Delayed recovery of giant kelp near its southern range limit in the North Pacific following El Niño

The northern distribution limit of giant kelp, Macrocystis pyrifera (Agardh), along the Pacific coast of North America is relatively stable near Point Año Nuevo, California, but its southern limit has varied over hundreds of kilometers along the Baja California peninsula during the past 20 years. The factors that drive this variability in the southern limit were examined by: (1) quantifying the abundances of M. pyrifera and the subsurface stipitate kelp, Eisenia arborea (Areschoug), near M. pyriferas then southern limit at Punta San Roque (27°15N; 114°42W) on ten occasions between August 1997 and June 2002 (prior to and following the 1997/1998 El Niño Southern Oscillation); (2) initiating a field manipulation at Punta San Roque in June 2000 to examine competition between these two kelp species; and (3) surveying 20 additional sites near Punta San Hipólito (27°01N; 114°00W) in June 2002 to determine whether M. pyrifera populations had recovered to their pre-1982/1983 El Niño southern range limit after ~20 years of absence. These observations and the experimental results suggest that M. pyrifera is removed from its southern limit by the high temperatures and large waves associated with El Niños, while E. arborea is able to survive and recruit in high densities and thereby delay or even prevent M. pyriferas recovery. An inverse relationship was observed between these two species over the 5-year study. The field experiment showed that at both 4 months and 1 year after areas occupied by E. arborea were cleared, substantial recruitment of M. pyrifera occurred, while recruitment was negligible in uncleared areas. This study suggests that the southern range limit of M. pyrifera is determined by increased mortality and recruitment failure following the warm temperatures and low nutrient conditions associated with El Niños, limited substrate availability, and by competition with the more persistent populations of E. arborea.Communicated by J.P. Grassle, New Brunswick     

Energy demand of larval and juvenile Cape horse mackerels, Trachurus capensis, and indications of hypoxia tolerance as benefit in a changing environment

Trachurus capensis is an important fisheries resource in the degraded Namibian upwelling ecosystem. Food supply and shoaling of hypoxic zones are hypothesised to influence the species’ recruitment success. This paper is the first to quantify energy requirements and hypoxia tolerance of larval and juvenile stages of a Trachurus species. We measured normoxic respiration rates of T. capensis with a size range from 0.001 to 20.8 g wet mass (WM) collected off Cape Town (33.9°S, 18.5°E, 12/2009) and in the northern Benguela (17–24°S, 11–15°E, 02/2011). Routine metabolic rate (RMR) and standard routine rate (SRR) (mg O₂ h^?1) followed the allometric functions RMR = 0.418 WM^0.774 and SRR = 0.275 WM^0.855, respectively. Larvae and juveniles had comparatively high metabolic rates, and the energy demand of juveniles at the upper end of the size range appeared too high to be fuelled by a copepod diet alone. T. capensis’ early life stages showed a high tolerance to hypoxic conditions. RMR in larvae did not change until 30 % O₂sat at 18 °C. In juveniles, critical oxygen saturation levels were low (P_C for SRR = 11.2 ± 1.7 % O₂sat and P_C for RMR = 13.2 ± 1.6 % O₂sat at 20 °C) and oxy-regulation effective (regulation index = 0.78 ± 0.09). A high hypoxia tolerance may facilitate the retention of larvae in near-shore waters providing favourable feeding conditions and allowing juveniles to exploit food resources in the oxygen minimum zone. These mechanisms seem to well adapt T. capensis to a habitat affected by spreading hypoxic zones and probably enhance its recruitment success.     

Population dynamics of the sea urchin Strongylocentrotus purpuratus in a Central California kelp forest: recruitment,mortality, growth,and diet

The population dynamics of Strongylocentrotus purpuratus inhabiting dense mats of geniculate coralline algae in a shallow central California Macrocystis pyrifera forest was examined. Sea otters had occupied the area for over two decades. Sea urchin density and size distribution were sampled during 1984–1986 to determine recruitment and mortality patterns. Growth rates were obtained from tetracycline-labeled individuals and changes in sizefrequency modes. Periodic collections were made to examine gut contents and the relationship of diet to food availability. The interaction of recruitment and mortality resulted in a dynamic population structure. Mean densities ranged from 6.5 to 12.7 urchins 0.25 m^-2. The population consisted primarily of urchins up to 40 mm in test diameter in a size distribution which changed from unimodal to bimodal and back over the 2 yr study. Mortality was temporally variable and related to test diameter. Growth rates were somewhat lower than most previous reports for the species and suggested that most of the population was made up of 1 and 2 yr-old individuals. Diet consisted largely of geniculate coralline algae, with fleshy brown algae becoming important when available as drift. This study showed high densities of small urchins can exist in a California kelp forest inhabited by sea otters, but regular recruitment may be necessary to maintain such populations.     

Stalked tunicates Boltenia ovifera form biogenic habitat in the rocky subtidal zone of Nova Scotia

The stalked tunicate Boltenia ovifera is widely distributed in the Arctic and North Atlantic oceans on rocky substrata at 10–300 m depth, although its ecological role in benthic communities is poorly understood. The distribution and abundance of B. ovifera were recorded at 10–100 m depth in towed-video transects in November 2011 and in June, July, and November 2012 off a wave-exposed headland near Halifax, Nova Scotia. Specimens also were collected at 30 m depth using SCUBA (44°26.88′N, 63°31.59′W) in September 2012. Areas dominated by B. ovifera had densities of 10–100 individuals m^?2 on rocky substrata at 30–60 m depth. These tunicate beds often were associated with sparse kelp (Agarum clathratum) and turf-forming red algae. A generalized additive model indicated that depth, substrate type, and benthic algal type were strong predictors of tunicate abundance. Twenty-two epibiotic species were found on specimens of B. ovifera, including juvenile conspecifics. Filter-feeding macroinvertebrates, including anemones and soft corals, were more abundant in areas with B. ovifera than in areas without these tunicates. Our findings suggest that beds of B. ovifera can act as biogenic habitat to enhance local species richness in the rocky subtidal zone of Nova Scotia.     

Ammonium excretion in a temperate-reef community by a planktivorous fish,Chromis punctipinnis (Pomacentridae), and potential uptake by young giant kelp,Macrocystis pyrifera (Laminariales)

The blacksmith Chromis punctipinnis, an abundant planktivorous damselfish off southern California, USA, shelters along rocky reefs at night. While sheltered, blacksmiths excrete ammonium that could, in turn, be utilized by nearby benthic macrophytes. Laboratory experiments during the summer and fall of 1983 and 1984 indicate that ammonium excretion at night ranged from 18.1 mol h^-1 by a 8.5 g (dry) fish, to 89.1 mol h^-1 by a 27.3 g fish; excretion rates generally declined throughout the night. Field measurements at night indicate that ammonium concentrations were significantly higher in rocky crevices occupied by blacksmiths than in unoccupied shelters, and the ammonium level in one shelter dropped after a blacksmith was experimentally removed. Young kelp plants (Macrocystis pyrifera) are capable of taking up ammonium at night. Ammonium levels in chambers containing both a blacksmith and a young kelp plant were significantly lower than in chambers containing only a fish, and ammonium levels dropped in ammoniumspiked chambers that contained kelp plants. Nighttime ammonium uptake rates by young kelp plants, which averaged 1.6 mol g^-1 (dry) h^-1, were only slightly lower than those during the day. Daytime excretion by blacksmiths occasionally results in elevated ammonium levels in the water column. On two of six days, ammonium concentrations in midwater foraging aggregations were slightly but significantly higher than in upcurrent controls; since blacksmiths typically aggregate at the incurrent margin of kelp beds, the ammonium is swept downcurrent and may be utilized by large M. pyrifera that extend through the water column. Thus, the activities of blacksmiths may results in the importation of extrinsic, inorganic nitrogen to primary producers on temperate reefs.     

The structure of subtidal algal and invertebrate assemblages at the Chatham Islands,New Zealand

We examined the distribution and abundance of organisms on subtidal rocky reefs at nine sites around the Chatham Islands, a remote group 780 km east of southern New Zealand. We sampled five depth strata ranging from 1 to<16 m to identify spatial patterns in the abundance of algae and invertebrates and to assess their variation within and among sites. This information is used to discuss hypotheses concerning community structure at this remote locality. Several patterns were apparent. The immediate subtidal was occupied by the southern bull kelp Durvillaea spp. A suite of 11 fucalean species were dominant to a depth of 10 m with an average abundance of 28 m^-2, while one species, Carpophyllum flexuosum, occurred mostly in deeper water. Only two laminarian species of algae were present at the islands. The indigenous Lessonia tholiformis was abundant at 2.5 to 15 m and was not found in deeper water, while the giant kelp Macrocystis pyrifera was abundant at two sites in 12 to 18 m. The commercially valuable abalone Haliotis iris was extremely abundant in shallow water, with an overall mean of 6 m^-2 at 5 m. The sea urchin Evechinus chloroticus was common, but reached high densities only in small (<25 m²) patches. The characteristic urchin-dominated zones reported in kelp beds world-wide were not seen. There was considerable site-to-site variation in the occurrence and abundance of individual species. Some differences between sites were associated with shelter from swell (e.g. M. pyrifera was found only in sheltered sites) and physical habitat (e.g. juvenile H. iris were found only beneath boulders inshore), but much of the variation could not be explained by physical or depth-related factors alone. We hypothesize that the differences in these kelp bed assemblages compared to mainland New Zealand are partially due to the high degree of endemism at the Chatham Islands. Local variation cannot be explained by herbivory, and is most likely the result of the various life-history characteristics of the major habitat-forming species, the large brown algae.     

Effects of micro- and mesograzers on intertidal macroalgal recruitment

This study examined the effects of a guild of micrograzing harpacticoid copepods (dominated by two species of Paradactylopodia sp. nov. and one species of Scutellidium sp. nov.) and a mesograzing periwinkle, Afrolittorina praetermissa, on the early recruitment of intertidal macroalgae on a wave-exposed, rocky shore. This is the first study, to our knowledge, to examine the effects of micrograzers (<500 μm) on intertidal macroalgal recruitment. Data showed that microscopic harpacticoid copepods altered the assemblages and reduced the densities of several macroalgal taxa, while A. praetermissa changed the assemblages and reduced both the density and number of macroalgal taxa. Recruitment of encrusting coralline algae was actually higher in copepod inclusions than exclusions, suggesting that copepods may be beneficial to the recruitment of this algal group. These results contribute to the understanding of grazing as a factor causing high mortality of algal recruits, but also highlight the need for more studies that examine the effects of micro- and mesograzers on the distribution and abundance of macroalgae.     

Survival of microscopic stages of a perennial kelp (<Emphasis Type="Italic">Macrocystis pyrifera</Emphasis>) from the center and the southern extreme of its range in the Northern Hemisphere after exposure to simulated El Niño stress

Many organisms survive stressful conditions through a tolerant life history stage. The life history known as the alternation of generations is typical of temperate kelps, producing diploid macroscopic stages, and both haploid and diploid microscopic stages, with the haploid stages thought to be stress tolerant. The survival of microscopic stages of the giant kelp Macrocystispyrifera during El Niño has been suggested, yet has never been tested. This mechanism could be critical for population persistence, particularly at the southern limit of the range in the Northern Hemisphere, which is greatly impacted by El Niño conditions. The purpose of this study was to determine if microscopic stages of giant kelp could survive and recover from El Niño-type conditions and whether those from a population near its southern limit were more tolerant than a population at the center of its range. Microscopic stages were exposed to a laboratory simulation of potential El Niño conditions (high temperature, with and without light and nitrate) for 8 weeks and then allowed to recover at optimal conditions (low temperature and high nitrate) for 8 weeks, while controls were left at optimal conditions the entire 16 weeks test period. Haploid developmental stages from both populations survived and recovered from stressful conditions with no population level effect, suggesting haploid stress-tolerance may be widespread. The more advanced the developmental stage, and the presence of nitrate, resulted in significantly greater recovery for haploids. Yet, none of these stages were able to go on to produce sporophytes, whereas all controls did. There was a large population-level effect for diploids, however, with only microscopic diploid stages (embryonic sporophytes) from the southern-limit population recovering from El Niño simulated stress, suggesting ecotypic adaptation for microscopic sporophytes. Diploid recovery was significantly greater with light. We propose that the diploid stage is the most likely to survive and recover after El Niño conditions, as it would avoid obligate egg and sperm encounters after the stress period. The survivorship of the microscopic diploid in a seed bank analogue may be how the isolated southern-limit populations are able to recover after mass disappearance during El Niño.     

Modeling kelp forest distribution and biomass along temperate rocky coastlines

We address the global deficit of data describing kelp forest distribution, relative covers and biomass by testing the ability of species distribution models to predict these attributes at locations where data are currently limited. We integrated biological ground truth data with high-resolution environmental datasets to develop generalized additive models that accurately predict the structure of Laminaria forests within the Bay of Morlaix (48°42′42″N, 3°55′40″W). Forest distribution and proportional covers were predicted using water depth, light availability, wave exposure and sediment dynamics. The biomass of individual kelp species was modeled by supplementing these same variables with measures of seafloor slope and benthic position. Biomass predictions for Laminaria digitata and Laminaria hyperborea contrast the physiological tolerances of these species to light and wave exposure gradients. As a direct management output, we produced high-resolution maps (25 m² grids) that closely match independent field data and provide vital information for marine spatial planning.     

Population genetic structure of the stalked barnacle <Emphasis Type="Italic">Pollicipes pollicipes</Emphasis> (Gmelin, 1789) in the northeastern Atlantic: influence of coastal currents and mesoscale hydrographic structures

Within its distribution range in the northeastern Atlantic, the stalked barnacle Pollicipes pollicipes shows a well-defined pattern of genetic variation, comprising (a) a subtropical/temperate northern assemblage, made up of populations distributed between 47°N and 28°N along the French, Iberian, North African and Canary Islands coastlines, and (b) a single isolated and highly divergent tropical population in the Cape Verde Islands (16°N), at the southernmost limit of the species’ distribution. However, within the northern assemblage several populations show a level of genetic differentiation that allows rejection of the hypothesis of genetic homogeneity. The congruence observed between genetic and hydrographic patterns suggests a crucial role of hydrodynamics, and of the dispersal of the planktonic larvae, in the determination of population structure. Along the southern European Atlantic coast, the Iberian Poleward Current and mesoscale hydrographic structures are, respectively, facilitating gene flow at the regional level and genetic differentiation at the local level. On the Atlantic coast of North Africa, the homogenizing equatorward flow of the Canary Current does not extend as far as the Cape Verde Islands. A demographic expansion is dated to the late Pleistocene, preceding the Eemian interglacial, and is oldest in the case of the long-standing Cape Verde population, sustained by a stable tropical habitat. The divergence between the Cape Verde population and the remaining populations is thus ancient, and suggests that oceanic current patterns may constitute a generalized physical barrier to the dispersal of marine organisms between Cape Verde and the rest of Macaronesia.     

Aspects of the ecology and growth of an intertidal juvenile population of Concholepas concholepas (Mollusca: Gastropoda: Muricidae) at Las Cruces,Chile

The habitat, density and growth rate for an intertidal population of Concholepas concholepas (Bruguière, 1789) were studied at Las Cruces, Chile (Lat. 33°30′S; Long. 71°38′W) during 1977–1978. The growth rate (3.67 mm month^-1) was determined in a newly settled group, whose average length was 11.3 mm, and whose age was estimated at 3 months. The densities found ranged between 1.1 and 107.3 “locos” m^-2. Based on these results, the time of settlement of C. concholepas was calculated; capsule deposition, maximum maturity and recruitment reported by other authors for different localities are discussed.     

Patterns of co-occurrence and interactions between age classes of the common triplefin,Forsterygion lapillum

Many marine organisms have pelagic larval stages that settle into benthic habitats occupied by older individuals; however, a mechanistic understanding of intercohort interactions remains elusive for most species. Patterns of spatial covariation in the densities of juvenile and adult age classes of a small temperate reef fish, the common triplefin (Forsterygion lapillum), were evaluated during the recruitment season (Feb–Mar, 2011) in Wellington, New Zealand (41°17′S, 174°46′E). The relationship between juvenile and adult density among sites was best approximated by a dome-shaped curve, with a negative correlation between densities of juveniles and adults at higher adult densities. The curve shape was temporally variable, but was unaffected by settlement habitat type (algal species). A laboratory experiment using a “multiple-predator effects” design tested the hypothesis that increased settler mortality in the presence of adults (via enhanced predation risk or cannibalism) contributed to the observed negative relationship between juveniles and adults. Settler mortality did not differ between controls and treatments that contained either one (p = 0.08) or two (p = 0.09) adults. However, post hoc analyses revealed a significant positive correlation between the mean length of juveniles used in experimental trials and survival of juveniles in these treatments, suggesting that smaller juveniles may be vulnerable to cannibalism. There was no evidence for risk enhancement or predator interference when adults were present alongside a heterospecific predator (F. varium). These results highlight the complex nature of intercohort relationships in shaping recruitment patterns and add to the growing body of literature recognizing the importance of age class interactions.     

Reproductive ecology of a hippolytid shrimp,Lebbeus virentova (Caridea: Hippolytidae) at the Von Damm Vent Field,Mid-Cayman Spreading Centre

Elucidation of life-history traits is essential to understand larval dispersal and population dynamics in marine benthic assemblages. This study is the first investigation of the life history of a recently described hippolytid shrimp from a deep-sea chemosynthetic environment, Lebbeus virentova Nye, Copley, Plouviez and Van Dover, 2013 at the Von Damm Vent Field (18°22N, 81°47W, ~2,300 m depth, Mid-Cayman Spreading Centre, Caribbean), using samples collected in February and June 2013. Lebbeus virentova is gonochoric and iteroparous. The sex ratio of L. virentova was significantly female biased (1:3) in February and June. The sampled population of L. virentova had a unimodal size–frequency distribution pattern in February and June, consistent with continuous recruitment and mortality. Continuous reproduction is indicated by a lack of synchrony in oocyte size–frequency distributions within both months, and asynchronous development of embryos among females, which may result in asynchronous larval release. A large embryo size in this species (2.65 ± 0.28 mm diameter) compared with other caridean shrimps suggests possible abbreviated larval development, as described in other species of the genus from non-chemosynthetic environments. Fecundity (26–94 embryos female^?1) was lower and embryo size larger in L. virentova compared with alvinocaridids at chemosynthetic environments. This suggests that there are phylogenetic constraints on reproductive features of decapods at hydrothermal vents.     

Vertical zonation of endosymbiotic zooxanthellae within a population of the intertidal sea anemone,Anthopleura uchidai

Intertidal organisms commonly form zonation bands along the shore. Environmental stressors often determine the vertical position of each zonation band. These stressors may similarly affect the distribution pattern of endogenous species in their intertidal hosts. To evaluate this possibility, we investigated the distribution pattern of endosymbiotic zooxanthellae in the genus Symbiodinium in a population of the intertidal sea anemone Anthopleura uchidai. We used molecular genetics to identify the Symbiodinium clades and found that A. uchidai has two clades of Symbiodinium, clades A and F. These Symbiodinium clades were disproportionally distributed along the vertical gradient of the intertidal shore. Anemones on the upper shore exclusively possessed clade F Symbiodinium while clade A Symbiodinium became dominant in the sea anemones on the lower shore. Photosynthesis activity assays showed that these Symbiodinium clades had similar net productivities at 23.3 and 31.8 °C at all irradiance levels. At 35 °C, however, clade A Symbiodinium exhibited substantially lower net productivities than clade F Symbiodinium, demonstrating that these Symbiodinium clades have distinct tolerances to thermal stress. These results suggest that the thermal gradient across tidal height is a major factor shaping the zonation pattern of Symbiodinium clades in A. uchidai.     

Feeding ecology of juvenile rockfishes off Oregon and Washington based on stomach content and stable isotope analyses

The feeding habits of pelagic, juvenile rockfishes (Sebastes spp.) collected off Oregon in 2002, and Oregon and Washington in 2006, were examined using stomach content and stable isotope analyses. Sampling occurred along a series of transects across the shelf between Crescent City, California (Lat. 41°54.0′), and Newport, Oregon (Lat. 44°39.0′), in 2002, and off Willapa Bay, Washington (Lat. 46°40.0′), and the Columbia River, Oregon (Lat. 46°10.0′), in 2006. Species composition varied both years with distance from shore, but the predominant species were darkblotched (Sebastes crameri), canary (S. pinniger), yellowtail (2006 only; S. flavidus), and widow (S. entomelas) rockfishes. Stomach content analysis revealed that darkblotched rockfish had highly variable diets, and canary, yellowtail, and widow rockfishes exhibited a high degree of overlap in 2006. Multivariate analysis showed significant differences in diet based on distance from shore where caught, fish size, and species. Stable isotope analysis indicated that all species were feeding at about the same trophic level within each year, with a 1.5 ‰ difference in δ¹⁵N between years and regions. The difference in δ¹⁵N values may indicate a greater contribution of mesotrophic zooplankton such as euphausiids, hyperiid amphipods, and chaetognaths to fish diets in 2006. Depleted ¹³C values were indicative of diets based on primary production from a more offshore origin, suggesting that these rockfish had previously inhabited offshore waters. These results add to our understanding of some of the important environmental factors that affect young-of-the-year rockfishes during their pelagic phase.     

Interactions among sea otters, sea stars, and suspension-feeding invertebrates in the western Aleutian archipelago

The structuring and organizing effects of apex predators on ecosystems are becoming increasingly well documented. The enhancement of kelp forests via sea otter predation on herbivorous sea urchins is among the earliest and best known examples. This study provides evidence for direct and indirect trophic interactions among sea otters, predatory sea stars, and filter-feeding mussels (Mytilus trossulus) and barnacles (Semibalanus cariosis). In western Massacre Bay at Attu Island (173°E, 53°N), subtidal transects showed sea star body size and biomass density declined markedly between 1983 and 1994 as sea otters reinhabited this area. Mussels and barnacles translocated from the rocky intertidal zone to shallow subtidal habitats to assess loss rates from sea star predation showed lower mortality rates after the arrival of sea otters. Prey mortality rates in subtidal caged controls were consistently low and similar to those of intertidal controls in both years. These findings elucidate a trophic pathway by which sea otters can influence ecosystems separate from the well-known sea otter/sea urchin/macroalgae cascade.     

Growth and production of new recruits and adult individuals of Ascophyllum nodosum in a non-harvested population at its southern limit (Galicia,NW Spain)

Populations near the geographic distribution limits of the species are considered to live under suboptimal conditions, and hence, slight environmental changes can be critical for their survival. The potential sensitivity to disturbances of the long-living macroalga Ascophyllum nodosum was analyzed by the determination of growth, recruitment, mortality, and production of biomass of a population near its southern distribution limit. Recruitment, survival and growth rates of <2 years old individuals were determined in a new population growing in experimentally denudated squares. Demographic data for >2 years old individuals were obtained from individuals in the original population after estimating their age from the number of gas bladders in the thallus. Growth and survival were described as continuous nonlinear functions of age applied to the population and were further used to make demography-based production estimates. Recruitment of A. nodosum in denudated substrates seemed to require a previous cover of other macroalgae (as Fucus vesiculosus) as the only cohort detected during the 26-month period of the study was observed after F. vesiculosus individuals started to increase. The low production estimates (2,033 g m^?2 for a 10 year period) and poor recruitment may indicate a slow recuperation of this population to denudation. However, the large variability observed in the estimated growth curves of different populations along this southern distribution area suggests a large influence of local conditions that may help to overcome environmental changes at regional scales.     

Carbon emission associated with respiration and calcification of nine gastropod species from the intertidal rocky shore of Western Europe

Intertidal rocky shores are characterized by vertical zonation that results from the interplay between environmental conditions, organism physiology, and species interactions. Metabolism of intertidal organisms is highly variable between species and it changes with vertical position along the intertidal gradient. The present study aimed to quantify the carbon metabolism of nine intertidal rocky shore gastropods, in order to clarify their respective roles in carbon production during emersion and immersion. The influences of monthly temperature variation and tidal level were tested for each species. Analyses were performed in the laboratory using the infrared gas analyzer method for measuring aerial respiration rates, and the dissolved inorganic carbon and total alkalinity technique for measuring aquatic respiration rate and calcification. Hourly carbon fluxes were calculated for the mean annual temperature of 13 °C measured in both air and underwater in the study area. Respiration rates were similar for emersion (8–25 μmol CO₂ g AFDW^?1 h^?1) and immersion (10–23 μmol DIC g AFDW^?1 h^?1). For all species, underwater respiration fluxes were more influenced by monthly temperature variation than by air fluxes, probably as an adaptation to the rapid changes occurring during emersion. Calcification was an important factor influencing annual carbon fluxes for all studied species; every species showed different calcification rates according to its size and position on the intertidal zone. Annual carbon emissions were calculated using the mean immersion/emersion time of each species. Intertidal gastropod carbon emission was primarily influenced by body biomass and their vertical position within the intertidal zone.     

Long-term trend in substratum occupation by a clonal, carbonate bryozoan in a temperate rocky reef in times of thermal anomalies

Studies over time provide opportunities to detect variations in the spatial and temporal patterns of clonal organisms and measure changes on their population dynamics related to extreme events. We assessed population dynamics for a bryozoan species dominating a subtidal rocky reef at Tino Island, in the eastern Ligurian Sea (NW Mediterranean). Using 9 years of annual photosurveys (1997–2005), rapid decline in Pentapora fascialis colony cover was shown at 11 and 22 m depths following the anomalous warming events in 1999 and 2003. An 86 % reduction in live colony portion was found after the 1999 warming event (2.3 °C higher than normal), with larger colonies being most affected. Effects from the 2003 event were delayed, and gradual cover decline occurred during the following 2 years. At the “Shallow” photostations, none of the larger colonies (>1,000 cm²) survived after the first cover decline. Availability of new substrate after the 1999 disturbance resulted in enhanced recovery through new colony production. At the “Deep” photostations, the population structure did not change over the duration of the monitoring period showing the same monomodal structure and same dominant size class (50–500 cm²). In the 4 years following the first cover decline, the deeper population regained colony cover to levels similar to pre-disturbance level, showing a good resilience. This 9-year monitoring analysis provided the temporal resolution needed to detect changes occurring in the P. fascialis population and will contribute to the assessment of long-term changes on benthic populations suffering during recent decades from dramatic increases in extreme events.     

Floating kelps in Patagonian Fjords: an important vehicle for rafting invertebrates and its relevance for biogeography

Floating macroalgae are common dispersal vehicles for associated benthic invertebrates. In order to investigate the importance of kelp rafts for species dispersal in the Patagonian Fjord Region (PFR), the abundance and distribution pattern of floating kelps (Macrocystis pyrifera, Durvillaea antarctica) and of the invertebrate fauna associated with M. pyrifera were evaluated during austral spring of 2002–2005, 2008 and 2010. In the southernmost Magellan Region (MR), benthic M. pyrifera were additionally sampled to compare the community structures in both conditions. Floating kelps were abundant throughout the entire PFR, harbouring a diverse and abundant invertebrate fauna. The density of floating kelps increased towards the south. In the MR, a loss of species was observed between benthic and floating condition (e.g. decapods, echinoderms, several peracarid species), but a high diversity of organisms from all major phyla were observed on rafts. Throughout the PFR, the predominant rafting species belonged to the peracarids, molluscs and annelids, but the community composition differed between floating samples from the northern and southern zones of the PFR. Relative abundances of peracarids were higher in northern zones, whereas molluscs and annelids dominated in the southern areas. Species of the peracarid genera Peramphithoe, Gondogeneia, Bircenna and Limnoria were shared between all areas. The results suggest that kelp rafts not only contribute to local population connectivity in the PFR, but could also be an important dispersal vehicle for rafting species along the PFR in north–south direction, crossing the biogeographic boundary around Taitao Peninsula. Furthermore, the MR appears to be an important stepping stone for species dispersal via kelp rafting in the subantarctic region.