Seasonal energetic constraints in Mediterranean benthic suspension feeders: effects at different levels of ecological organization

The aim of this work is to examine the role of food as a constraining factor at different levels of ecological organization in benthic littoral ecosystems. In the search for patterns in ecological systems, it has recently been documented that seasonality in the dynamics of benthic suspension feeders (BSF) in the Mediterranean Sea is characterized by summer dormancy. We review recent studies on seston availability in the water column and feeding and respiration by BSF in the Mediterranean. The objective is to assess whether a pattern across particular studies exists that could provide evidence of food as a constraining factor. A pattern emerges from these organism‐level studies, one which indicates the seasonal occurrence of an energy shortage in the taxa exhibiting summer dormancy. This energy shortage is closely related to low food availability and suggests that an energetic constraint underlie the summer dormancy phenomenon. The seasonal occurrence of summer energy shortage also appears to affect the dynamics of BSF at population and community level. In this sense, in late summer 1999, a mass mortality event of BSF affected several hundreds of kilometers in the Ligurian Sea (NW Mediterranean). The fact that the mass mortality event occurred in late summer and especially affected the taxa that exhibit energy shortage – such as anthozoans and sponges – suggests, that energetic constraints may contribute to the understanding of the mass mortality event. The energy shortage phenomenon may provide a mechanism to understand how the occurrence of anomalous climatic conditions may have induced the mass mortality of some BSF taxa. This review points out the existence of a common energy shortage phenomenon mainly related to low food availability as an important determinant of the dynamics of most BSF taxa at organism level in the Mediterranean. But this determinant also affects the dynamics of BSF at population and community level. Therefore, the extant data point up the crucial role of food as a constraining factor for benthic littoral ecosystems in oligotrophic areas like the Mediterranean, with important seasonal variations in seston abundance and composition.     

Are Antarctic suspension-feeding communities different from those elsewhere in the world?

This paper reviews the trophic ecology of benthic suspension feeders in Antarctic shelf communities, studied within SCAR's EASIZ Programme, in comparison with published information from other seas. Dense benthic suspension-feeder communities capture large quantities of particles and may directly regulate primary, and indirectly, secondary production in littoral food chains. Most work has been performed in temperate and tropical seas; however, little is known about suspension feeders in cold environments. Recent studies on Antarctic littoral benthic suspension feeders suggest the period of winter inactivity may last only a few weeks. This contrasts with the hypothesis that in Antarctic communities there is a prolonged period of minimal activity lasting at least 6 months during the austral winter. Results from other oceans may explain how dense benthic communities could develop under such conditions. Alternative food sources, i.e. the "fine fraction", sediment resuspension, lateral advection and efficient food assimilation may play a significant role in the development of suspension-feeder dominated, very diversified, high biomass and three-dimensionally structured communities on the Antarctic shelf.     

Appearance and evolution of marine benthic communities in the Early Palaeozoic

The development of marine benthic communities in the Early Palaeozoic occurred mainly in the shallow water epicontinental seas. It included those stages of the Cambrian and Ordovician evolutionary radiations that were dominated by a high rate of morphogenesis, when new food and territory resources were not limited. This provided many opportunities for coadaptation of emerging organisms. At the time of the Cambrian radiation, the body plans of all animals were formed, while in the Ordovician, the maximum rank of emerging taxa did not exceed the level of class. The beginning of each radiation was explosive. Vendian benthic communities developed in cold seas and in the shallowest areas of warm seas, where organic matter from the surface layers was available at the bottom because of the absence of a thermocline. The Cambrian radiation began with the appearance of pelagic suspension feeders, because of which much of the primary production could penetrate the thermocline and settle at the bottom. This allowed the occupation of warmer seas and greater depths. At the same time, the productivity of the pelagic region sharply increased because of the emergence of positive feedback between the producers and consumers in the water, leading to increased water transparency and elongation of trophic chains. Arthropods, the first suspension feeders, were the launch group of the Cambrian radiation. Cambrian benthic suspension feeders could seize only the smallest particles, mostly bacteria, and dissolved organic matter. This food resource was contained in the thin bottom water layer. Therefore, the food grasping structures of all the Cambrian suspension feeders were near the bottom, without forming tiers. The Ordovician evolutionary radiation began with the launch of the Pelmatozoan echinoderms, which were the first benthic suspension feeders to begin feeding on plankton. The exploitation of this resource led to the creation of a 1-m tier above the bottom and an increase in their calcite productivity. Positive feedback emerged between the grounds and the community of its inhabitants and considerably changed the composition and diversity of grounds, which sharply increased the diversity of benthos. The appearance of positive feedback between different components of ecosystems resulted in explosive evolution in both the Cambrian and Ordovician.     

Seasonality of feeding activity in Antarctic suspension feeders

The feeding activity of four benthic suspension-feeding groups (bryozoans, hydroids, polychaetes and holothurians) was monitored in situ every month for a 2-year period at Signy Island in the maritime Antarctic. The bryozoans were monitored at species level, whereas the other taxa could be differentiated only to genus. A marked seasonal variation in feeding activity was observed in most taxa. Although environmental parameters such as sea water temperature, fastice duration and water column chlorophyll concentrations suggested that winter in the maritime Antarctic lasts for about 6 months, many animals ceased feeding only for a short period of 2 or 3 months around the middle of the austral winter (June/July). These suspension feeders must therefore be efficient at utilising the low concentration of the microplankton existing in the water column for much of the year. Comparison with environmental variables suggested several possible cues for changes in feeding activity, but these cues may differ between taxa. Photoperiod and changes in disturbance by water movement (both mediated by ice), and food concentration are likely to be important environmental cues for polar suspension feeders.     

Invaders are not a random selection of species

We assembled information on 119 species of freshwater macroinvertebrate invaders in North America and Europe, and compared them to all native freshwater species in North America and Europe. We tested whether the invaders were a random or selected group among taxa (phylum or class), water quality requirements, and feeding habit. We found that freshwater macroinvertebrate invaders are not a random selection of species, and are over-represented by molluscs and crustaceans, while taxa richness of native communities are dominated by insects. Over 35% of native species of aquatic invertebrates in North America are only able to live in areas with excellent or very good water quality, and are intolerant of organic pollution. In contrast, all invaders are tolerant of at least moderate amounts of organic pollution. There was a significant difference in the distribution of feeding habits between native species and invaders: collector-filterers (including suspension feeders) were 2.5–3 times more abundant, and predators were 3–4 times less abundant among invaders than among native invertebrates. The ongoing spread of exotic species affects the biodiversity of selected taxa, shifts communities toward greater tolerance of organic pollution and increases the numbers of suspension feeders, thereby enhancing benthic pelagic coupling in waterbodies with high densities of invaders. Because these processes are very similar in Europe and North America, we suggest that the observed patterns may have a common global effect.     

Intertidal and subtidal soft-bottom macro- and meiofauna of the Kongsfjord (Spitsbergen)

The intertidal and subtidal soft bottom macro- and meiofauna of a glacier fjord on Spitsbergen was studied after complete ice melt in June 2003. The abundances of the benthic fauna were within the range reported from estuaries and similar intertidal areas of boreal regions. The high proportion of juveniles in the eulittoral zone indicated larval recruitment from subtidal areas. The macrobenthic fauna can be divided into an intertidal and a subtidal community, both being numerically dominated by annelids. Deposit feeders were numerically predominant in intertidal sites, whereas suspension feeders were most abundant in the subtidal area. Among the meiofauna, only the benthic copepods were identified to species, revealing ecological adaptations typical for intertidal species elsewhere.     

Global diversity of marine isopods (except asellota and crustacean symbionts)

The crustacean order Isopoda (excluding Asellota, crustacean symbionts and freshwater taxa) comprise 3154 described marine species in 379 genera in 37 families according to the WoRMS catalogue. The history of taxonomic discovery over the last two centuries is reviewed. Although a well defined order with the Peracarida, their relationship to other orders is not yet resolved but systematics of the major subordinal taxa is relatively well understood. Isopods range in size from less than 1 mm to Bathynomus giganteus at 365 mm long. They inhabit all marine habitats down to 7280 m depth but with few doubtful exceptions species have restricted biogeographic and bathymetric ranges. Four feeding categories are recognised as much on the basis of anecdotal evidence as hard data: detritus feeders and browsers, carnivores, parasites, and filter feeders. Notable among these are the Cymothooidea that range from predators and scavengers to external blood-sucking micropredators and parasites. Isopods brood 10-1600 eggs depending on individual species. Strong sexual dimorphism is characteristic of several families, notably in Gnathiidae where sessile males live with a harem of females while juvenile praniza stages are ectoparasites of fish. Protandry is known in Cymothoidae and protogyny in Anthuroidea. Some Paranthuridae are neotenous. About half of all coastal, shelf and upper bathyal species have been recorded in the MEOW temperate realms, 40% in tropical regions and the remainder in polar seas. The greatest concentration of temperate species is in Australasia; more have been recorded from temperate North Pacific than the North Atlantic. Of tropical regions, the Central Indo-Pacific is home to more species any other region. Isopods are decidedly asymmetrical latitudinally with 1.35 times as many species in temperate Southern Hemisphere than the temperate North Atlantic and northern Pacific, and almost four times as many Antarctic as Arctic species. More species are known from the bathyal and abyssal Antarctic than Arctic GOODS provinces, and more from the larger Pacific than Atlantic oceans. Two areas with many species known are the New Zealand-Kermadec and the Northern North Pacific provinces. Deep hard substrates such as found on seamounts and the slopes are underrepresented in samples. This, the documented numbers of undescribed species in recent collections and probable cryptic species suggest a large as yet undocumented fauna, potentially an order of magnitude greater than presently known.     

Temperate and tropical algal-sea anemone symbioses

Abstract. In this review, we seek to develop new insights about the nature of algal‐sea anemone symbioses by comparing such associations in temperate and tropical seas. Temperate seas undergo pronounced seasonal cycles in irradiance, temperature, and nutrients, while high irradiance, high temperature, and low nutrients are seasonally far less variable in tropical seas. We compare the nature of symbiosis between sea anemones (= actinians) and zooxanthellae (Symbiodinium spp.) in both regions to test tropical paradigms against temperate examples and to identify directions for future research. Although fewer anemone species are symbiotic in temperate regions, they are locally dominant and ecologically important members of the benthic community compared to the tropics. Zooxanthella densities tend to be lower in temperate anemones, but data are limited to a few species in both temperate and tropical seas. Zooxanthella densities are far more stable over time in temperate anemones than in tropical anemones, suggesting that temperate symbioses are more resistant to fluctuations in environmental parameters such as irradiance and temperature. Light‐saturated photosynthetic rates of temperate and tropical zooxanthellae are similar, but temperate anemone hosts receive severely reduced carbon supplies from zooxanthellae during winter months when light is reduced. Symbiont transmission modes and specificity do not show any trends among anemones in tropical vs. temperate seas. Our review indicates the need for the following: (1) Investigations of other temperate and tropical symbiotic anemone species to assess the generality of trends seen in a few “model’ anemones. (2) Attention to the field ecology of temperate and tropical algal‐anemone symbioses, for example, how symbioses function under seasonally variable environmental factors and how zooxanthellae persist at high densities in darkness and winter. The greater stability of zooxanthella populations in temperate hosts may be useful to understanding tropical symbioses in which bleaching (loss of zooxanthellae) is of major concern. (3) Study of the evolutionary history of symbiosis in both temperate and tropical seas. Continued exploration of the phylogenetic relationships between host anemones and zooxanthella strains may show how and why zooxanthellae differ in anemone hosts in both environments.     

Protein, carbohydrate, lipid concentrations and HSP 70–HSP 90 (stress protein) expression over an annual cycle: useful tools to detect feeding constraints in a benthic suspension feeder

In the present paper we suggest an effect of seasonal variations in food availability on two ecophysiological parameters in a warm temperate benthic suspension feeder: the tissue concentrations of proteins, carbohydrates and lipids on the one hand, and the expression of stress proteins (HSP 70 and 90, inducible and/or constitutive) on the other hand. The concentrations of biomacromolecules have already been used to describe bentho-pelagic and reproductive processes, but this is the first time that stress protein expression is suggested to be directly related with food constraints in marine organisms. Paramuricea clavata (Cnidaria: Gorgonacea) express HSP 70 and 90 (constitutive and/or inducible) throughout the seasonal cycle, and HSP 70 levels are twice as high as the levels of HSP 90. In summer and autumn, when seston availability to suspension feeders was low, P. clavata showed low levels of carbohydrates and lipids, but high levels of HSPs expression. The levels of HSP 70 and 90 expression fit with negative exponential functions of carbohydrate and lipid concentrations. We suggest a direct effect of food availability on the studied ecophysiological parameters while the effect of temperature may be rather indirect. HSP expression as well as the tissue concentrations of carbohydrate and lipids may be used as biomarkers of environmental changes and seston availability to benthic suspension feeders.     

Dormancy in the Copepoda — an overview

Dormancy affects copepods in their anatomy, physiology, genetics, population biology, community ecology, evolution and local and geographic distribution. It is known from freeliving representatives of three copepod taxa, namely the Harpacticoida, Cyclopoida and Calanoida. Species showing dormancy occur in various realms and habitats, both freshwater and marine, being benthic, planktic or ice-dwelling. Depending on the taxon, dormancy occurs at various times of the year, prevailing in higher and temperate latitudes. Copepod dormancy is expressed in various ontogenetic stages, such as resting eggs, arrested larval development, juvenile and adult encystment, or arrested development of nonencysted copepodids or adults. Ecologically, dormancy is an energy saving trait, allowing the individual to bridge periods of environmental harshness. Adverse environmental conditions could be abiotic (e.g. desiccation, temperature, oxygen availability) or biotic in nature (e.g. food availability, predation). Diapause s. str. is initiated, maintained and terminated by triggering factors (e.g. photoperiod, temperature, chemical cues, population density/physiological factors). The dormant state and emergence patterns directly affect reproduction, population dynamics, community composition, coexistence and distribution of copepods, as well as the phenology of their predators and living food items. Populations having dormancy, in most cases belong to and affect communities of two realms: the water column and the bottom. Dormant stages may provide means for dispersal as well as for staying in special localities. The variability of dormancy permits flexible and complex life histories. Dormancy is subjected to and on the other hand affects copepod evolution.     

Pleistocene fish otoliths from the Mediterranean Basin: a synthesis

An overview of upper pPliocene and pleistocene otolith assemblages is compiled on the basis of both literature data and newly collected material from several sections located mainly in southern Italy. One hundred and five taxa are listed. Additional comments are provided for taxa subject to discussion. The composition and affinities of the Mediterranean Pleistocene otolith associations (consisting mainly of deep sea fishes) is checked against the available data for Pre-Messinian, Pliocene, and Recent Mediterranean fishes. This analysis is based on the recorded nominal species, but some taxa that could be identified at the generic level only are also included when they are relevant from a biogeographic point of view. Some new data on Piacenzian (middle Pliocene) and Gelasian (upper Pliocene) otoliths are also provided because this time interval still constitutes a major gap in the knowledge of the stratigraphic range of Plio-Pleistocene fish taxa. The Pleistocene deepwater fish fauna of the Mediterranean shows a markedly more oceanic character than the present-day one, a tendency that was all ready observed from early Oligocene till Pliocene times. Starting from the late Piacenzian, an increasing number of Atlantic taxa progressively invaded the Mediterranean including several subpolar and temperate forms. This invasion became stronger in the Gelasian. From this period up to the middle and late Pleistocene, the Mediterranean deepwater fish fauna (mesopelagic and benthopelagic) is enriched by taxa which today are typical for the north Atlantic cold deep water. This faunal shift seems to be correlated with the evolution of the Plio-Pleistocene Mediterranean paleoceanographic setting as well as with the global climatic deterioration.     

Seasonal Variability of Particulate Organic Matter in a Mountain Stream in Central Spain

The aim of this paper was to study the influence of environmental characteristics of the Mediterranean climate on seasonal variability of particulate organic matter abundance in a mountain stream. Coarse and fine fractions of both suspended and benthic particulate organic matter were determined on 14 occasions between February 1998 and November 1999 in a second‐order Mediterranean stream in Central Spain (Arroyo Mediano). Temporal variability of suspended organic matter followed a seasonal pattern, attributed to litter‐fall inputs, instream processing, and the hydrological regime. Suspended organic matter (SOM) and its seasonal variability fall well within the range reported for streams in temperate non‐Mediterranean deciduous forest. However, we found no seasonal trend in benthic organic matter (BOM) storage, and it seems that the amount of BOM remained fairly constant throughout the year. Reach retention (evaluated as the ratio between BOM and SOM per m²) was higher in summer during reduced stream flow, mainly due to coarse particulate organic matter storage. These observations do not differ from those reported for other headwater streams in temperate forested biomes, from which we conclude that there was no evidence of a Mediterranean influence on particulate organic matter dynamics in the Mediano stream, nor probably in other headwater Mediterranean streams. (© 2006 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

A method to identify estuarine macroinvertebrate pollution indicator species in the Virginian Biogeographic Province

Macroinvertebrates are commonly used as biomonitors to detect pollution impacts in estuaries. The goal of this research was to identify estuarine benthic invertebrates that could be used as indicator species to detect presence or absence of pollution in the Virginian Biogeographic Province using available monitoring data from the U.S. Environmental Protection Agency's Environmental Monitoring and Assessment Program. The data were first subdivided by habitat. Then we summarized the benthic community using principal coordinates analysis and projected the results onto a pollution gradient. Then we compared medians and corrected interquartile ranges to identify indicator species that were sensitive to pollution (or lack thereof) and showed low variation in response. Our data set was divided into smaller subsets that ranged from 108 to 532 stations per habitat. Out of 707 species in the Virginian Province, 67 were identified as pollution indicator species; 37 pollution sensitive taxa and 30 pollution tolerant taxa. The pollution sensitive taxa were represented by more phyla than were the pollution tolerant taxa. The pollution tolerant taxa were dominated by deposit feeders while the pollution sensitive taxa were not dominated by animals using any one feeding strategy. The pollution sensitive taxa included many crustaceans, known to be sensitive to pollution, while the pollution tolerant taxa included many tubificid oligochaetes, which are common indicators of pollution. Our empirically based results corresponded well with other studies that identified tolerance values or indicator species. One advantage of this technique is that it can be used on smaller data sets, assuming that there are not major habitat differences among the samples. We believe that this parsimonious technique can be applied to other coastal areas where mid-size (100–500 stations) monitoring data sets are available.     

Effects of the duration of cold stratification on early life stages of the Mediterranean alpine plant Silene ciliata

Cold stratification provided by snow cover is essential to break seed dormancy in many alpine plant species. The forecast reduction in snow precipitation and snow cover duration in most temperate mountains as a result of global warming could threaten alpine plant populations, especially those at the edge of their species distribution, by altering the dynamics of early life stages. We simulated some effects of a reduction in the snow cover period by manipulating the duration of cold stratification in seeds of Silene ciliata, a Mediterranean alpine specialist. Seeds from three populations distributed along an altitudinal gradient were exposed to different periods of cold stratification (2, 4 and 6 months) in the laboratory and then moved to common garden conditions in a greenhouse. The duration of the cold stratification treatment and population origin significantly affected seed emergence percentage, emergence rate and seedling size, but not the number of seedling leaves. The 6‐month and 4‐month cold stratification treatments produced higher emergence percentages and faster emergence rates than seeds without cold stratification treatment. No significant cold stratification duration x seed population origin interactions were found, thus differential sensitivity to cold stratification along elevation is not supported.     

Humidity-regulated dormancy onset in the Fabaceae: a conceptual model and its ecological implications for the Australian wattle Acacia saligna

Background and aims

Seed dormancy enhances fitness by preventing seeds from germinating when the probability of seedling survival and recruitment is low. The onset of physical dormancy is sensitive to humidity during ripening; however, the implications of this mechanism for seed bank dynamics have not been quantified. This study proposes a model that describes how humidity-regulated dormancy onset may control the accumulation of a dormant seed bank, and seed experiments are conducted to calibrate the model for an Australian Fabaceae, Acacia saligna. The model is used to investigate the impact of climate on seed dormancy and to forecast the ecological implications of human-induced climate change.

Methods

The relationship between relative humidity and dormancy onset was quantified under laboratory conditions by exposing freshly matured non-dormant seeds to constant humidity levels for fixed durations. The model was field-calibrated by measuring the response of seeds exposed to naturally fluctuating humidity. The model was applied to 3-hourly records of humidity spanning the period 1972–2007 in order to estimate both temporal variability in dormancy and spatial variability attributable to climatic differences among populations. Climate change models were used to project future changes in dormancy onset.

Key Results

A sigmoidal relationship exists between dormancy and humidity under both laboratory and field conditions. Seeds ripened under field conditions became dormant following very short exposure to low humidity (<20 %). Prolonged exposure at higher humidity did not increase dormancy significantly. It is predicted that populations growing in a temperate climate produce 33–55 % fewer dormant seeds than those in a Mediterranean climate; however, dormancy in temperate populations is predicted to increase as a result of climate change.

Conclusions

Humidity-regulated dormancy onset may explain observed variation in physical dormancy. The model offers a systematic approach to modelling this variation in population studies. Forecast changes in climate have the potential to alter the seed bank dynamics of species with physical dormancy regulated by this mechanism, with implications for their capacity to delay germination and exploit windows for recruitment.     

Paleolatitudes in the Devonian of Brazil and the Frasnian-Famennian mass extinction

Differences in faunal composition of contemporary Devonian marine communities in Brazilian sedimentary basins suggest variation because of climatic gradients. Temperatures may have ranged from subarctic conditions in the Parana Basin, a type of Devonian Hudson's Bay, to temperate in the Amazon and adjoining basins, analogous to a modern “north Atlantic” climate. These gradients were paralleled in other parts of South America. The Malvinokaffric cold climate fauna was dominated by groups oforganisms that survived the worldwide late Devonian mass extinctions. Invertebrate groups absent in the Malvinokaffric regions of Brazil, Bolivia and Argentina (i.e. those restricted to Devonian equatorial belts of North America, Eurasia and Australia) were decimated in the late Devonian. These two parallel observations suggest that late Devonian extinction may have been the result of drastic cold spells that killed off reefal and peri-reefal life. By elimination of crucial benthic community components, the trophic structure of the shallow marine ecosystem of the time was upset. Subsequent repopulation of the Carboniferous seas was accomplished by hardy, eurythermal invertebrate taxa present in cold as well as tropical regions. Cold spells and mass mortality in very shallow waters may go some way to explain the production of the widespread black shale environments so typical of marine regressive phases in the late Devonian of the western hemisphere.     

Shifts in Antarctic megabenthic structure after ice-shelf disintegration in the Larsen area east of the Antarctic Peninsula

The aim of this study was to contribute to a general understanding of the response of the Antarctic macrobenthos to environmental variability and climate-induced changes. The change in population size of selected macrobenthic organisms was investigated in the Larsen A area east of the Antarctic Peninsula in 2007 and 2011 using ROV-based imaging methods. The results were complemented by data from the Larsen B collected in 2007 to allow a conceptual reconstruction of the environment-driven changes before the period of investigation. Both Larsen areas are characterised by ice-shelf disintegration in 1995 and 2002, respectively, as well as high inter-annual variability in sea-ice cover and oceanographic conditions. In 2007 one ascidian species, Molgula pedunculata, was abundant north and south of the stripe of remaining ice shelf between Larsen A and B. Population densities decreased drastically in the Larsen A between 2007 and 2011, coincident with the decrease in Corella eumyota, another ascidian. Among the ophiuroids, the population of deposit feeders increased, while suspension feeders halved their abundance. Current measurements indicated a northward flow between the Larsen B and Larsen A, suggesting that a major physical forcing on benthic population development comes from the South. The results demonstrate that Antarctic macrobenthic populations can exhibit dramatic population dynamics. Analyses of sea-ice dynamics, salinity, temperature and surprisingly ice-shelf disintegration history, however, did not provide any clear evidence for environmental drivers underlying the apparent changes.     

Nord‐ und Südpolarmeer im Klimawandel. Ein biologischer Vergleich

Impacts of climate change on polar seas The polar seas in the Arctic and Antarctic are characterized by extreme cold and the prevalence of sea ice, which provides a unique polar habitat but also strongly affects the pelagic and benthic biota beneath. Life conditions for the marine fauna and flora differ considerably between the Arctic and Southern Oceans, as a result of contrasts in geography, geological history, as well as seasonal dynamics in light regime, sea ice cover and, hence, biological production. Climate change is particularly obvious in the Arctic Ocean and off the Antarctic Peninsula where warming results in a rapid shrinkage of the summer sea ice cover. Such decline threatens the sea‐ice communities and their associated fauna and will also have far reaching effects for the plankton and benthos of the polar seas.     

Clade perseverance from Mesozoic to present: a multidisciplinary approach to interpretation of pattern and process

Two clades of marine bryozoans, cyclostomes and cheilostomes, exemplify the benefits of applying a multidisciplinary approach to the interpretation of long-term evolutionary patterns. The cyclostome bryozoans were dominant in the Mesozoic; since that era, they have decreased in absolute terms and the cheilostomes have come to exceed them in both abundance and diversity. Many studies of living assemblages of the encrusting members of these two clades indicate that cheilostomes are superior space competitors, but paleontological studies suggest that competition between the two taxa has not been escalating over geological time. Both clades occur throughout the world's oceans and seas, and recent work in the geographical extremes has shown that the relative success of the clades varies markedly from place to place. In this study, the importance of differential patterns of recruitment and cumulative space occupation in the two clades was evaluated over four years and in two environments, one temperate and one polar. In both of these environments, peaks of recruitment and space occupation by the two clades were out of phase. The different strategies and outcomes of spatial competition are examined, largely using data from the literature. Only recently has it been realized that tied outcomes of competition are stable alternative results and not simply transitory phases. Many competitive encounters involving cyclostomes result in ties, implying that their strategy is based on persistence rather than dominance. When different indices and models are used to analyze competition data from the two clades, the interpretation varies markedly with methodology. The differences in patterns of recruitment, space occupation, and spatial competition have influenced both our understanding of how the two clades have persisted alongside each other and our perception of cheilostome superiority. Analysis of fluid dynamics has shown that small differences in the mechanical structure of typical members of each clade lead to fundamental differences in water movement. For animals that rely on water motion for transport of nutritional and excretory elements (suspension feeders), small changes in current velocity and direction can have a major impact. Preliminary chemical analysis of the excurrent stream leaving cheilostome colonies has shown it to be laden with excretory products, which can interfere and mix with a neighbor's feeding currents. Clearly, spatial competition involves more than a simple mechanical "showdown."     

Effects of dredging on critical ecological processes for marine invertebrates,seagrasses and macroalgae,and the potential for management with environmental windows using Western Australia as a case study

Dredging can have significant impacts on benthic marine organisms through mechanisms such as sedimentation and reduction in light availability as a result of increased suspension of sediments. Phototrophic marine organisms and those with limited mobility are particularly at risk from the effects of dredging. The potential impacts of dredging on benthic species depend on biological processes including feeding mechanism, mobility, life history characteristics (LHCs), stage of development and environmental conditions. Environmental windows (EWs) are a management technique in which dredging activities are permitted during specific periods throughout the year; avoiding periods of increased vulnerability for particular organisms in specific locations. In this review we identify these critical ecological processes for temperate and tropical marine benthic organisms; and examine if EWs could be used to mitigate dredging impacts using Western Australia (WA) as a case study. We examined LHCs for a range of marine taxa and identified, where possible, their vulnerability to dredging. Large gaps in knowledge exist for the timing of LHCs for major species of marine invertebrates, seagrasses and macroalgae, increasing uncertainty around their vulnerability to an increase in suspended sediments or light attenuation. We conclude that there is currently insufficient scientific basis to justify the adoption of generic EWs for dredging operations in WA for any group of organisms other than corals and possibly for temperate seagrasses. This is due to; 1) the temporal and spatial variation in the timing of known critical life history stages of different species; and 2) our current level of knowledge and understanding of the critical life history stages and characteristics for most taxa and for most areas being largely inadequate to justify any meaningful EW selection. As such, we suggest that EWs are only considered on a case-by-case basis to protect ecologically or economically important species for which sufficient location-specific information is available, with consideration of probable exposures associated with a given mode of dredging.