanese anchovy (Engraulis japonicus) were common in the diet of harbour seals, but made up a very small proportion of fisheries catches in Erimo. The importance of other prey species varied between seasons. Sculpins, greenlings and forage fishes were the top three prey groups for harbour seals in spring by percent modified frequency of occurrence, whereas the most important target groups by percentage mass caught by fisheries were codfishes, cephalopods and other invertebrates. Sculpins and rockfishes remained as important prey items for harbour seals in summer and autumn, in addition to codfishes, snailfishes and blennies. In contrast, the main groups caught by fisheries in summer were other fishes, particularly Japanese amberjack (Seriola quinqueradiata) and various species of sharks and eels, and invertebrates. By autumn, Erimo fisheries had focussed on catching salmon. Salmon were taken by harbour seals as well during this period, but at a relatively lower frequency compared to other prey groups. The results of our study show that although harbour seals consumed several of the prey species targeted by fisheries, the relative importance of these species to seals and fisheries and the seasons in which they were targeted were different.     

Trophic ecology of Pomatoschistus microps within an intertidal bay (Roscoff,France), investigated through gut content and stable isotope analyses

The diet of Pomatoschistus microps has been studied using both gut content and stable isotope analyses. In the Roscoff Aber Bay (Brittany, France), this fish is commonly found on sandy muddy intertidal flats. Gut content analyses were also interpreted using trophic indices. Owing to the large diversity of prey consumed, these indices emphasised the opportunistic feeding behaviour of P. microps. Here, this species fed mainly on endofauna with meiofauna being of high relative importance. The main biotic components of its trophic habitat, characterized by δ¹³C and δ¹⁵N, provided evidence of a major trophic pathway based on drift Enteromorpha sp. Trophic positions estimated by both diet analyses and isotopic analyses led to similar results. In this bay, P. microps is a first‐order predator with a low degree of omnivory. Despite a preferential consumption of the amphipod Corophium arenarium, we assumed that this goby behaves as a generalist feeding on a uniform variety of endofauna taxa.     

Insights into the diet of a poorly known species: pygmy killer whale Feresa attenuata from Cape Verde,West Africa

Despite having a worldwide distribution in tropical waters, knowledge on pygmy killer whales Feresa attenuata, including diet, is poor, with only a few studies carried out to date. The presence of otoliths and beaks in stomachs that have been examined indicate that the diet of F. attenuata includes squid, octopus and fish. In this study, the stomach contents of two F. attenuata recently stranded in Cape Verde, West Africa, were examined. A variety of fish hard parts and otoliths were recovered, in addition to the upper beak from an unidentified cephalopod and the remains of some marine arthropods. Lanternfish (Myctophidae) and deepbody boarfish Antigonia capros were identified from otoliths and comprised 95.4% of prey items. Whereas the sizes of the myctophids in the two whale stomachs were similar, those of other prey items differed. The findings provide important information on the diet of F. attenuata in the Cape Verde region.     

Perceptions and attitudes of stakeholders towards the wildlife refuge of Ilha dos Lobos,a marine protected area in Brazil

At the north coast of Rio Grande do Sul state, southern Brazilian coast, the Wildlife Refuge of Ilha dos Lobos (29°20′S; 49°43′W), is a full protection area and a seasonal resting refuge for South American sea lions (Otaria flavescens). The regular presence of sea lions on the island is not only a touristic attraction in the region, but also a problem to the fishery community. This community faces economic impacts in its activities due to fish depredation caused by sea lions. Because of this issue, the perceptions and attitudes of different stakeholders from the region are important to be considered by managers, so that a realistic and decentralized action plan can be designed. The goal of this study was to analyze how different local stakeholders perceived the Wildlife Refuge of Ilha dos Lobos. Additionally, how these stakeholders differed from each other in terms of attitudes, knowledge and perceptions. In October 2011, January and March 2012, face-to-face interview surveys were conducted with fishermen and tourists, and drop-off questionnaires surveys with business, and tourism professional and teachers, in a total of 150 questionnaires, 30 for each group. The results indicated existence of differences across the five stakeholder groups. It was also suggested that non-direct stakeholders had a more favorable view of the Wildlife Refuge of Ilha dos Lobos and the sea lion, whereas direct stakeholder (the fishermen) saw it as an obstacle to their fisheries activities.     

南海狮的回声定位

国外对加州海狮(Zalophus californianus)的回声定位进行了一些研究,但其它种类海狮未见报导.我们在对加州海狮进行了一些实验后,对南海狮(Ostoriabyronia)进行了研究,以对其回声定位能力进行判断和评价,并与加州海狮进行对照比较.     

Distribution of sea cucumbers in relation to sediment characteristics in coral reef lagoons and adjacent waters around Ishigaki Island,southern Japan

To elucidate which environmental factors affect lagoonal‐scale sea cucumber distributions in Ishigaki Island, Okinawa, Japan, intertidal and subtidal areas of three coral reef lagoons were classified into several ground divisions by bottom characteristics, and sea cucumber densities therein were compared with the composition of sediment cover, grain size and organic content, and coverage of macroalgae, seagrass, and massive corals. Holothuria atra, Holothuria leucospilota, Stichopus chloronotus, and Synapta maculata had highest densities in the nearshore areas but were rare in reef flats, probably because of wave disturbance and low areal cover of sand sediment as potential feeding environments. No relationship was observed between sea cucumber densities and sediment organic content or grain size. Thus, even if these sea cucumbers have selectivity for habitats with a high sediment organic content, the effect of such selectivity on the distribution seems to be limited to relatively small areas. The sea cucumber distributions can be classified by bottom sediment/biota composition into bedrock (H. leucospilota), sand (H. atra), and lagoonal types (St. chloronotus and Sy. maculata). These habitat selections were possibly related to various aspects of sea cucumber ecology such as refuge from predators or turbulence, or settlement and nursery place, which have implications for importance of the complexity of lagoonal‐scale topography and sediment/biota conditions for the coexistence of various holothurian species.     

Regional and species specific sexual reproductive patterns of three zooxanthellate scleractinian corals across the Eastern Tropical Pacific

Sexual reproduction of zooxanthellate scleractinian corals in the Eastern Tropical Pacific (ETP) is influenced by the interactive effects of regional and local oceanographic conditions, as well as a variety of other environmental factors. Differences in spatial and temporal gamete development and reproductive patterns of three widespread scleractinian corals of this region—Pocillopora damicornis (branching colony morphology, characterized as hermaphrodite broadcaster), Pavona gigantea (massive colonies, characterized as gonochoric broadcaster/sequential co‐sexually hermaphroditic) and Porites panamensis (encrusting colonies, characterized as gonochoric brooder)—were evaluated at local and regional scales across the ETP. This area extended from the Gulf of California (24°N) to the southern coast of Ecuador (–1°S), including offshore islands, using existing data pooled from prior studies. Predictive models were employed based on environmental variables: sea surface temperature, daylight hours, diffuse attenuation co‐efficient at 490 nm and photosynthetically active radiation. Datasets were extracted from satellite images ( https://oceancolor.gsfc.nasa.gov ) and analysed using WAM_STATIST ver. 6.33® software to obtain monthly average data from each site. The spatial (region, sub‐region and site) and seasonal (wet, dry) variation in reproductive activity (% colonies with gametes) differed among the three species; significant interactions were season × sub‐region for P. damicornis, season × site (sub‐region) for P. gigantea, and season × site for P. panamensis, for which sub‐region was not considered as a factor. The predictive models also suggest that gamete production/maturation of P. damicornis and P. gigantea is influenced by local differences in sea surface temperature and daylight hours. Porites panamensis was not correlated with any environmental variable examined. Variations in local and regional reproductive developmental patterns are likely an acclimatization response by each species imposed by the timing and duration of local stressor events. This analysis has provided insights into the diverse local and regional physical drivers that affect species responses and acclimatization in sexual reproduction across the ETP.     

Molecular evidence for Myripristis jacobus and Scarus taeniopterus new to Bay of Bengal: Sporadic appearance or preliminary colonization?

Individual specimens of Myripristis jacobus Cuvier, 1829 and Scarus taeniopterus Lesson, 1829 were collected from Gopalpur‐on‐sea, Odisha coast, Bay of Bengal. Both the fish samples were assigned species level tags using the conventional taxonomic methods. The capture of both the fish species is of particular interest in that they were new to the Bay of Bengal. Molecular analysis using the mitochondrial cytochrome oxidase subunit I (COI) gene showed high confidence sequence similarity in species identification. The resultant phylogenetic relationships strongly support the monophyly of both the genus with congruent clustering of both the species according to their morphological identification. In addition, the time tree produced is in complete agreement with the recent studies. These studies conclusively prove that both the species might have been originated during the middle to the late Miocene period. Furthermore, both fish species are typical inhabitants of stigmatized coral reefs, confined to the reef regions. These fish species perhaps migrated to the Bengal Bay from the Andaman and Nicobar Islands reef regions because of climatic changes.     

The feeding habits of slope dwelling macrourid fishes in the eastern North Pacific

The diet of slope dwelling macrourid fishes in the eastern North Pacific is poorly known. We collected several hundred stomach samples to investigate the feeding habits of Coryphaenoides acrolepis and Albatrossia pectoralis, the two dominant slope dwelling macrourids off the continental United States. Coryphaenoides acrolepis exhibited a pronounced ontogenetic shift in diet. Specimens <15 cm pre-anal fin length (PAF) consumed primarily polychaetes, amphipods, cumaceans and mysids, while larger individuals consumed increasingly larger, more pelagic prey such as fish, squid, and large crustaceans. Scavenging was also very important to specimens >15 cm with scavenged food constituting approximately 20% of the weight of total prey and occurring in approximately 20% of fish 21–29 cm. Albatrossia pectoralis consumed primarily midwater fish and squid, and we believe that it feeds in the water column. There were significant differences between the diets of A. pectoralis and C. acrolepis suggesting some degree of niche separation between macrourid species on the continental slope of the eastern North Pacific. Both species are at the top of the food web on the upper continental slope and, because of their abundance, may exert significant pressures on their prey populations.     

Predominance of parallel‐ and cross‐predation in anglerfish

Several studies in the last 20 years have revealed that morphological asymmetry in fish can be characterized as ‘antisymmetry’. Antisymmetry is a lateral dimorphism in which each population consists of individuals with well‐developed left sides (lefties) and well‐developed right sides (righties). This dimorphism influences predator–prey interactions. In some piscivorous fishes, it has been found that predators can catch more prey of the opposite morphological type to themselves (cross‐predation) than of the same morphological type (parallel‐predation). Our previous work clarified that the predominance of cross‐predation is caused by lateralized behaviors of predators and prey that correspond to their morphological antisymmetry. Moreover, based on the results of our behavioral observations, we hypothesized that parallel‐predation can predominate when predators encounter the potential prey frontally. To test this hypothesis, in the present study we investigated the relationship between lateral morphological types of anglerfish (Lophiomus setigerus) and those of the prey fishes found in their stomachs. Anglerfish attract potential prey using their first dorsal fin (illicium) as a lure, and their frontal encounters with potential prey fishes were photographed in situ and observed in an aquarium. The results of a stomach contents analysis indicated that parallel‐predation predominated in five benthopelagic prey fish species (perches and eels). By contrast, five benthic prey fishes (gobies and weevers) exhibited the predominance of cross‐predation. These results not only demonstrate the predominance of parallel‐predation in a natural fish community, but also suggest that the relationship between morphological types of predator and prey species can be reversed depending on the lifestyle of prey.     

The intermingling of benthic macroinvertebrate communities during a period of shifting range: The “East of Nantucket” Atlantic Surfclam Survey and the existence of transient multiple stable states

A survey of the region eastward of Nantucket provided an opportunity to examine the cold temperate–boreal boundary along the high‐energy Great South Channel. Here described are the benthic macroinvertebrate community types encountered, with a focus on the influence of climate change on the range boundaries of the benthic biomass dominants and the potential existence of transient multiple stable states. The survey identified three primary community types. The shallowest sites were occupied by a surfclam‐dominated community, comprising an abundance of large (≥150 mm) surfclams, and a few common attached epibiota primarily attached to exposed surfclam shell. Two communities exist at intermediate depths, one dominated by submarket and small market‐size surfclams (<150 mm) and the other, created by mussel mats and their attendant epibiota, crabs, sea urchins, and other mobile epifauna. Mussels are a foundational species, establishing a hard‐bottom terrain conducive to these other denizens in soft‐bottom habitat. Cobbles were nearly ubiquitous, rocks were routinely recovered, and boulders were encountered occasionally. Slow growing attached epibionts were exceedingly rare and mobile epifauna were not obviously associated with these large sedimentary particles; nor were the surfclam or mussel communities. The frequency of barnacle scars suggests sediment scour under the high‐flow regime characteristic of the surveyed region, which voids the habitat potential of these sedimentary particles. The abundance of surfclam shell indicates that surfclams have inhabited the shoaler depths for an extended time; limited shell at deeper sites supports the inference from the absence of large animals that these sites are relatively newly colonized and represent further evidence of an offshore shift in range brought on by increasing bottom water temperatures. The dichotomous nature of the two primary community types at mid‐depths suggests that these two communities represent multiple stable states brought on by the interaction of an invading cold temperate species with the receding boreal fauna resulting in a transient intermingling of species, which, however, structure the habitat into exclusionary stable states rather than overlapping in a co‐occurrence ecotone.