Empirical evidence for large‐scale human impact on intertidal aggregations,larval supply and recruitment of Pyura praeputialis around the Bay of Antofagasta,Chile

In northern Chile, Pyura praeputialis is an invasive species inhabiting rocky intertidal and subtidal habitats restricted exclusively to the Bay of Antofagasta where it forms extensive aggregations. The negative impact of Pyura gathering on mid‐intertidal abundances of this species has recently been reported at the south‐eastern end of this bay. In the present study we have increased sampling sites to cover the entire bay toward the north‐western end and the northern section, where a coastal marine reserve for the scallop fishery partially restricts shellfish gathering. Therefore, the sampling sites were chosen to represent different levels of shellfish gathering access along the northern shore of the bay. Long‐term monitoring (1999–2014) of changes in tunicate cover and the abundances of larvae and recruits at seven sites are reported. The opening of a remodelled artificial and recreational beach in 2012, on the central‐eastern shore of the bay, has increased accessibility to rocky intertidal platforms that started to be massively visited by Pyura gatherers from the summer of 2013. This allowed for the implementation of an intensive short‐term monitoring program of changes in tunicate cover and the abundances of their larvae and recruits. When gathering access was present the reduction in intertidal cover was generalized to the entire bay and followed by reductions in larvae and recruits. However, these reductions were not found in sites with more restricted gathering access. We conclude that continuous extraction by Pyura gatherers followed by reductions of conspecific larvae and recruits are the main drivers behind the reduced abundance of P. praeputialis in the entire bay of Antofagasta. Thus, if gathering is not stopped important ecosystem services provided by this tunicate in the bay may be threatened. Similar consequences may be expected if other massive and irreversible reductions in other species of the Pyura complex, that inhabit other coasts in the southern hemisphere, occur. The controversy concerning the impacts of invasive species and whether they cause negative, positive or neutral impacts to original ecosystems and fisheries is discussed.     

Recent introduction of the dominant tunicate,Pyura praeputialis (Urochordata,Pyuridae) to Antofagasta,Chile

The large sessile tunicate Pyura stolonifera (Pleurogona: Stolibranchiata: Pyuridae), has been regarded as a complex taxon with disjointed distributions, including Australia (Pyura stolonifera praeputialis), South Africa (Pyura stolonifera stolonifera) and South America (Chile, Antofagasta: Pyura sp., the 'piure de Antofagasta'), and has been cited under at least five taxonomic combinations. The 'piure de Antofagasta' is a competitively dominant species in rocky intertidal habitats and shows a limited geographical range (60-70 km) exclusively inside the Bay of Antofagasta. Using cytochrome oxidase I (COI) mitochondrial sequence data from Pyura specimens of the three taxa we tested whether the Chilean taxon represents: (i) a Gondwana relict; (ii) a more recently divergent species; or (iii) a recently introduced species. The results suggest that the Chilean taxon is a recent introduction to Chile from Australian populations and that Pyura stolonifera praeputialis, from Australia, and the 'piure de Antofagasta' are geographical populations of a single species: Pyura praeputialis; whereas the South African taxon represents a second species: Pyura stolonifera.     

Effects of inter-annual temperature variability,including ENSO and post-ENSO events,on reproductive traits in the tunicate Pyura praeputialis

Reproduction in tunicates is considered to be particularly vulnerable to changes in seawater temperature. In the present study we investigated the effects of sea surface temperature and temperature anomalies on reproductive traits of the non-native sessile tunicate Pyura praeputialis. Reproductive traits of this species were investigated over the course of 67 months, based on samples collected at two localities (eastern and western shorelines) of the Bay of Antofagasta. The study period included years with different oceanic and atmospheric conditions: a warm event, El Niño (June 1997 to July 1998); a cold event, La Niña (August 1998 to December 2000); and post-La Niña (January 2000 to December 2002). We compared two common indices (condition and gonadosomatic) and histological sections to evaluate the maximum reproduction index of this introduced species that dominates a large part of the rocky intertidal habitat in the Bay of Antofagasta. We found sexually mature individuals all year-round and a decrease in both reproductive indices matching the reduction in the sea surface temperature during the austral autumn and winter months. The results suggest that gonad development in this species is sensitive to sea surface temperature and thermal anomalies. We conclude that future thermal anomalies or projected global average sea surface warming associated with climate change might have no negative consequences on these reproductive traits of P. praeputialis. This suggests this species is ecologically resilient and that the ecosystem services that this species provides for other invertebrate and algal species will be maintained.     

Allometric differences on the shell shape of two scorched mussel species along the Atlantic South American Coast

Scorched mussels (Bivalvia: Mytilidae) are an important component of rocky shore assemblages around the world. The mussel beds of the mid-intertidal coasts of the south-western Atlantic are composed of two species of scorched mussels, which dominate the physiognomy and structure of the rocky shore communities. The present study investigates the variation in the shell-shape allometric trajectories of both species of mussels throughout their distributional range in the southwestern Atlantic coast, from latitude 34°S to 53°S. Shells of Brachidontes rodriguezii were collected at two Uruguayan localities and four Argentinean localities, while shells of Perumytilus purpuratus were collected at six Argentinean localities. Shell shapes of the specimens were studied by geometric morphometrics using landmark and semi-landmark methods. Different evolutionary histories and thermal regimes characterize these two species. Brachidontes rodriguezii seems to have a long history in the region and is present in a restricted area, the warm temperate region of the south-western Atlantic, while P. purpuratus seems to have a recent history and is present in a wide thermal range, involving the cold-temperate regions of the south-western Atlantic and south-eastern Pacific and the warm-temperate region of the south-eastern Pacific. Based on these considerations, we prompt the hypothesis that allometric trajectories of P. purpuratus shell shape is more variable and adapted to a specific habitat than B. rodriguezii. The impact of allometry was larger in P. purpuratus than in B. rodriguezii. Graphical evidence was gathered, showing that these two species differ in allometric growth forms and that the allometric shell morphology changes reflect environmental constraints and adaptation. While the intertidal mussel beds of the South American coasts are ecologically similar in appearance, we found that the allometric variation of the scorched mussels occurs in response to diverging evolutionary processes such as phenotypic plasticity in P. purpuratus and canalization in B. rodriguezii.     

Foraging behavior of the American oystercatcher Haematopus palliatus pitanay (Murphy 1925) on the intertidal ascidian Pyura praeputialis (Heller 1878) in the Bay of Antofagasta, Chile

Oystercatcher foraging behavior has been described for diverse intertidal prey such as limpets, mussels, and oysters. This paper describes foraging behavior of the American oystercatcher, Haematopous palliatus pitanay, on attached and wave-dislodged ascidians, Pyura praeputialis (prey with a restricted geographic range of 70 km) in the Bay of Antofagasta, Chile. Stabbed holes on the top of the ascidian's tunic, probing excursions, handling time, and five prey-handling sequential stages (striking, hammering, prying, cavity food searching, and swallowing) are described and measured. The need to determine ascidian profit-ability for oystercatcher species in Australia and Chile is highlighted. Received: January 19, 2000 / Accepted: July 23, 2000     

Intertidal mussel beds from the South-western Atlantic show simple structure and uniform appearance: does environmental harshness explain the community?

Communities of the rocky mid-intertidal zone of the South-western Atlantic are uniform in appearance, dominated by dense monocultures of small-size mussels (Brachidontes rodriguezii and Perumytilus purpuratus). To explain this, two hypotheses have been advanced in the literature: environmental harshness due to high potential evaporation and historical contingency after the Last Glacial Maximum. In this study of Uruguayan and Argentine shores, we address the implications and predictions of these two hypotheses from a biogeographic perspective by studying the regional distribution and composition of mid-intertidal mussels. We conducted an extensive latitudinal sampling survey (21 locations, 34–54°S), along with a compilation of available information on mussel bed composition and mussel predators present along the coastline. Then we constructed latitudinal profiles of ecologically significant environmental variables with specific emphasis on potential evaporation, a proxy for desiccation stress. The results show that mussel beds are composed of two species of small mussels, which coexist over a biogeographic transition zone (40–42°S) related to sea surface water temperature. The distribution of mussels along the coastline studied is not consistent with the environmental harshness hypothesis. In addition, in the Central Patagonian zone (44–50°S), two invertebrate predators also inhabit the intertidal rocky shores. However, these localities showed higher environmental harshness (potential evaporation rate) than non-Patagonian localities. We suggest that further attention should be given to historical contingency in order to advance towards a hypothesis consistent with current knowledge on the post-glacial biogeographic history of the South-western Atlantic.     

Relationship between the aerobic and anaerobic metabolic capacities and the vertical distribution of three intertidal sessile invertebrates: Jehlius cirratus(Darwin) (Cirripedia), Perumytilus purpuratus (Lamarck) (Bivalvia) and Mytilus chilensis (Hupé) (Bivalvia)

In protected areas of the southern Chilean coastline, the barnacle Jehlius cirratus dominates the upper intertidal zone, the bivalve Perumytilus purpuratus is the dominating species in the intermediate zone and Mytilus chilensis in the middle-lower zone. Varying degrees of aerial respiration were observed in the laboratory for these three sessile invertebrates: 80–100% of the immersed oxygen uptake for Jehlius cirratus, 30–50% for Perumytilus purpuratus and 5–15% for Mytilus chilensis. The pyruvate kinase/phosphoenolpyruvate carboxykinase (PK/PEPCK) ratio (adductor muscle) was higher for P. purpuratus than for M. chilensis. J. cirratus had no PEPCK activity in total soft tissues, under the assay conditions used in this study, and PK activity was higher for the cirriped than that of both bivalves. Succinate was produced in both mussels when exposed to hypoxic conditions, but it was not detected in J. cirratus. The kinetic parameters of the PK revealed that M. chilensis presented the highest _0.5 value for PEP and J. cirratus the lowest. Alanine inhibited the PK from M. chilensis to a greater extent (higher n_H value) and did not affect the PK of the cirriped. This latter enzyme also presented a marked substrate inhibition, above PEP concentrations of 0.5 mM. The evidence suggests that organisms inhabiting the upper intertidal zone could utilize an aerobic metabolism, given their high aerial respiration capacity, and that the succinate anaerobic pathway could be a more important metabolic strategy in species of the middle-lower intertidal zone. The physiological and biochemical capacities of these three species could thus be correlated with their vertical distribution in the intertidal rocky shore.     

Ecological implications of invasive tunicates associated with artificial structures in Puget Sound, Washington, USA

The non-native tunicates Didemnum vexillum, Ciona savignyi, and Styela clava are of concern to resource managers of Puget Sound, Washington, USA because they have been shown to threaten native species diversity and shellfish aquaculture in other regions. Invasive tunicates in Puget Sound occur mainly on man-made structures such as floating docks and aquaculture facilities. We conducted studies of the three species of concern and a fourth introduced tunicate, Botrylloides violaceus, that occur on these structures to evaluate their effects on mussels and native invertebrate communities. Because most studies of community effects of tunicates have dealt with sessile fouling organisms, we focused instead on epibenthic organisms such as meiofaunal harpacticoid copepods and macrofaunal polychaetes and amphipods that are known to be important prey for juvenile salmon and other small fish. Similar studies have shown mixed results, with negative, positive, or no effects depending on the species. We also found few community-level effects. Abundances of several species were lower when tunicates were present, but only at some of the sites. Several other species, including a non-native isopod, were significantly more abundant in the presence of tunicates. However, in most cases results were not statistically significant and more intensive, controlled sampling or experiments may be needed to demonstrate any consistent tunicate effects. Although invasive tunicates cause problems for mussel growers elsewhere, we did not find negative effects on mussels at four sites in Puget Sound. Given the large impacts known to accompany tunicate invasions elsewhere and their relatively recent invasions into Puget Sound, monitoring of their populations and effects should continue in the region.     

Mytilid mussels: global habitat engineers in coastal sediments

Dense beds of mussels of the family Mytilidae occur worldwide on soft-bottoms in cold and warm temperate coastal waters and have usually been considered hot spots of biodiversity. We examined intertidal mussel beds at four distant locations around the globe with the same sampling method, to find out whether this “hot spot” designation holds universally. We studied species assemblages within the matrices of byssally interconnected mussels engineered by Mytilus edulis in the North Sea, by mixed Perumytilus purpuratus and Mytilus chilensis at the southern Chilean coast, by Musculista senhousia in the Yellow Sea and by Xenostrobus inconstans at the coast of southern Australia. In all cases, species assemblages inside mussel beds were significantly different from those outside with many species being restricted to one habitat type. However, species richness and diversity were not generally higher in mussel beds than in ambient sediments without mussels. In the North Sea (M. edulis) and at the Chilean coast (P. purpuratus, M. chilensis), mussel beds have markedly higher species numbers and diversities than surrounding sediments, but this was not the case for mussel beds in Australia (X. inconstans) and the Yellow Sea (M. senhousia) where numbers of associated species were only slightly higher and somewhat lower than in adjacent sediments, respectively. In conclusion, although soft bottom mytilid mussels generally enhance habitat heterogeneity and species diversity at the ecosystem level, mussel beds themselves are not universal centres of biodiversity, but the effects on associated species are site specific.     

Complementary influences of co-occurring physical ecosystem engineers on species richness: insights from a Patagonian rocky shore

Structural modification of the environment by physical ecosystem engineers often allows for the occurrence of species that are not able to establish in unengineered habitats, thus leading to increased species richness at the landscape-level (i.e., areas encompassing engineered and unengineered habitats). Unlike previous studies that focused on the contribution of a single engineering species to landscape-level species richness, this study evaluates whether co-occurring engineers—i.e., intertidal mussels (primarily Perumytilus purpuratus) and rock boring bivalves (Lithophaga patagonica)—contribute to landscape-level species richness in a similar or complementary way. Our results show that both mussel and L. patagonica patches harbor a substantial number of invertebrate species in addition to those occurring in the unenegineered rock substrate. However, the distinctive habitat patches created by each engineer add exclusive subsets of species to the study area, which implies that mussel and L. patagonica patches contribute complementarily to overall species richness in our intertidal landscape. Here we postulate that complementary engineering effects on landscape-level species richness will occur when the engineered patches structurally differ from each other and, thus, vary in their relative ability to modulate two or more abiotic conditions and/or resources that prevent species establishment in the unengineered state. In spite of its inherently small spatial scale (500 m), our study highlights the potential for complementary engineering impacts at the larger scales that are usually implied in biodiversity conservation and management (tens to hundreds of kilometers) and outlines a simple conceptual basis and approach to address them.     

A new species of Proctoeces and reinstatement of Proctoeces humboldti George-Nascimento and Quiroga 1983 (Digenea: Fellodistomidae) based on molecular and morphological evidence

The most studied digenean of marine organisms in Chile is by far Proctoeces humboldti, a parasite of the intestine of the clingfish Sicyases sanguineus and gonad of the keyhole limpet Fissurella spp. (progenetic metacercariae). The mussel Perumytilus purpuratus has been suggested as the first intermediate host for this digenean. In a study examining the parasites of S. sanguineus from central Chile, we found specimens of Proctoeces showing significant morphological differences with P. humboldti. To assist in the resolution of the taxonomic identification of these specimens, as well sporocysts obtained from the mussel P. purpuratus from central and northern Chile, phylogenetic studies using DNA sequences from the SSU rRNA, as well the LSU rRNA and Cox 1 gene were performed. Results showed that the clingfish S. sanguineus is a host for two species of Proctoeces (P. humboldti and P. syciases n. sp.) along the northern and central Chilean coast, without geographic separation; the mussel P. purpuratus is the first intermediate host for P. syciases n. sp. but not for P. humboldti in central and northern Chile. Fissurellids (Archaeogastropoda) along the Chilean coast harbor only progenetic stages of P. humboldti, but there is no evidence of progenesis for P. syciases. The reinstatement of Proctoeces humboldti is strongly suggested.     

Differences in the activity rhythms of juvenile gobiid fish,Chasmichthys gulosus,from different tidal localities

Activity patterns of intertidal gobiid fish,Chasmichthys gulosus, collected from different tidal localities, were recorded in a constant environment. Gobies collected from a rocky shore with a large tidal range and a regular tidal regime displayed a ca. 12 hour rhythm in their activity pattern. Gobies from a rocky shore with a small tidal range and an irregular tidal regime, however, did not clearly display any such rhythm. The ecological implications of such activity rhythms are discussed from the viewpoints of feeding and avoidance of predators.     

Hard rock versus soft bottom: the fauna associated with intertidal mussel beds on hard bottoms along the coast of Chile,and considerations on the functional role of mussel beds

The fauna associated with hard bottom mussel beds along the exposed Pacific coast of Chile was examined. The abundance of adult (>10 mm body length) purple mussels Perumytilus purpuratus varied between 32 and 75 individuals per 50 cm², and their biomass between 4.8 and 8.6 g AFDW per 50 cm² at eight sampling sites between Arica (18°S) and Chiloé (42°S). At all sampling sites, the associated fauna was dominated by suspension-feeding organisms (cirripeds, spionid and sabellid polychaetes, a small bivalve) followed by grazing peracarids and gastropods. Predators and scavengers also reached high abundances while deposit- and detritus-feeding organisms were of minor importance. The majority of organisms associated with these hard bottom mussel beds feed on resources obtained from the water column or growing on the mussels rather than on materials deposited by the mussels. This is in contrast to the fauna associated with mussel beds on soft bottoms, which comprises many species feeding on material accumulated by mussels (faeces and pseudofaeces) and deposited within the mussel bed. Many of the organisms dwelling between mussels both on hard bottoms and on soft bottoms have direct development, but organisms with pelagic development also occur abundantly within mussel beds. We propose that species with direct development are disproportionately favoured by the structurally complex habitat with diverse interstitial spaces between the mussels, which provides ample shelter for small organisms. We conclude that mussels on hard-bottoms primarily provide substratum for associated fauna while mussels on soft bottoms provide both substratum and food resources. Electronic Publication     

Process of invasiveness among exotic tunicates in Prince Edward Island, Canada

Over the past decade, four exotic tunicates (Styela clava, Ciona intestinalis, Botrylloides violaceus and Botryllus schlosseri) have been reported in the Brudenell estuary in Prince Edward Island (PEI), Canada. Styela clava was the first exotic tunicate to arrive in 1997, rapidly establishing, spreading, invading, and eventually becoming a nuisance in several estuaries of PEI. In the Brudenell estuary, S. clava remained the only exotic nuisance tunicate until 2003. In the fall of 2004, the vase tunicate C. intestinalis, was reported in low abundance, followed by the two colonial species, B. schlosseri and B. violaceus, reported in the spring of 2005. The abundance of C. intestinalis rapidly increased post-introduction, eventually replacing S. clava as the foremost nuisance species on mussel farms in the estuary. To date, C. intestinalis continues to colonize this estuary at epidemic proportions, resulting in the continuing drop of S. clava abundance. The current abundance of C. intestinalis is estimated at 5 cm⁻², which is similar to S. clava abundance at its height in 2003. The 2006 abundance of S. clava is estimated to have fallen to near 0 cm⁻². The dominance of C. intestinalis as a fouling organism on mussel farms is considered a serious threat to this aquaculture industry, mainly due to its unmanageable weight. The process of the detection, establishment, invasiveness, and eventual rise to nuisance level of exotic tunicates in the Brudenell River is presented.     

Patterns of organizations of intertidal and shallow subtidal vegetation in wave exposed habitats of central Chile

Wave-exposed rocky intertidal habitats of central Chile exhibit zonation of algal morphologies rather than strict patterns of species zonation. In low shore areas, there is a vertical sequence of perennial belts of calcareous crusts, kelp-like forms and expanded cushions or non-calcareous crusts. The calcareous crusts are represented by species of Mesophyllum, the kelp-like forms include Lessonia nigrescens and Durvillaea antarctica, while the cushions are represented by Gelidium chilense and G. lingulatum and the noncalcareous, expanded crusts by Codium dimorphum. Thin and thick blades, represented by Iridaea laminarioides, Ulva rigida and Porphyra columbia and filamentous forms including Ceramium rubrum, Centroceras clavulatum and Polysiphonia spp. are more patchy than the lower, perennial belts. They may, however, form distinct temporal monocultures at upper intertidal levels. Upper and lower limits of the various zones are set by interactions of several factors, the relative importance of which can change seasonally. When some of the factors restricting species distribution are experimentally removed, other interactions among factors become limiting.Within each zone, species are morphologically similar, with the abundance of species being regulated by symmetric competitive interactions. Competition is often asymmetric at the boundaries of zones except when adults of small-sized forms interact with morphologically similar juveniles of larger forms. Irrespective of their extremely different morphologies, the permanent, zone-forming algal species generally combine escape from grazers or defensive adaptations with clear competitive abilities. Nevertheless, there is a clear competitive hierarchy which is expressed in vertical displacements and zonation. The lowershore habitats could potentially be occupied by any of the different types of algae. Fast growth and large size allow the kelps to occupy this zone pushing the calcareous crust dominated-zone down into shallow subtidal areas and displacing the cushions and fleshy crusts into the low and middle intertidal regions. In turn, these last forms can displace thick and thin foliose forms and filaments to upper levels on the shore. Displaced forms may exist as patches at various levels of the shore.     

Comparative evaluation of the DNA damage response in two Peruvian marine bivalves exposed to changes in temperature

The comparison of temperature responses of two mytilids from the high (Brachidontes purpuratus) and low (Semimytilus algosus) intertidal zone of the Peruvian coast was carried out focusing on the production of micronucleus and nuclear abnormalities in gill tissue. Two temperatures (23 and 11°C) were evaluated, in presence of three mitomycin C concentrations as a stressor (0.02, 0.04 and 0.06 × 10^?6), simulating hypothetical El Niño and La Niña conditions. Responses to extreme temperatures between both species were significantly different (P = 0.008). Frequency of micronuclei and nuclear abnormalities in S. algosus did not differ statistically among the temperature treatments, whereas in B. purpuratus there was an observed significant decrease in nuclear abnormalities (P = 0.012) and micronuclei frequencies (P = 0.002) between both temperature treatments. The low frequency of micronuclei observed at high temperature and mitomycin C suggests a better efficiency to stress resistance, as occurs during El Niño events, of the species from the higher intertidal zone of B. purpuratus compared to S. algosus from the lower intertidal zone.     

Algae-based biotopes of the Azores (Portugal): spatial and seasonal variation

The increasing importance of coastal management created the need for a systematic classification and characterization of marine communities. Accurate quantitative methodologies for rocky shore algae-based biotope definition, were developed and tested on the Islands of São Miguel and Santa Maria (Azores). Shores of both islands were surveyed, covering all rocky substrate types. Biotopes were defined by assessing the associated habitat and species characteristics, using ANOSIM and SIMPER analysis, respectively. A total of ten biotopes were identified. Generally both islands’ biotopes are characterized by the same taxa/ecological categories, in summer and in winter. However, association between these taxa/ecological categories and the shore height at which they occur differs geographically and temporally. There is a generalized gradual succession of taxa/ecological categories from upper intertidal down to deepest subtidal, although geographical differences occur. Diversity is highest at the land–water interface and decreases towards both extremes (upper intertidal and deepest subtidal level). The strongest evidence of seasonal variation occurs at the upper intertidal. The methodology used proves to be effective in broad scale shoreline assessment of biological communities in warm-temperate coastal marine environments, and thus suitable for the purpose it was developed for. As a consequence it should be applied to the remaining islands of the Azorean archipelago as well as to other macaronesian islands, e.g. Madeira and the Canaries.     

BUCEPHALID TREMATODE INFECTIONS IN MYTILID BIVALVES FROM THE ROCKY INTERTIDAL OF SOUTHERN CHILE

Bucephalid trematode sporocysts were found embedded amidst themantle tissue and viseral mass of intertidal mytilid bivalvesfrom Southern Chile. An investigation into the prevalence ofinfection revealed that <1% of Perumytilus purpuratus wereinfected whereas 20–32% of the Semimytilus algosus examinedharboured sporocysts. No sporocysts were found in an adjacentpopulation of Choromytilus chorus. Prevalence of infection wasrelated to size of S. algosus present at a particular site,and generally increased with host size. The majority of parasitizedmussels appeared to be at an advanced stage of infection, inthese animals destruction of the host's gonadal tissue was evident.Such parasitic castration combined with high prevalence of infectionsuggests that the reproductive potential of this mytilid maybe severely impaired. The implications of these results forthe mariculture industry are also discussed. (Received 19 February 1991; accepted 11 June 1991)     

Mass mortality in a population of the mussel Mytilus edulis L. Caused by high temperature on rocky shores

Mass mortality in a population of the mussel Mytilus edulis L. occurred in early August, 1981, on rocky intertidal shores in Mutsu Bay, northern Japan, due to unusually high temperatures. The decomposed tissues disappeared within 3 days after death. Higher mortality was observed in the upper part of the Mytilus zone than in the lower zone. As temperatures rose the mussels expelled water contained in the shells and 50% individuals died within 1 h from high body temperature rising up to >40°C. Septifer (Mytilisepta) virgatus (Wiegmann), which occupied the zone above Mytilus, showed low mortality (maximum 16.8%). Heat tolerance seems stronger in Septifer than in Mytilus. Many small animals including polychaetes, amphipods and nemerteans etc., lived among the empty shells and when the shells were removed by wave action, these animals disappeared from the bared rock surface. One year after the mass mortality the rock surface was covered by the barnacle Chthamalus challengeri Hoek. The ecological effect of mass mortality on the intertidal community is discussed.     

Habitat structure is more important than nutrient supply in modifying mussel bed assemblage in an upwelling area of the Peruvian coast

Upwelling intensity modifies coastal primary production and influences individual traits of habitat-forming species. Along the Peruvian coast, beds of the mytilid Perumytilus purpuratus provide structurally complex habitats that harbour many organisms. We predict that in the nutrient-rich system of Central Peru, the modification of structural complexity would have stronger effects on the Perumytilus community than nutrient addition. We experimentally examined the effects of nutrient addition on the Perumytilus-dominated assemblage and we evaluated the potential effect of varying shell size on the Perumytilus-dominated assemblage. Nutrient addition to the mussel bed with slow-release fertilizers caused no changes in the total macro- and microalgal biomass and did not affect abundances or composition of the assemblage. To explore the effect of structural complexity on the Perumytilus assemblage, we manipulated mussel size with experimental bags containing small and large individuals. Predators, grazers and mobile organisms were more abundant among smaller mussels, with smaller gap volume, whereas the barnacle Jehlius cirratus was more abundant on larger mussels. In conclusion, point-source nutrient addition to the mussel bed did not enhance primary production. However, the modification of structural characteristics related to mussel size induced changes in the faunal assemblage. Thus, it seems that in this nutrient-rich system, nutrient enhancement would not significantly affect Perumytilus and its assemblage, whereas structural habitat seems to play an important role in shaping this community.