The Palaeobiogeography Of Silurian Ramphoprionid Polychaete Annelids

Silurian palaeobiogeography of the Ramphoprionidae (Annelida; Polychaeta) is outlined. Members of the family are reported from northern Europe (Estonia, Sweden and England), the Russian Arctic (Severnaya Zemlya), and the Canadian Arctic (Baillie–Hamilton Island). These areas were located fairly close to the palaeoequator during the Silurian and a faunal exchange is indicated by the broad, intercontinental distribution of the family. A few taxa may, however, have been endemic. Ramphoprionids had long stratigraphical ranges and were generally fairly rare during the Silurian, infrequently exceeding ten per cent of the jawed polychaete faunas.     

Taphonomic implications of polychaete jaws recovered from gut contents of the queen triggerfish

Remains of jawed polychaetes were recovered from stomach contents of queen triggerfish ( Balistes vetula L.) collected by spearing from Came Bow Cay, Belize, Central America. Jaws in the nereidids, glycerids, and arabellids recovered are perfectly preserved, and might be passed by the fish without noticeable alteration. Jaws in the onuphids are fully articulated, but the outer wall-layer has been digested away, leaving only the inner aragonitic layer intact. A low fossilization potential for onuphid jaws is indicated, consistent with laboratory studies of jaw composition. Based on literature accounts, fish predation may have a considerable influence on polychaete jaw (scolecodont) assemblages in sediment.     

Polychaetes as surrogates for marine biodiversity: lower taxonomic resolution and indicator groups

Due to conservation needs, reliable rapid-assessment methods for mapping of biodiversity are needed. One approach is to use surrogates, i.e. quantities that correlate strongly with the number of species, but are easier to obtain. The purpose of this paper is to test two polychaete surrogates, one for higher taxa and one for indicator groups, that will facilitate prediction of species richness in marine soft-bottom communities. Soft sediment is an important habitat which covers most of the ocean bottom. Data on polychaetes from the North Atlantic were used since polychaetes are often numerically dominant in the benthic assemblages, both with regard to the number of species and their abundance. In the polychaete assemblages along the Norwegian coast, richness at the genus, family and order level were significantly, linearly correlated to total species richness (r 0.92). Polychaetes in the order Terebellida were found to be a good indicator of polychaete species richness and to a lesser extent also of whole benthic assemblages. The group Terebellida is potentially well suited as an indicator group, since it contains long-lived, large species that are easy to sort from the sediment and it is well defined taxonomically. Although promising as proxies for species richness in marine biodiversity studies, the use of lower taxonomic resolution and indicator groups requires further investigations in more local areas where there are conservation issues.     

Epibiontic and endobiontic polychaetes of Geodia cydonium (Porifera,Demospongiae) from the Mediterranean Sea

Polychaete assemblages associated to the sponge Geodia cydonium were investigated at two sampling sites in the Mediterranean Sea: Porto Cesareo Basin (Apulia) and Marsala Lagoon (Sicily), both characterized by sheltered hydrodynamic conditions. Samples were seasonally performed during 1997, in order to compare the assemblages coming from the two localities studied, considering separately the internal and external tissues of the sponge, and with the aim of evaluating the influence of sponge size on polychaete colonization. The examined sponge is characterized by a peculiar stratification of its tissues: an external thick and hard layer, the cortex, and an internal softer one, the choanosome. Statistical analysis showed that this was the main factor controlling polychaete assemblage, with the internal tissue, less rich and diversified, appearing impoverished with respect to the external layer. A similarity in species composition was observed between sites, even though some differences were evidenced in the abundance of some species, mainly reflecting differences in local environmental conditions. Species richness and density increased with the increasing sponge size. Such a situation is particularly evident at Porto Cesareo, where sponges are covered by an algal layer which is particularly rich on the largest specimens, thus suggesting that most of the species of polychaetes were linked more to the neighbouring environment than to the sponge itself.     

Polychaete assemblages and sediment pollution in a harbour with two opposing entrances

The harbour at Ceuta is one of the most important harbours in the Strait of Gibraltar. The sediments are moderately polluted with organic matter and heavy metals but the harbour has two opposing entrances and a connecting channel which increases water renewal and dissolved oxygen across the harbour. For these special conditions, the value of the soft bottom polychaete community as a bioindicator, and possible advantages of the presence of two harbour entrances on biotic assemblages, were studied. Twenty-one stations were selected, and 27 variables were measured in the sediment of each station. The polychaete species richness and Shannon diversity values were similar inside and outside the harbour. Nevertheless, the Pielou evenness index was significantly higher in the external stations due to high densities of some species of polychaetes such as Pseudomalacoceros tridentata and Capitella capitata inside the harbour. The multivariate approach based on polychaete species composition was much more sensitive than univariate analyses at discriminating between internal and external stations. The pollution gradient and granulometric parameters were the main factors affecting polychaete distribution. Polychaete species richness and diversity in sediments inside Ceuta harbour were higher than in conventional harbours due to the positive effects of increased water renewal. These results should be taken into consideration in design, construction and remodelling of future harbours.Communicated by: H.-D. Franke     

Spatiotemporal variation in benthic polychaetes (Annelida) and relationships with environmental variables in a tropical estuary

Annelida constitute a dominant functional component in soft-bottom macrobenthic communities and reveal a wide range of adaptability to different marine and coastal habitats. Analyses in different polychaete assemblages and their responses to habitat conditions reflect the biological effects of marine pollution and habitat disturbance. The present study is designed to study colonization and community structure of polychaetes in two ecologically distinct locations of the Sundarban Biosphere Reserve on the northeast coast of India. Polychaete assemblages are characteristically different at the two sites in the extreme northern (Ghusighata) and southern (Gangasagar) portions of the Biosphere Reserve. Levels of heavy metals in polychaete body tissues also reveal interspecific and regional variations. The predominant polychaete fauna exhibited a distinct and unique assemblage of two types: (i) Mastobranchus indicus – Dendronereides heteropoda in the sewage-fed substratum at Ghusighata and (ii) Lumbrinereis notocirrata – Ganganereis sootai – Glycera tesselata at Gangasagar at the mouth of the Hugli estuary where chronic anthropogenic stress and contamination with agricultural and industrial effluents occur. The faunistic composition of polychaetes and their potential for the accumulation of heavy metals from the ambient medium are distinctly different. The study demonstrates that textural composition of the sediments, together with hydrodynamic and geotechnical properties, seem to have the greatest control to quantify the differences of the polychaete community in the two study stations. An in-depth comparative study of polychaete community structure at multiple spatial scales is strongly recommended for future environmental impact assessment in this fragile environment.     

Biodiversity and distribution of polychaetes and molluscs along the Dese estuary (Lagoon of Venice, Italy)

Mollusc and polychaete distribution in the mud flats along the Dese estuary (lagoon of Venice, Italy) was analysed in May, August and October to evaluate taxonomic biodiversity and zonation pattern of soft macrobenthos in an area of the lagoon hardly known from this point of view. Throughout the studied area most of the species were polychaetes, particularly Streblospio shrubsolii (Buchanan), which showed greater abundance from May to October. Molluscs and polychaetes showed a zonation of marine species which colonize brackish environments in the outer and intermediate areas of the estuary and of brackish water species in the inner one. In the outer area different assemblages were attributed to different sediment types. This distribution pattern, put in relation to the distance from the sea, was evident in May and partly blurred in August and October: the assemblages of the inner area of the estuary invaded the intermediated area in August and affected outer area in October because of spreading of the brackish water species S. shrubsolii, which became dominant. The distribution pattern of molluscs and polychaetes can be explained in terms of confinement as the leading factor influencing water quality and dynamics of the ecosystem. It is also needed to be taken into consideration the role played by competition due to different degree of opportunism of the most abundant species.     

Distribution of polychaete feeding guilds in sedimentary environments of the Campeche Bank, Southern Gulf of Mexico

The aim of this study was to analyze the trophic structure of the polychaete assemblages found in the Campeche Bank, southern Gulf of Mexico and to examine the effect of the sediment composition on the spatial distribution of the feeding guilds. In all, 2,662 organisms belonging to 160 species and 16 feeding guilds were identified. Filter-feeders (Fabricinuda trilobata and Bispira melanostigma) dominated. Five groups of stations were defined based on feeding guilds: one, in the southwest, characterized by motile jawed burrowers (17.14% contribution); the second, from the southeast to the northwest, characterized by seven guilds (45.25%), mainly filter-feeders and surface deposit-feeders; the third, in the southwest, characterized by three guilds (42.13%), mainly discretely motile tentaculate filter-feeders and motile unarmed burrowers; group four, in the east, was characterized by sessile tentaculate filter-feeders (63.68%); and group five, in the center and to the north, was characterized by four guilds (53.69%), mainly discretely motile tentaculate filter-feeders. The variety of feeding guilds was higher in the northwest with seven guilds, and the lowest variety was found in the east and south with only one or two guilds. Contrary to the starting hypothesis, the sediment composition was not the main factor that determined the distribution of the polychaete feeding guilds. Instead, salinity and depth were more important for the spatial arrangement of the trophic groups. The feeding guilds of polychaetes proved to be more sensitive to environmental changes than density or diversity.     

Les Pectinidae miocènes des faluns (Ouest de la France). Intérêts biostratigraphiques des associations

The Miocene Pectinidae from the “faluns” (W France). Biostratigraphical interests of the associations. New sampling was realized throughout neogene outcrops occurring in Western France (mainly, “faluns” of Langhian to Tortonian age from Touraine, Anjou and Blésois). It has been known for long that the species or associations of the representatives of the family Pectinidae vary in each outcrop, but it was still necessary to get original data, which were collected precisely in their original level. Such a work is still in progress in Mediterranean areas. It seems that the systematic framework of several species has still to be detailed; the recently re-discovered species may be 12 in number. The species may characterize four main assemblages: rich and poorly diversified faunas, collected in microconglomerates, deposited in shallow environments; rich and diversified association, corresponding to the bryozoa facies deposited in deeper environments (“savignean” facies Auct.); poor assemblage, collected in shallow water, shelly beds (“pontilevian” facies Auct.); nearly absent Pectinidae in the shallow water Arca facies (“lublean” Auct.). Generally, the species belonging to the genus Pecten are only common in the savignean facies, in which the diversity consequently increases. Actually, the biochronological framework of the “faluns” is correct, mainly related to the mammal zonation. The associations of Pectinidae collected in the “faluns” may be of several interests.

• •Several memberships of the observed faunas have a long stratigraphical range (Langhian–Pliocene), wide geographical distribution (from Britain to Mediterranean areas), and may be ecologically tolerant. They belong to the Crassadoma multistriata and Aequipecten radians species. In Western France, A. radians may be related to quite shallow environment. Opposite, the species Gigantopecten ligerianus may be related to the deepest environments.
• •Several species or association may success to each other through time, which may offer a regional stratigraphical interest, in the “faluns” from Western France: P. subarcuatus—P. sp. 1 (Langhian), P. sp. 2 (close to praebenedictus, Serravalian to Tortonian.), P. praebenedictus s. s. (younger Tortonien), P. jacoabeus or maximus (Messinian to Pliocene). G. ligerianus was recently recollected regionally in sediments of Langhian age only, if the local deposition environments are deep enough. Note that numerous shells belonging to this species exist in historical collections, from Doué area (obs. pers. M.B.).

     

Polychaete diversity in the Scotia Arc benthic realm: Are polychaetes tracers for faunal exchange?

The Scotia Arc is the only shallow-water and island bridge linking nowadays Patagonia and the Antarctic. The Antarctic Circumpolar Current as an oceanographic peculiarity makes this region an interesting biogeographic transition zone, because this frontal system traditionally is said to isolate the Antarctic fauna from that of the adjacent northern ecosystems. Based on benthos samples from three expeditions onboard R/V Polarstern, we studied distribution patterns of 200 polychaete species and 34 major benthic taxa in order to evaluate the role of polychaetes in the benthic realm of this part of the Southern Ocean. ANOSIM test distinguished three station groups: the central eastern Scotia Sea, the continental shelf off South America and stations at the tip of the Antarctic Peninsula. These station groups differed in organism densities and diversities with stations at the tip of the Antarctic Peninsula hosting the most diverse and dense community. The polychaete diversity patterns in the three assemblages evidenced closer connectivity between the tip of the Antarctic Peninsula and the central eastern Scotia Sea than between the continental shelf off South America with either the stations off the tip of the Peninsula or the central eastern Scotia Sea. This is probably supported by the Polar Front, which divides the island chain into two branches. Species distribution and community patterns of polychaetes appear to be associated with oceanographic and sediment conditions in this region. Most of the shared species showed the capability to tolerate differences in hydrostatic pressure. We suggest that the islands of the Scotia Sea may constitute a bridge for exchange of benthic species, particularly for polychaetes with eurybathic distribution and high dispersal capabilities.     

Polychaetes and oligochaetes associated with intertidal rocky shores in a semi-enclosed industrial and urban embayment,with the description of two new species

The species composition and relative abundance of the annelid benthic macrofauna (Polychaeta and Oligochaeta) inhabiting the rocky intertidal zone of the ria of Ferrol (Galicia, NW Spain) were studied during field collections, from 2000 to 2002. A total of 14,619 specimens (11,585 polychaetes belonging to 76 species and 24 families and 3,034 oligochaetes belonging to 18 species and two families) were collected from 98 quantitative samples taken from 13 sampling sites. The general spatial distribution of the annelid fauna reflects an increase of the diversity from the inner to the outer part of the ria. The general patterns found in the annelid composition suggest that the assemblages were dominated by oligochaetes in the inner sheltered sampling sites and polychaetes in the outer more exposed sites. Several faunistical and taxonomical remarks on selected species are presented. Two new species of oligochaetes are described: Coralliodrilus artabrensis sp. n. and Pirodrilus fungithecatus sp. n. (Naididae, Phallodrilinae). A new biological index, based on the oligochaete/polychaete ratio (O/P), is proposed as tool to evaluate environmental quality and to monitor future changes in the environment.     

Fossil plankton depth zones

Varied evidence from the known depth correlated distribution of benthic shelly facies communities in the Ordovician and Silurian clearly shows that graptolite taxonomic diversity increased from the nearshore regions to the shelf margin regions, as well as into the deeper portions of epicontinental basinal areas. We then consider a method for determining the depth distribution of individual graptolite taxa. This method depends on correlating the first appearance of graptolite taxa as one departs from shoreline in terms of the underlying benthos, arrayed in benthic assemblages. The first appearance departing from shoreline corresponds with the upper depth limits of each taxon in terms of the underlying depth indicated by the underlying, associated benthos. The depth of maximum abundance for each taxon should correspond to that point in the increasing relative abundance of that taxon where relative increase in abundance ceases. The lower depth limit of each taxon should correspond to that point on the relative abundance curve where increase in total relative abundance ceases.Using information about the stratigraphic ages indicated by the graptolite faunas, zone-by-zone, combined with that provided by the communities present within the associated and formerly underlying benthic assemblages should provide a more refined stratigraphic zonation than that obtained without combining both.     

Facies dependence of spore assemblages and new data on sedimentary influence on spore taphonomy

Palynological studies have been undertaken on Lower Carboniferous (Mississippian) rocks encountered in boreholes from the North Sea to the Baltic Sea (Rügen) in Northern Germany and from an outcrop at the northern border of the Rhenish Slate Mountains in western central Germany. The studied sections belong to different facies realms within the neritic shelf facies. They begin with very near shore coastal clastic facies in the North Sea and onshore Northern Germany, ranging into carbonate platform facies at Rügen, and then into platform slope facies at the northern margin of the Rhenish Slate Mountains. The focus of this study is the analysis of the relationship between sedimentary facies and spore assemblages; therefore, spores were considered only as sedimentary particles. Thus, it was necessary to create a new spore classification scheme based only on relevant sedimentary features. The new scheme distinguishes three classes of miospore based on shape, with each class subdivided into three subclasses based on features of ornament. The diversity of the spore assemblages and the total palynofacies have also been studied. Correlating palynological results from sections representing different sedimentary facies has led to a better understanding of the influence of sedimentary facies on spore assemblages. Correlations have been made at two stratigraphical levels. Regional correlation is undertaken on the Upper Viséan from the coastal clastic facies (North Sea, Northern Germany), the carbonate platform facies (Rügen) and the marginal platform slope facies (Velbert). Local correlation within the carbonate platform facies has been attempted using samples of Mid-Viséan TS zone from four boreholes on Rügen. Certain important differences have been observed. The local correlation at Rügen shows a decrease in spore frequency and spore assemblage diversity related to increasing resedimentation within subsided blocks of the platform. The composition of spore assemblages, strongly dominated by round-shaped spores, is nearly the same in all sections. The regional correlation shows clear proximal–distal trends for the total number of spores and also the diversity and composition of spore assemblages. A strong decrease in triangular and transitional spores from proximal to distal facies was observed. In conclusion, spore assemblages seem to be strongly influenced by sedimentary facies as well as by different sedimentary processes within the facies. Facies dependence of spore assemblages is shown to be an important factor for stratigraphical use of spores and for establishing vegetational and climatic trends.     

Distribution and diversity of polychaetes collected by trawling in Admiralty Bay: an Antarctic glacial fiord

The aim of the study is to determine patterns of distribution, species richness and diversity of polychaetes collected by trawling in Admiralty Bay. Forty-seven species of polychaetes were found in 75 samples collected in 1988 along a depth range from 15 to 500 m. Four polychaete assemblages were distinguished. The most diverse and peculiar assemblage was found in the deepest sublittoral of the central basin from ca. 300–500 m with main constant species Laetmonice producta, Sternaspis sp. and Thelepus cincinnatus. Shallow water assemblages had low diversity and species richness. The most common and eurytopic epifaunal polychaetes were Aglaophamus trissophyllus, Barrukia cristata, Harmothoe spp., Flabelligera mundata and Perkinsiana littoralis. The low species richness in shallow waters of Admiralty Bay is probably associated with depth, influence of glacial streams and suspended matter inflow which is particularly distinct in the shallow water fiord areas.     

福建沿岸红树林湿地多毛类生态分布

根据2009年至2012年在福建沿岸5块典型红树林湿地所作的调查资料,分析了福建沿岸红树林湿地多毛类的物种多样性、生态分布特点以及与环境因子的关系。研究区域春、秋两季共记录多毛类动物45种,其中沙蚕科、海稚虫科和小头虫科3个科种类最为丰富,种类属性为低盐或广盐性种类。多毛类平均密度和生物量分别为190个/m2和2.17 g/m2,样地×季节双因素方差分析表明,密度在不同样地间差异显著,密度和生物量的季节变化均为春季显著高于秋季。此外,林外光滩的多毛类数量要高于林内,不同样地的摄食群组成各异。红树林断面的平均种类数和多样性指数H'与沉积物粘土含量呈显著负相关,与多毛类类群的大尺度空间分布特征关联最为紧密的因子为地理纬度。     

Aspects of recent advances in the Ordovician stratigraphy and palaeontology of China

The continuous and richly fossiliferous Ordovician succession of China (particularly South China) comprises a heterogeneous suite of litho- and bio-facies, which has been a main focus of stratigraphical and palaeontological research in recent years. Among the seven GSSPs established in China, three are within the Ordovician System, and the GSSP of the Darriwilian Stage at Huangnitang, Changshan County, western Zhejiang Province, was the first “golden spike” in China and the first for the Ordovician System. A series of case studies have revealed that: (1) the Ordovician radiation of some fossil groups on the Upper Yangtze Platform (e.g., brachiopods and graptolites) reached their first α-diversity acme in the Didymograptellus eobifidus Biozone, four zones earlier than the global trend; (2) the β-diversity peak was attained 3–4 zones later than the α-diversity peak; (3) many brachiopod communities or faunas first occurred in the central part of the Upper Yangtze Platform and subsequently expanded to both more offshore and near-shore facies; (4) diachroneity existed in many aspects of the radiation. The end-Ordovician mass extinction was a severe event in South China. Two pulses of the extinction are recognized for a number of major fossil groups, some being most strongly affected during the first pulse whereas the others suffered during the second pulse. Macroevolution during the Ordovician–Silurian transition has been investigated in detail, and the role of the Lazarus effect has been found to be less important than previously believed.     

Marine ostracod provinciality in the late ordovician of palaeocontinental laurentia and its environmental and geographical expression

Background

We examine the environmental, climatic and geographical controls on tropical ostracod distribution in the marine Ordovician of North America.

Methodology/Principal Findings

Analysis of the inter-regional distribution patterns of Ordovician Laurentian ostracods, focussing particularly on the diverse Late Ordovician Sandbian (ca 461 to 456 Ma) faunas, demonstrates strong endemicity at the species-level. Local endemism is very pronounced, ranging from 25% (e.g. Foxe basin) to 75% (e.g. Michigan basin) in each basin, a pattern that is also reflected in other benthic faunas such as brachiopods. Multivariate (ordination) analyses of the ostracod faunas allow demarcation of a Midcontinent Province and a southern Marginal Province in Laurentia. While these are most clearly differentiated at the stratigraphical level of the bicornis graptolite biozone, analyses of the entire dataset suggest that these provinces remain distinct throughout the Sandbian interval. Differences in species composition between the provinces appear to have been controlled by changes in physical parameters (e.g. temperature and salinity) related to water depth and latitude and a possible regional geographic barrier, and these differences persist into the Katian and possibly the Hirnantian. Local environmental parameters, perhaps operating at the microhabitat scale, may have been significant in driving local speciation events from ancestor species in each region.

Conclusions/Significance

Our work establishes a refined methodology for assessing marine benthic arthropod micro-benthos provinciality for the Early Palaeozoic.     

Densities of polychaetes in habitat fragments depend on the surrounding matrix but not the complexity of the remaining fragment

Biogenically engineered habitats are constantly changing in space and time, resulting in changes to the landscape and the ecology of associated taxa. Using patchily distributed biogenic habitats on intertidal rocky shores the influences of different aspects of the landscape were investigated, that is, the surrounding matrix and the habitat itself. Experiments to test the effect of the matrix were carried out by transplanting assemblages in artificial habitats from one type of biogenically engineered matrix (created by the tubeworm Galeolaria caespitosa) to another (created by the oyster Saccostrea glomerata). Change to the surrounding matrix resulted in the densities of polychaete worms in central fragments of habitat changing to become more similar to their new surroundings. To test how the habitat influenced predator–prey interactions, polychaetes were transplanted from a complex habitat (created by the turfing alga Corallina officinalis) to less complex habitat (created by the tubeworm G. caespitosa), with or without the presence of the predatory polychaete, Perinereis amblyodonta. This experiment also tested the influence of habitat replacement on densities of polychaetes, regardless of any new interactions between species. Despite coralline turf being a much more complex habitat than Galeolaria, P. amblyodonta successfully preyed on other species of polychaetes in both habitats. Furthermore, in the absence of P. amblyodonta, survival of polychaetes did not differ between the habitats. These types of experiments will be useful for testing similar hypotheses in different landscapes and will assist in gaining a more general understanding of habitat modification.     

Three-dimensional reconstruction of mushroom body neuropils in the polychaete species <Emphasis Type="Italic">Nereis diversicolor</Emphasis> and <Emphasis Type="Italic">Harmothoe areolata</Emphasis> (Phyllodocida,Annelida)

Mushroom bodies are prominent brain neuropils present in most arthropod representatives. Similar structures in the brain of certain polychaete species are possibly homologous to these structures. Using three-dimensional reconstruction techniques, we investigated the structural composition of the mushroom body neuropils in the polychaete species Nereis diversicolor and Harmothoe areolata. Comparative analysis revealed a common organization of neuropil substructures in both species that closely matches the basic assembly of arthropod mushroom bodies. Concurring with earlier homology assessments, these neuroarchitectural similarities provide support for a common origin of mushroom body neuropils in polychaetes and arthropods. Beyond that, differences in the morphological differentiation of neuropil substructures indicate polychaete mushroom bodies to show a high degree of morphological variability, thus impeding the quest for a common ground pattern of these brain centers.     

Environmental factors structuring polychaete communities in shallow rocky habitats: role of physical stress versus habitat complexity

Polychaetes inhabiting 12 different hard bottom habitats were studied. A total of 157 species belonging to 32 families were identified. Differences among habitats in polychaete density, species richness, and diversity were analysed, as well as the relationships between these ecological indices and depth range, slope and in-bay/out-bay gradient. A high faunal homogeneity was found: all biotopes were dominated by a low number of eurytopic species. Intertidal habitats and subtidal ones with scarce algal cover were typified by vagile polychaetes (syllids, nereids), while sessile polychaetes (serpulids, sabellids) appeared typically among subtidal large macrophytes, habitats with a calcareous substrate and shaded habitats. Multivariate analyses showed that habitat complexity, determined by physical disturbance, is the main structuring factor for polychaete populations. Biotopes with the highest structural complexity displayed a high number of companion species increasing ecological indices and denoting a well-structured habitat. On the other hand, communities such as those in the upper intertidal, mainly controlled by physical environmental variables, showed a poorer polychaete fauna, dominated by ubiquitous species and a few well-adapted specialists.