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Abstract

In the frame of the COST ACTION ‘EMBOS’ (Development and implementation of a pan-European Marine Biodiversity Observatory System), coverage of intertidal macroalgae was estimated at a range of marine stations along the European coastline (Subarctic, Baltic, Atlantic, Mediterranean). Based on these data, we tested whether patterns in macroalgal diversity and distribution along European intertidal rocky shores could be explained by a set of meteo-oceanographic variables. The variables considered were salinity, sea surface temperature, photosynthetically active radiation, significant wave height and tidal range and were compiled from three different sources: remote sensing, reanalysis technique and in situ measurement. These variables were parameterized to represent average conditions (mean values), variability (standard deviation) and extreme events (minimum and maximum values). The results obtained in this study contribute to reinforce the EMBOS network approach and highlight the necessity of considering meteo-oceanographic variables in long-term assessments. The broad spatial distribution of pilot sites has allowed identification of latitudinal and longitudinal gradients manifested through species composition, diversity and dominance structure of intertidal macroalgae. These patterns follow a latitudinal gradient mainly explained by sea surface temperature, but also by photosynthetically active radiation, salinity and tidal range. Additionally, a longitudinal gradient was also detected and could be linked to wave height.

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... Holthuijsen, 2010;Reguero et al., 2012;Camus et al., 2013;Guillou and Chapalain, 2015;Rattray et al., 2015). Such models give a framework for local studies and open up new prospects to determine the effect of waves on the shores (Cefalì et al., 2016;Puente et al., 2016). Although aforementioned approaches have provided valuable insights into the relationship between hydrodynamics and intertidal communities, they are of little use at local scale, where small-scale topography (i.e. ...
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The World Register of Marine Species is an over 90% complete open-access inventory of all marine species names. Here we illustrate the scale of the problems with species names, synonyms, and their classification, and describe how WoRMS publishes online quality assured information on marine species. Within WoRMS, over 100 global, 12 regional and 4 thematic species databases are integrated with a common taxonomy. Over 240 editors from 133 institutions and 31 countries manage the content. To avoid duplication of effort, content is exchanged with 10 external databases. At present WoRMS contains 460,000 taxonomic names (from Kingdom to subspecies), 368,000 species level combinations of which 215,000 are currently accepted marine species names, and 26,000 related but non-marine species. Associated information includes 150,000 literature sources, 20,000 images, and locations of 44,000 specimens. Usage has grown linearly since its launch in 2007, with about 600,000 unique visitors to the website in 2011, and at least 90 organisations from 12 countries using WoRMS for their data management. By providing easy access to expert-validated content, WoRMS improves quality control in the use of species names, with consequent benefits to taxonomy, ecology, conservation and marine biodiversity research and management. The service manages information on species names that would otherwise be overly costly for individuals, and thus minimises errors in the application of nomenclature standards. WoRMS' content is expanding to include host-parasite relationships, additional literature sources, locations of specimens, images, distribution range, ecological, and biological data. Species are being categorised as introduced (alien, invasive), of conservation importance, and on other attributes. These developments have a multiplier effect on its potential as a resource for biodiversity research and management. As a consequence of WoRMS, we are witnessing improved communication within the scientific community, and anticipate increased taxonomic efficiency and quality control in marine biodiversity research and management.
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This study examined the interactive effects of grazing by limpets and inclination of the substratum in maintaining differences between mid-shore and low-shore assemblages of algae in the northwest Mediterranean, at different scales of space and through time. Alternative models leading to different predictions about these effects were proposed and tested. Limpets were excluded by fences from areas of the substratum at mid levels on the shore. The response of algal assemblages to this manipulation was compared with control and enclosure plots at the same level, and with unmanipulated plots in the low shore where limpets are less abundant. The effects of limpets were examined at several replicated sites (0.1–4 km apart) for each slope of the substratum (nearly horizontal vs vertical), at different locations (hundreds of kilometres apart) and at different times. Individual taxa responded differently to limpet exclusion. The percentage cover of the coarsely branched and filamentous algae increased significantly in exclosure plots, in some loser reaching values found on the low shore. These patterns, however, varied greatly from shore to shore and significant effects were found both on horizontal and vertical substrata. Multivariate analyses indicated that grazing by limpets accounted for about 20% of the differences between mid-shore and low-shore assemblages. This effect was independent of substratum inclination and was consistent in space and time, suggesting that physical conditions were not as stressful for macroalgae on vertical substrata as initially supposed. Variable recruitment of algae is proposed as a possible explanation for the lack of consistency in the effects of limpets at the scale of the shore. The results of this study emphasize the need for multiple-scale analyses of the interactive effects of physical and biological factors to understand the organization of natural assemblages.
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An ecological classification at the local scale may be a useful tool for conservation planning and for the implementation of specific management programmes in a region. For this purpose, a methodology previously applied on a small scale has been adapted to classify the coast of Cantabria (N Spain). This methodology includes a physical classification and biological validation. The shoreline was divided into 1 km stretches, and the abiotic variables (sea surface temperature, photosynthetically active radiation, significant wave height and coastal morphology) were recorded for each stretch. A hierarchical classification was proposed, with a first level that encompassed a grouping of quantitative variables based on SOM and k-mean analysis and a second level that subdivided the previous groups according to the categorical variable ‘coastal morphology’. To validate the classification using biological data, cover of intertidal macroalgal species was obtained at 14 sites along the study area, and several statistical analyses were applied to test the ecological significance of this classification. Three physical units were obtained (western (W), central (C) and eastern (E) coast), based on abiotic variables. Each group was then subdivided into subunits according to its coastal morphology (cliffs or wave-cut platforms). A general agreement between the macroalgal distribution and physical units was accomplished. In the lower intertidal, Bifurcaria bifurcata and Halopteris scoparia dominated the western and central areas, whereas Corallina spp./Ellisolandia elongata and Gelidium spp. were most abundant towards the east. In contrast, throughout the middle intertidal, Corallina. spp./E. elongata were the dominant taxa. The classification system developed in this study completes a hierarchical framework for classifying the NE Atlantic coast, a promising approach that permits the application of the most suitable resolution in each case study that could be applicable to a wide range of coastal areas.
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Coastal ecosystems are highly complex and driven by multiple environmental factors. To date we lack scientific evidence for the relative contribution of natural and anthropogenic drivers for the majority of marine habitats in order to adequately assess the role of different stressors across the European seas. Such relationship can be investigated by analysing the correlation between environmental variables and biotic patterns in multivariate space and taking into account non-linearities. Within the framework of the EMBOS (European Marine Biodiversity Observatory System) programme, hard bottom intertidal communities were sampled in a standardized way across European seas. Links between key natural and anthropogenic drivers and hard bottom communities were analysed using Boosted Regression Trees modelling. The study identified strong interregional variability and showed that patterns of hard bottom macroalgal and invertebrate communities were primarily a function of tidal regime, nutrient loading and water temperature (anomalies). The strength and shape of functional form relationships varied widely however among types of organisms (understorey algae composing mostly filamentous species, canopy-forming algae or sessile invertebrates) and aggregated community variables (cover or richness). Tidal regime significantly modulated the effect of nutrient load on the cover and richness of understorey algae and sessile invertebrates. In contrast, hydroclimate was more important for canopy algae and temperature anomalies and hydroclimate separately or interactively contributed to the observed patterns. The analyses also suggested that climate-induced shifts in weather patterns may result in the loss of algal richness and thereby in the loss of functional diversity in European hard bottom intertidal areas.
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Ecological classification of coastal waters has become increasingly important as one of the basic issues in the biology of conservation. Management and protection of coastal areas take place at different spatial scales. Thus, proper classification schemes should integrate equivalent information at various levels of definition in order to show its feasibility as a useful tool for assessment of coastal environments at the required scales. In this work, a global approach applied to the classification of the NE Atlantic coast is analysed in order to discuss pros and cons regarding different conceptual and technical issues for effective implementation of such a management tool. Using the hierarchical system applied at three different geographic scales: Biogeographic (NE Atlantic coast), Regional (Bay of Biscay) and Local (Cantabria region), five different topics were considered for debating strengths and weaknesses of the methodological alternatives at those spatial scales, using for validation the rocky shore macroalgae as a representative biological element of benthic communities. These included: (i) the spatial scales; (ii) the physical variables and indicators; (iii) the classification methodologies; (iv) the biological information; and (v) the validation procedure. Based on that analysis, the hierarchical support system summarized in this paper provides a management framework for classification of coastal systems at the most appropriate resolution, applicable to a wide range of coastal areas. Further applications of the physical classification for management of biodiversity in different environmental scenarios are also analysed.
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The boundary between land and sea, i.e. the littoral zone, is home to a large number of habitats whose distribution is primarily driven by the distance to the sea level but also by other environmental factors such as littoral’s geomorphological features, wave exposure, water temperature or orientation. Here we explore the relative importance of those major environmental factors that drive the presence of littoral rocky habitats along 1100 km of Catalonia’s shoreline (Spain, NW Mediterranean) by using Geographic Information Systems and Generalized Linear Models. The distribution of mediolittoral and upper infralittoral habitats responded to different environmental factors. Mediolittoral habitats showed regional differences drawn by sea-water temperature and substrate type. Wave exposure (hydrodynamism), slope and geological features were only relevant to those mediolittoral habitats with specific environmental needs. We did not find any regional pattern of distribution in upper infralittoral habitats, and selected factors only played a moderate role in habitat distribution at the local scale. This study shows for the first time that environmental factors determining habitat distribution differ within the mediolittoral and the upper infralittoral zones and provides the basis for further development of models oriented at predicting the distribution of littoral marine habitats.
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The ecological classification of coastal waters has become an important issue in ecosystem water quality assessment. Previous studies have suggested that abiotic variables seem to be a suitable alternative to biological data for classifying coastal areas at different scales. The study presented here proposes a downscaling methodology for the classification of coastal waters at a regional scale within the NE Atlantic based on standardized data and objective decision rules. Physical variables (temperature, wave exposure, tidal range and radiation) were selected because of their ecological role, availability and statistical decision rules. This information was based on satellite data and mathematical modelling of natural coastal processes. The N and NW Spanish coastline was subdivided into 41 20-km segments that were classified according to physical variables using the self-organizing map and k-means algorithms. To validate the classification with biological data, 21 sites representing the entire range of physical typologies in the study area were simultaneously and consistently sampled. Intertidal macroalgae were identified in each of 10 quadrats of 50 × 50 cm for two to three transects per site, according to a stratified sampling procedure. The coverage of macroalgae was obtained by photographic analysis. The physical classification shows four typologies: Lower Rias, Upper Rias, West Cantabric and East Cantabric. Statistical analyses confirmed the ecological significance of these typologies at the tidal levels where seaweeds were the major structural element (lower and middle intertidal). According to the biological data, the greatest differences were found between the Upper Rias and the rest of the N and NW Iberian Peninsula coast. Thus, the classification methodology has potential application as a management tool.
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Ecosystems may shift abruptly between alternative states in response to environmental perturbations [1-3]. Early warning indicators have been proposed to anticipate such regime shifts, but experimental field tests of their validity are rare [4-6]. We exposed rocky intertidal algal canopies to a gradient of press perturbations and recorded the response of associated assemblages over 7 years. Reduced cover and biomass of algal canopies promoted the invasion of algal turfs, driving understory assemblages toward collapse upon total canopy removal. A dynamic model indicated the existence of a critical threshold separating the canopy- and turf-dominated states. We evaluated common indicators of regime shift as the system approached the threshold, including autocorrelation, SD, and skewness [7]. These indicators captured changes in understory cover due to colonization of algal turfs. All indicators increased significantly as the system approached the critical threshold, in agreement with theoretical predictions [2, 8, 9]. The performance of indicators changed when we superimposed a pulse disturbance on the press perturbation that amplified environmental noise. This treatment caused several experimental units to switch repeatedly between the canopy- and the turf-dominated state, resulting in a significant increase in overall variance of understory cover, a negligible effect on skewness and no effect on autocorrelation. Power analysis indicated that autocorrelation and SD were better suited at anticipating a regime shift under mild and strong fluctuations of the state variable, respectively. Our results suggest that regime shifts may be anticipated under a broad range of fluctuating conditions using the appropriate indicator. Copyright © 2015 Elsevier Ltd. All rights reserved.
Article
Introduces the physical and chemical characteristics of the Baltic Sea, emphasising the almost complete absence of tides, the low salinities and the seasonal pulses in freshwater (governed by river inflow) and nutrient concentrations (governed by pelagic activities). Most of the Baltic marine organisms live close to their limit of tolerance to low salinity. Current environmental stress on the ecosystem becomes all the more significant with a paucity of potential replacement species. Communities on rocky substrates in the eulittoral and sublittoral, and on sandy and muddy substrates, are described. Comments are made on the distribution of biomass, both of plants (predominantly macroalgae) and animals (blue mussel Mytilus edulis comprising >90% of the biomass in most areas of the N Baltic). Functional aspects of the littoral system are reviewed. Grazing pressure on macroalgae is generally low because of the absence of large herbivores. -P.J.Jarvis
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Appropriate integration of remote sensing technologies into ecosystem services concepts and practices leads to potential practical benefits for the protection of biodiversity and the promotion of sustainable use of Earth's natural assets. The last decade has seen the rapid development of research efforts on the topic of ecosystem services, which has led to a significant increase in the number of scientific publications. This systematic review aims to identify, evaluate and synthesise the evidence provided in published peer reviewed studies framing their work in the context of spatially explicit remote sensing assessment and valuation of ecosystem services. Initially, a search through indexed scientific databases found 5920 papers making direct and/or indirect reference to the topic of “ecosystem services” between the years of 1960 and 2013. Among these papers, 211 make direct reference to the use of remote sensing. During the search we aimed at selecting papers that were peer-reviewed publications available through indexed bibliographic databases. For this reason, our literature search did not include books, grey literature, extended abstracts and presentations. We quantitatively present the growth of remote sensing applications in ecosystem services’ research, reviewing the literature to produce a summary of the state of available and feasible remote sensing variables used in the assessment and valuation of ecosystem services. The results provide valuable information on how remotely sensed Earth observation data are used currently to produce spatially-explicit assessments and valuation of ecosystem services. Using examples from the literature we produce a concise summary of what has been done, what can be done and what can be improved upon in the future to integrate remote sensing into ecosystem services research. The reason for doing so is to motivate discussion about methodological challenges, solutions and to encourage an uptake of remote sensing technology and data where it has potential practical applications.
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Dry/wet condition has a large interannual variability. Decision-makers need to know the onset, duration, and intensity of drought, and require droughts be monitored at a daily to weekly scale. However, previous tools cannot monitor drought well at this short timescale. The Palmer Drought Severity Index has been found dissatisfactory in monitoring because of its complexity and numerous limitations. The Standardized Precipitation Index (SPI) always asks for a timescale, and precipitation is averaged over the period of the scale. Because of this, the SPI cannot be used for short scales, e.g., several days, and what it tells is the overall drought situation of the period. The weighted average of precipitation (WAP) developed by Lu (Geophys Res Lett 36:L12707, 2009) overcomes the deficiency of the SPI; it does not require a timescale, and can provide the drought (and flood) extent of each day. Therefore, the WAP can monitor drought at scales from daily to weekly, monthly, and any longer scale, and is really “flexible and versatile for all timescales”. In this study, the standardized WAP (SWAP) is used to monitor the 2011 drought over China. Drought swept the country during the year from north to south and from east to west. In spring, a once-in-a-fifty-year drought occurred over the Yangtze River basin and the southern region, causing serious shortage of drinking water for people and livestock, as well as tremendous losses in agriculture and the shipping industry. Results show that the SWAP, with its monthly mean plots, can well reproduce the seasonal shift of the 2011 drought across the country. The animation of daily plots demonstrates that the SWAP would have been able to monitor the day-to-day variation of the spring drought around the Yangtze River basin. It can provide the details of the drought, such as when the drought emerged over the region, how long it maintained there (though drought area may move back and forth with extension and contraction of the area), and when the drought relieved over the basin.
Article
The use of conspicuous organisms as indicators of the limits of intertidal zones is examined for shores of the British Isles. Except in restricted areas the upper limit of laminarians provides an adequate boundary between the infralittoral fringe and the midlittoral zone. On the majority of shores which lack laminarians, suitable alternative forms usually exist and can be used. The use of the upper limit of barnacles to indicate the upper limit of the midlittoral zone is hampered by differences in the vertical distribution of the two barnacles which are present in different parts of the British Isles (Balanus balanoides and Chthamalus stellatus). The pattern of high-level zonation is described from areas possessing Chthamalus or Balanus alone, and from `transitional shores' on which both species are present at high levels. Possible methods of delimiting the zones by reference to organisms other than barnacles, or by reference to tidal levels, are discussed but considered to be generally unsuitable. The evidence available from many coastlines of the world suggests that although supra- and midlittoral zones can be universally recognized and are characterized by the same types of organisms, the exact level of the boundary between the zones must be expected to fluctuate to some extent from one shore to another, because of variation in the ability of the different species of barnacles (or comparable indicators) to tolerate the more arid conditions of the upper shore. The British Isles are considered to be an area of considerable complexity; but since no single method of delimiting the zones is applicable everywhere, it is proposed to retain the original demarcation line, the upper limit of barnacles, rather than to introduce any additional local features into the scheme of littoral zonation.
Article
A methodology to classify rocky shores along the North East Atlantic (NEA) region was developed. Previously, biotypes and the variability of environmental conditions within these were recognized based on abiotic data. A biological validation was required in order to support the ecological meaning of the physical typologies obtained. A database of intertidal macroalgae species occurring in the coastal area between Norway and the South Iberian Peninsula was generated. Semi-quantitative abundance data of the most representative macroalgal taxa were collected in three levels: common, rare or absent. Ordination and classification multivariate analyses revealed a clear latitudinal gradient in the distribution of macroalgae species resulting in two distinct groups: one northern and one southern group, separated at the coast of Brittany (France). In general, the results based on biological data coincided with the results based on physical characteristics. The ecological meaning of the coastal waters classification at a broad scale shown in this work demonstrates that it can be valuable as a practical tool for conservation and management purposes.
Article
Wind wave reanalyses have become a valuable source of information for wave climate research and ocean and coastal applications over the last decade. Nowadays, wave reanalyses databases generated with third generation models provide useful wave climate information to complement, both in time and space, the instrumental measurements (buoys and alimetry observations). In this work, a new global wave reanalysis (GOW) from 1948 onwards is presented. GOW dataset is intended to be periodically updated and it is based on a calibration of a model hindcast with satellite altimetry data, after verification against historical data. The outliers due to tropical cyclones (not simulated due to insufficient resolution in the wind forcing) are identified and not taken into account in the process to correct the simulated wave heights with the altimeter data. The results are validated with satellite measurements in time and space. This new calibrated database represents appropriately the wave climate characteristics since 1948 and aims to be the longest and up-to-date wave dataset for global wave climate variability analysis as well as for many coastal engineering applications.
Article
The Water Framework and Habitats Directives require the evaluation of both the conservation and ecological status of macroalgae communities at water body or habitat level. However, assessments of macroalgal communities are highly time-consuming, both in terms of sampling effort and laboratory processing. These constraints have brought about their oversight in many marine monitoring programs, especially in subtidal environments. By using data from intertidal and subtidal macroalgae assemblages of Mouro Island (North coast of Spain) we wanted to identify possible cost-effective methods for monitoring this biological indicator, based on both high taxa levels and use of representative taxa. Multivariate analyses were applied using different data transformations. The results show that macroalgal communities are robust to aggregation to genus or even family level. Moreover, the outcomes show that transformation types introduce higher variation in the multivariate pattern of samples than the taxonomic level to which organisms are identified. Also, the study supports the use of representative taxa as a surrogate to overall community structure. Therefore, we conclude that a rapid-assessment by means of field evaluations, based on coverage of representative taxa, is a reliable alternative for the assessment of macroalgae status. In addition this procedure allows evaluation at a broader spatial scale (water body or habitat level) than traditional quantitative sampling procedure does.
Article
The Northern Kattegat front separates Kattegat surface water or around 26 PSU and Skagerrak water of around 34 PSU. The front plays an important role in the water exchange between Kattegat and Skagerrak, and consequently it is also important for the exchange of dissolved and suspended matter such as nutrients, organic material and Gelbstoff. In order to investigate the dynamics of the Northern Kattegat front two surveys were performed, one in the period 2–12 March, 1992 and in one in the period 21 September–2 October, 1992. Vertical profiles of temperature, salinity and current were measured along cross-sections. The data from the two surveys and the analyses of the data concerning the dynamics of the Northern Kattegat front are presented. First, a conceptual model of the current dynamics, based mainly on Stigebrandt, A. (1987) (Tellus, 39A, 170–177), is proposed, where the front is considered to be a ‘rotational baroclinic control’ guiding the current in the Kattegat. The model is evaluated on the basis of the two surveys. Secondly, an equation describing the outflow from the Kattegat to the Skagerrak is derived and evaluated.
Article
A computationally efficient relocatable system for generalized inverse (GI) modeling of barotropic ocean tides is described. The GI penalty functional is minimized using a representer method, which requires repeated solution of the forward and adjoint linearized shallow water equations (SWEs). To make representer computations efficient, the SWEs are solved in the frequency domain by factoring the coefficient matrix for a finite-difference discretization of the second-order wave equation in elevation. Once this matrix is factored representers can be calculated rapidly. By retaining the first-order SWE system (defined in terms of both elevations and currents) in the definition of the discretized GI penalty functional, complete generality in the choice of dynamical error covariances is retained. This allows rational assumptions about errors in the SWE, with soft momentum balance constraints (e. g., to account for inaccurate parameterization of dissipation), but holds mass conservation constraints. While the dynamical calculations involve elevations alone, depth-averaged currents can be directly assimilated into the tidal model with this approach. The efficient representer calculation forms the basis for the Oregon State University (OSU) Tidal Inversion Software (OTIS). OTIS includes software for generating grids, prior model covariances, and boundary conditions; for time stepping the nonlinear shallow water equations to generate a first guess or prior solution; for preliminary processing of TOPEX/Poseidon altimeter data; for solution of the GI problem; and for computation of posterior error bars. Approximate GI solution methods, based on using a reduced set of representers, allow very large datasets to be inverted. OTIS regional and local GI tidal modeling (with grids containing up to 10(5) nodes) require only a few hours on a common desktop workstation. Use of OTIS is illustrated by developing a new regional-scale (1/68) model of tides in the Indonesian Seas.
Article
The intertidal vegetation of the 'Abra de Bilbao' (Basque coast, N Spain) was studied following a pollution gradient. Under the effect of pollution, several signs of alteration are detected in the vegetation, which responds by simplifying its structure. Species which require high quality environmental characteristics such as Cytoseira tamariscifolia, Cystoseira baccata, Bifurcaria bifurcata, Gelidium sesquipedale, Gigartina pistillata and Laurencia obtusa disappear, whereas Corallina elongata, Chondria coerulescens, Caulacanthus ustulatus and Gelidium pusillium become dominant in successive steps as pollution increases. In general, large perennial algae are replaced by small turfing algae. In addition, an intensification of the herbivorous activity (mainly limpets) occurs on the soft caespitose vegetation. Species richness and algal abundance suffer a reduction. In the most degraded environmental conditions, vegetation is replaced by filter-feeders (mussels and barnacles), and cyanobacteria proliferate. Pollution also alters the zonation pattern reducing in the number of vegetation belts.
Article
Mainly on the basis of the distribution patterns of 42 species of the recently revised genus Cladopkora (Chlorophyceae) in the north Atlantic Ocean, it appeared possible to distinguish 10 phytogeographic distribution groups of wide applicability. Experimentally determined critical temperatures limiting essential events in the life histories of 17 benthic algal species were used to infer possible phytogeographic boundaries; these appeared to fit closely the phytogeographic boundaries derived from field-distribution data. For a temperate species, at least six different boundaries can be postulated and should be checked in the northern hemisphere: (1) the ‘northern lethal boundary’ (corresponding to the lowest winter temperature which a species can survive); (2) the ‘northern growth boundary’ (corresponding to the lowest summer temperature which, over a period of several months, permits sufficient growth); (3) the ‘northern reproductive boundary’ (corresponding to the lowest summer temperature permitting reproduction over a period of several months); (4–6) the corresponding southern boundaries. Photoperiodic responses may influence the temperature responses. Many phytogeographic boundaries appear to be of a composite nature. For instance, the southern boundary of Laminaria digitata follows the European 10°C February isotherm (which corresponds to the highest winter temperature permitting fertility in the female gametophyte, i.e. to the ‘southern reproductive boundary’), and the American 19°C summer isotherm (corresponding to the ‘southern lethal boundary’). Thus, experimental evidence supports the validity of eight of the following 10 distribution groups (for distribution groups 2 and 6, such evidence could not be found): (1) the amphiatlantic tropical-to-warm temperate group, with a north-eastern extension (examples: Gracilaria foliifera and Centroceras clavulalum); (2) the amphiatlantic tropical-to-warm temperate group, with a north-western extension (example: Hypnea musciformis); (3) the amphiatlantic tropical-to-temperate group (example: Sphacelaria rigidula =furcigera); (4) the amphiatlantic temperate group: the Cladophora rupestris type (examples: Callithamnion hookeri, Dumontia contorta; Laminaria saccharina is transitional to type 10, I., digitata to types 5 and 10); (5) the amphiatlantic temperate group: the Cl. albida type (examples: Scytosiphon lomentaria, Petalonia fascia); (6) the tropical western Atlantic group; (7) the north-east American tropical-to-temperate group (example: Gracilaria tikvahiae); (8) the north-east American temperate group and the corresponding Japanese temperate group (examples: Campylaephora hypneoides and Sargassum muticum); (9) the warm-temperate Mediterranean-Atlantic group, and the corresponding warm-temperate Californian group (examples: Saccorhiza polyschides, Laminaria hyperborea, I., ockroleuca, Macrocystis pyrifera, Hedophyllum sessile); (10) the Arctic group (examples: Saccorhiza dermatodea and Sphacelaria arctica). Distribution groups 6, 9 and 10 have comparatively narrow temperature ranges with a span of 18 22°C between their lethal boundaries and of 5 12°C between their reproductive or growth boundaries. These narrow temperature ranges limit the species in these groups to the tropics; the temperate coasts on the eastern sides of the north Pacific and north Atlantic Oceans and in the southern hemisphere; and to the Arctic, respectively. The narrow temperature ranges of group 9 make the species in this group unfit for life on the western temperate coasts of the north Pacific and north Atlantic Oceans, where algae must cope with annual temperature fluctuations of more than 20°C. Conversely, algae in group 8 (containing the numerous Japanese endemic species) are characterized by wide temperature spans (e.g. 29°C between ‘lethal boundaries’, 12–19°C between ‘growth and/or reproductive boundaries’) and must be potentially capable of occupying wide latitudinal belts on temperate coasts along the east sides of the north Pacific and north Atlantic Oceans. Algae ‘escaped’ from Japan, such as Sargassum muticum, conform to this picture. Apparently Japanese algae do not have the capacity for long distance dispersal. The corresponding east American coasts (30–45 N) harbour very few endemic species, probably as a result of the adverse nature of these sediment coasts for benthic macroalgae and their functioning as a barrier to latitudinal displacements of the flora during glaciations. The remaining distribution groups (1,2,3,4,5,7) are characterized by wide temperature spans and wide distributions, often in both the Atlantic and Pacific Oceans and in both hemispheres. Six temperate species (in distribution groups 4, 5 and 9) with an amphiaequatorial distribution have similar winter-temperature maxima permitting reproduction and corresponding with winter isotherms of 15–17°C; their upper lethal temperatures are more dissimilar and correspond with summer isotherms of 20–30°C. Their amphiaequatorial distribution can be explained by assuming glacial temperature drops along east Pacific and east Atlantic equatorial coasts in narrow belts of intensified upwelling during the presumably intensified glacial circulation of the ocean gyres.
Article
Ambient UV radiation has substantially increased during the last decades, but its impact on marine benthic communities is hardly known. The aim of this study was to globally compare and quantify how shallow hard-bottom communities are affected by UV during early succession. Identical field experiments in 10 different coastal regions of both hemispheres produced a consistent but unexpected pattern: (i) UV radiation affected species diversity and community biomass in a very similar manner, (ii) diversity and biomass were reduced to a larger extent by UVA than UVB radiation, (iii) ambient UV levels did not affect the composition of the communities, and (iv) any UV effects disappeared during species succession after 2–3 months. Thus, current levels of UV radiation seem to have small, predictable, and transient effects on shallow marine hard-bottom communities.
Article
this is an unreviewed manuscript, primarily intended for informal exchange of information among ncep staff members MMAB Contribution No. 276.
Article
Wave exposure is one of the fundamental variables of the coastal environment and many coastal processes are affected by waves and wave-induced water motion. Directly or indirectly, waves affect the lives of many coastal macroalgae and seagrasses. Although the idea that waves can affect macrophyte distribution and biodiversity is not new, it remains poorly examined. We looked at how a gradient of wave exposure, determined using the surf similarity number, influences the species richness of macrophytes among phyla and functional form groups. We suggest that wave exposure is a biologically and statistically significant variable and that the species richness of most phyla and groups decrease with increasing exposure. More interestingly, the analysis suggests that the Phaeophyta and the thick-leathery functional form group, which are associated with important coastal habitats (i.e. submarine forests), increases in species richness as exposure increases. It appears that the effect of wave exposure varies with phyla and functional form group. Consequently, we reinforce the belief that studies that address the biodiversity and distribution of macrophytes must include wave exposure as a measured variable, to provide any meaningful synthesis of the ecology of these organisms.
Article
Experimentally determined lethal temperatures and temperatures limiting growth or reproduction in the life histories of 15 benthic algal species were used to infer possible phytogeographic boundaries in the North Atlantic Ocean. These appeared to correspond closely with phytogeographic boundaries based on distribution data. Many boundaries appeared to be of a composite nature. For instance, the southern boundary ofNemalion helminthoides is interpreted as a southern reproduction boundary on the N. Atlantic E. shore and a southern lethal boundary on the N. Atlantic W. shore. The northern boundary on both sides of the ocean is a northern reproduction boundary.N. helminthoides is a typical representative of the amphiatlantic temperate distribution group, to which seven other of the fifteen investigated species belong (Chondrus crispus, Desmarestia aculeata, D. viridis, Monostroma grevillei, Acrosiphonia arcta with a comparable composite southern boundary;Rhodochorton purpureum with a southern lethal boundary).Polysiphonia ferulacea andDictyota dichotoma are treated as representatives of the amphiatlantic tropical-to-warm-temperate distribution group, andP. denudata as representative of the amphiatlantic tropical-to-temperate group.P. harveyi belongs to the N.E. American temperate group and is bounded by a northern reproduction boundary and a southern reproduction boundary. This is one of the very few species endemic to N.E. America. This poor endemism is ascribed to the vast adverse sediment shores and their additional acting as barriers to glacial northsouth displacements of the flora; it is not related to the wide annual temperature fluctuations (>20 C) typical for N.E. America. The temperate algal flora of Japan, however, which is extremely rich in endemic species is subject to equally wide annual temperature fluctuations.Bonnemaisonia hamifera is such a Japanese endemic, which has been accidentally introduced into the North Atlantic Ocean where its life history seems to be disrupted: it is maintained mainly by vegetative propagation of the heteromorphic tetrasporophyte. The species of the warm temperate Mediterranean-Atlantic group are probably too stenothermous for life on N.E. American shores; they need annual temperature fluctuations<20 C.Acrosymphyton purpuriferum seems to belong to this group, but arguments are presented to unite this species withA. caribaeum and to range it under the amphiatlantic tropical-to-warm-temperate group.Clathromorphum circumscriptum belongs to the Arctic distribution group and has a southern reproduction boundary across the ocean along the 3 C February isotherm. This species is able to survive temperatures of about 20 C. Five amphiequatorial temperate species discussed in this paper and four in another related paper have similar maximum winter temperatures of 14–17 C (mean monthly values) allowing reproduction. Their amphiequatorial distribution can be explained by assuming similar low temperatures in the euphotic zone along E. Pacific and E. Atlantic equatorial coasts i.e. in narrow inshore belts of intensified upwelling during the presumably intensified glacial circulation of the ocean gyres.
Article
Experimentally determined ranges of thermal tolerance and requirements for completion of the life history of some 60 seaweed species from the North Atlantic Ocean were compared with annual temperature regimes at their geographic boundaries. In all but a few species, thermal responses accounted for the location of boundaries. Distribution was restricted by: (a) lethal effects of high or low temperatures preventing survival of the hardiest life history stage (often microthalli), (b) temperature requirements for completion of the life history operating on any one process (i.e. [sexual] reproduction, formation of macrothalli or blades), (c) temperature requirements for the increase of population size (through growth or the formation of asexual propagules). Optimum growth/reproduction temperatures or lethal limits of the non-hardiest stage (often macrothalli) were irrelevant in explaining distribution. In some species, ecotypic differentiation in thermal responses over the distribution range influenced the location of geographic boundaries, but in many other species no such ecotypic differences were evident. Specific daylength requirements affected the location of boundaries only when interacting with temperature. The following types of thermal responses could be recognised, resulting in characteristic distribution patterns: (A) Species endemic to the (warm) temperate eastern Atlantic had narrow survival ranges (between ca 5 and ca 25°C) preventing occurrence in NE America. In species with isomorphic life histories without very specific temperature requirements for reproduction, northern and southern boundaries in Eur/Africa are set by lethal limits. Species with heteromorphic life histories often required high and/or low temperatures to induce reproduction in one or both life history phases which further restricted distribution. (B) Species endemic to the tropical western Atlantic also had narrow survival ranges (between ca 10 and ca 35°C). Northern boundaries are set by low, lethal winter temperatures. Thermal properties would potentially allow occurrence in the (sub) tropical eastern Atlantic, but the ocean must have formed a barrier to dispersal. No experimental evidence is so far available for tropical species with an amphi-Atlantic distribution. (C) Tropical to temperate species endemic to the western Atlantic had broad survival ranges (<0 to ca 35°C). Northern boundaries are set by low summer temperatures preventing (growth and) reproduction. Thermal properties would permit occurrence in the (sub)tropical eastern Atlantic, but along potential “stepping stones” for dispersal in the northern Atlantic (Greenland, Iceland, NW Europe) summer temperatures would be too low for growth. (D) In most amphi-Atlantic (tropical-) temperate species, northern boundaries are set by low summer temperatures preventing reproduction or the increase of population size. On European shores, species generally extended into regions with slightly lower summer temperatures than in America, probably because milder winters allow survival of a larger part of the population. (E) Amphi-Atlantic (Arctic-) temperate species survived at subzero temperatures. In species with isomorphic life histories not specifically requiring low temperatures for reproduction, southern boundaries are set by lethally high summer temperatures on both sides of the Atlantic. None of the species survived temperatures over 30°C which prevents tropical occurrence. Species with these thermal responses are characterized by distribution patterns in which southern boundaries in Eur/Africa lie further south than those in eastern N America because of cooler summers. In most species with heteromorphic life histories (or crustose and erect growth forms), low temperatures were required for formation of the macrothalli (either directly or through the induction of sexual reproduction). These species have composite southern boundaries in the north Atlantic Ocean. On American coasts, boundaries are set by lethally high summer temperatures, on European coasts by winter temperatures too high for the induction of macrothalli. Species with this type of thermal responses are characterized by distribution patterns in which the boundaries in Eur/Africa lie further north than those in eastern N America because of warmer winters.