Phoronid phylogenetics (Brachiopoda; Phoronata): evidence from morphological cladistics,small and large subunit rDNA sequences,and mitochondrial cox1

A matrix of 24 morphodevelopmental characters and an alignment of small subunit (SSU) and large subunit (LSU) rDNA nuclear and cox1 mitochondrial gene sequences (～4500 sites) were compiled from up to 12 phoronids including most named taxa, but probably constituting only a portion of worldwide diversity. Morphological data were analysed by weighted parsimony; sequence data by maximum and Bayesian likelihood, both with Phoronis ovalis as the local outgroup. Morphological and sequence‐based phylogenies were similar, but not fully congruent. Phoronid rDNAs were almost free from mutational saturation, but cox1 showed strong saturation unless distant outgroups and P. ovalis were omitted, suggesting that many phoronid divergences are old (≥100 Myr). rDNA divergence between named phoronid taxa is generally substantial, but Phoronopsis harmeri (from Vladivostock) and Phoronopsis viridis (from California) are genetically close enough to be conspecific. In another alignment, of 24 taxa, phoronid rDNAs were combined with data from brachiopods and distant (molluscan) outgroups. The relative ages of divergence between phoronids and their brachiopod sister‐groups, of the split between the P. ovalis and non‐ovalis lineages, and of other phoronid splits, were estimated from this alignment with a Bayesian lognormal uncorrelated molecular clock model. Although confidence limits (95% highest probability density) are wide, the results are compatible with an Early Cambrian split between phoronids and brachiopods and with the Upper Devonian latest age suggested for the P. ovalis/non‐ovalis split by the putative phoronid ichnofossil, Talpina. Most other ingroup splits appear to be ～50–200 Myr old. Inclusion of phoronids with brachiopods in the crown clade pan‐Brachiopoda suggests that a distinctive metamorphosis and absence of mineralization are ancestral phoronid apomorphies. Worldwide diversity and possible associations between character‐states and life‐history attributes deserve comprehensive further study.     

On the organization of the lophophore in phoronids (Lophophorata: Phoronida)

The organization of the lophophore is the main feature used for the identification of phoronid species. The structure of the lophophore and tentacles in seven phoronid species (Phoronis ovalis, P. ijimai, P. hippocrepia, P. svetlanae, P. australis, Phoronopsis harmeri, and Ph. malakhovi) collected in different areas of the World Ocean was studied. Two new patterns of the phoronid lophophore structure were found: “transition to horseshoe-shaped” (as in P. ovalis from Aniva Bay and in P. ijimai from the coast of Iturup Island, Sea of Okhotsk) and “transition to spiral” (in burrowing specimens P. hippocrepia from Aniva Bay, P. svetlanae and Ph. harmeri from Vostok Bay, Sea of Japan). For the first time it was shown that phoronid species with different types of the lophophore structure possess different kinds of tentacles. Thus, five types of phoronid tentacles were identified that vary in the shape of their cross section: rounded, oval, ellipsoid, rectangular, and skittle-shaped. A correlation was found between lophophore organization and the type of tentacles in phoronids. A table of the correlation between body size, lophophore organization, tentacle structure, and mode of life in different phoronid species is proposed.     

Intertidal distribution of infauna in a central California lagoon: the role of seasonal blooms of macroalgae

This study examined the roles of seasonal blooms of green algae, Ulva expansa (Setchell) Setchell et Gardner, and biotic disturbance by burrowing ghost shrimp, Callianassa gigas Dana, and foraging rays, on the intertidal distributions of a phoronid, Phoronopsis viridis Hilton, and a tellinid bivalve, Macoma nasuta (Conrad). Algal removal experiments in 1984 and 1986 demonstrated that heavy seasonal algal cover in the lower zone significantly reduced the abundances of both Phoronopsis and Macoma. Growth of Macoma transplanted into the algal zone was significantly lower in plots with algal cover than in plots regularly cleared of algae. Algal cover did not significantly affect early recruitment of either Phoronopsis or Macoma. Neither ghost shrimp nor rays appeared to reduce the abundances of phoronids or clams, although ray disturbance did result in a significant increase in the proportion of phoronids regenerating dorsal body parts. These results indicate that seasonal algal blooms are capable of producing discrete patterns of infaunal distribution in intertidal sedimentary habitats.     

Effects of a macroalgal mat (Ulva lactuca) on estuarine sand flat copepods: an experimental study

The effects of a macroalgal mat (Ulva lactuca) on intertidal sand flat copepods were studied in Jamaica Bay, NY. In 1999, Ulva was removed from paired quadrats in the algal mat. Density and species richness were estimated from sediment cores over seven biweekly periods and compared with Ulva-removal quadrats and with an unvegetated reference site. In a separate analysis at the same site, species composition and density of all copepods within the algal mat were determined from replicated quadrat samples over seven dates. Emergence traps were deployed in all treatments in 1999 to quantify copepod movements from sediments into the water column during flooding tide. Dissolved oxygen (DO) was continuously monitored over several 96-h periods in and above the algal mat. In 2000, three biomass levels of Ulva (0, 100, 200 g DW per m²) were added to paired quadrats in the unvegetated zone. Density and species richness were estimated over eight time periods. Ulva removal resulted in significant increases in density and species richness, but levels remained lower than at the unvegetated site. Likewise, addition of Ulva biomass produced significant declines in copepod density. However, the differences between low- and high-Ulva-addition treatments were insignificant. Increases in phytal copepods after Ulva addition did not compensate for losses of epibenthic and infaunal species. The overall effect of the Ulva mat was a net loss of copepod density, including the loss of nearly all infaunal species. Anoxia in sediments and within the algal mat are correlated with declines in infaunal species and with escape by some infaunal species from the sediment into the water column.     

Spatial associations among palatable and unpalatable macroalgae: A test of associational resistance with a herbivorous amphipod

Associational resistance is the process by which plants may gain protection from spatial associations with neighbouring plants. We tested whether association with an unpalatable alga, Dictyopteris acrostichoides, affects the abundance and colonisation behaviour of the herbivorous amphipod Peramphithoe parmerong on its preferred host alga Sargassum linearifolium. Despite predictions, natural densities on S. linearifolium when surrounded by D. acrostichoides were higher than on isolated individuals of S. linearifolium. Colonisation experiments in the laboratory and the field tested the hypotheses that the observed variation in field abundance with algal neighbourhood was due to variation in the size of habitat patches, physical obstruction of host finding by D. acrostichoides and variation in the relative abundance of S. linearifolium and D. acrostichoides. None of these possible mechanisms was found to significantly alter rates of amphipod colonisation on the scales of individuals selecting among algal pieces in the laboratory or among habitat patches in the field. The failure of colonisation processes to explain observed variation in natural amphipod densities suggests that post-colonisation processes such as survival or emigration may vary with the spatial associations among algae.     

Ultrastructural study of spermatogenesis in<Emphasis Type="Italic"> Phoronopsis harmeri</Emphasis> (Lophophorata,Phoronida)

The process of sperm development in Phoronopsis harmeri was studied by electron microscopy. Developing spermatogenical cells are aggregated around the capillaries of the haemal plexus. The spermatogonia, which are situated around the capillary walls of the caeca, are remarkable for the presence of germ-line vesicles and contain their centrioles near the cell membrane. The spermatocytes and spermatids are flagellated cells arranged in clusters. During spermiogenesis the basal body/flagellum complex migrates to the apical pole of the spermatid. The acrosome-like structure arises from material produced by the Golgi complex. It lacks a surrounding membrane and has a fibrillar content. The nucleus elongates and the condensation of chromatin is caused by an activation of 'initiation centres'. The late spermatid and the spermatozoon appear as two-armed 'V'-shaped cells in which one arm contains the nucleus and posteriorly located mitochondria, and the other one is the axoneme. Spermatogenesis of P. harmeri is an interesting example of gamete differentiation where advanced sperm structure is combined with a plesiomorphic pattern of sperm development characterized as 'flagellate spermatogenesis'. Communicated by H.-D. Franke     

Inter‐ and intraspecific plasticity in distribution patterns of immunoreactive compounds in actinotroch larvae of Phoronida (Lophotrochozoa)

Recent molecular data place Phoronida within the protostome superclade Lophotrochozoa, where they have been suggested to form a monophyletic assemblage with Brachiopoda and/or Nemertea. Herein, the anatomy of the nervous system and the structure of the apical organ are described for two larval stages of Phoronis muelleri in order to contribute to the discussion concerning the evolution of lophotrochozoan nervous systems. Specimens were investigated by confocal laser scanning microscopy using antibodies against the serotonin‐like immunoreactive (serotonin‐lir), the FMRF‐amide‐like immunoreactive (FMRFamide‐lir) and the small cardioactive peptide B‐like immunoreactive (small cardioactive peptide B‐lir) compounds of the nervous system. Consistent with larvae of other phoronid species, we found a complex apical organ that consists of numerous serotonin‐lir flask‐shaped cells, additional bi‐ or multipolar serotonin‐lir cells and several FMRFamide‐lir perikarya. A detailed comparison between our results and those of a previous study on the same species shows significant differences in the innervation of the preoral lobe, the tentacles and the telotroch. Our work is the first to prove the presence of small cardioactive peptide B in phoronids. In larvae of P. muelleri, it is expressed in neurites along the margin of the preoral hood, in the mesosome, in the tentacles, in the trunk as well as in the apical organ. A positive signal for this peptide is also known from molluscs, annelids and arthropods, indicating that it was also part of the protostomian groundplan. In contrast to a recent study on another phoronid species, Phoronopsis harmeri, we did not find a ventral neurite bundle in the larval stages investigated herein, thus leaving the question open whether this structure was part of the phoronid groundplan or evolved de novo in P. harmeri.     

Macroalgal-mediated transfers of water column nitrogen to intertidal sediments and salt marsh plants

In many temperate estuaries, mats of opportunistic macroalgae accumulate on intertidal flats and in lower elevations of salt marshes, perhaps playing a role in linking water column nitrogen (N) supply to these benthic habitats. Using a flow-through seawater system and tidal simulator, we varied densities (equivalent to 0, 1, 2, or 3 kg m⁻² wet mass) of ¹⁵N-labelled macroalgae (Enteromorpha intestinalis) on estuarine sediments in microcosms with/without pickleweed (Salicornia virginica) to assess N transfers from algae. In the 6-week experiment, macroalgal biomass increased from initial levels in the lower density treatments but all algae lost N mass, probably through both leakage and decomposition. With all densities of algae added, sediments and pickleweed became enriched in ¹⁵N. With increasing mat density, losses of algal N mass increased, resulting in stepwise increases in ¹⁵N labeling of the deeper sediments and pickleweed. While we did not detect a growth response in pickleweed with macroalgal addition during the experiment, N losses from algal mats that persist over many months and/or recur each year could be important to the mineral nutrition of N-limited marsh plants. We conclude that N dynamics of intertidal sediments and lower salt marsh vegetation are linked to the N pools of co-occurring macroalgae and that further study is needed to assess the magnitude and importance of N transfers.     

Habitat size, flora, and fauna: Interactions in a tidal saltwater marsh

Anthropogenic habitat fragmentation is increasingly problematic in both terrestrial and aquatic systems. Fragmentation reduces the size of habitat patches, so examining the effect of patch size on community structure can provide insight into the potential effects of fragmentation. In this study, we examined the effect of habitat size on the density of Spartina alterniflora shoots in tidal saltwater marshes, as well as on the two predominant macrofaunal species, the marsh periwinkle Littoraria irrorata and fiddler crabs Uca spp. We estimated the density of shoots in three different marsh habitats, (1) large island marshes, (2) small island marshes, and (3) large fringing marshes, in Indian Field Creek, York River, Chesapeake Bay. We manipulated shoot density in each of the marsh types to distinguish between the effects of marsh grass density and marsh type on crab and Littoraria densities in the system. We found significant differences in grass density among the three marsh types as well as significant species-specific effects of grass density, marsh type, and distance from edge on faunal abundance. Decreasing the shoot density resulted in a decrease in Littoraria density in the large marshes. Littoraria density increased with distance from edge in the small marshes and in the first 5 m of the fringing marshes, then decreased with distance from edge after 5 m in the fringing marshes. Shoot density had a negative effect on crabs in both the large and small marshes. These results suggest that fragmentation would have a negative effect on the community structure by lowering the densities of both the flora and fauna.     

Community and population-level responses to disturbance in a sandflat community

The responses of soft sediment infauna were investigated in an intertidal sandflat to determine patterns of recolonization and succession at the community and population level. Experimental disturbance plots, 1 m², were initiated in August and sampled for 4.5 months along with ambient sediments. Sediment grain-size was used as a general indicator of the physical state of the disturbance patches, and grain-size distributions among disturbance and ambient patches became similar after ∼2.5 months. Recolonization varied among the dominant infaunal taxa. Densities of infauna that were most abundant in the habitat, primarily syllid polychaetes, did not recover to ambient levels until 3-4 months after disturbance, when ambient densities were falling to winter lows. Multivariate analysis indicated that community recovery occurred by the end of the study period after 4.5 months. Although community structure recovered by the end of the study, the population structure of the dominant species Parapionosyllis longicirrata remained significantly different among ambient and disturbed patches. On all sampling dates except one, disturbance patches had a higher number of larger individuals than ambient sediments. Previous studies have shown that bedload transport of juveniles and adults, and other processes, can cause recolonization to be relatively rapid on intertidal sandflats. However, our results indicate that recovery times may be on the order of months at large disturbance sizes. Therefore, rapid responses may occur primarily in the case of small-scale (<1 m²) disturbance patches. Secondly, recovery at the community level does not necessarily mean that population-level characteristics of species comprising the community have recovered. Population-level differences may be longer lasting than indicated by community level indicators of recovery.     

The fauna of the South China Sea include unknown phoronid species: new records of larvae and adults

The Phoronida is a phylum of marine invertebrates that have a worldwide distribution and that form huge benthic aggregations in many aquatic areas. Although there are only 11 recognized species of phoronids, many species clearly remain to be described. The matching of larval and adult stages of the same species will help indicate which larvae belong to described species and which represent undescribed species whose adults have yet to be found. Larvae of four phoronid species were collected in Nha Trang Bay and investigated by light microscopy and molecular methods. Three of the described larvae belong to new phoronid species that have yet to be named. These larvae have unique morphological features and occupy separate positions on the phylogenetic tree of phoronids. One of the described larvae belongs to Phoronis hippocrepia or to a closely related species. Observations made with a new macrophotographic method indicated that Phoronopsis californica also occurs in Nha Trang Bay. Thus, according to records of larvae and adults, Nha Trang Bay contains at least nine phoronid species: Phoronopsis californica, Phoronopsis malakhovi, Phoronis australis, Phoronis hippocrepia (or a closely related species), three unknown phoronid species whose larvae have unique morphological features and that apparently belong to the genus Phoronis, and two species, whose adults have unique set of morphological features. The richness of phoronids in this area suggests that the Indo-Pacific is a centre of phoronid biodiversity.     

Demography, competitive interactions and grazing effects of intertidal limpets in southern New Zealand

Population dynamics and the effects of intraspecific competition on limpet growth and maintenance of bare patches were investigated for the intertidal limpet Cellana ornata (Dillwyn) at a boulder-dominated site and on a rocky platform near Kaikoura (South Island), New Zealand. Distribution and abundance patterns of C. ornata were described in relation to other biota and tidal height. C. ornata occurred almost exclusively in patches devoid of macroalgae, particularly in the mid-tidal zone. Both adult and juvenile limpets were most abundant on the tops of boulders, where their numbers were positively correlated with barnacle cover, which averaged 77%. The size structure and growth patterns of C. ornata were different between populations. Mark-recapture studies showed that the slopes of annual growth increments regressed on initial sizes were similar at both sites but that the annual increments on the platform were about 6 mm greater than on boulders. Growth virtually ceased at 27 mm for limpets at the barnacle-dominated boulder site and at 40 mm at the platform site. Recruiting cohorts had 20% survival on boulders and 37% on the platform during their first year. The largest size classes at both sites had around 57% annual survival. To test the effects of varying limpet densities on the growth and mortality of limpets and the maintenance of bare patches, densities of C. ornata were experimentally increased at both sites. Beyond a density of 4 per 0.25 m², sizes and survival of limpets were reduced at both sites, but the effect was more pronounced at the boulder site. Limpets at the boulder site were more effective at maintaining bare space than those on the reef platform. Enclosing limpets in plots with and without barnacles showed that C. ornata and a co-occurring species (Cellana denticulata (Martyn)) grazed more effectively and had greater growth in cleared plots. Overall, there was considerable variation in the demographics of C. ornata between populations driven by site-specific characteristics.     

富营养化山仔水库沉积物微囊藻复苏的受控因子

山仔水库作为福建省福州市重要的饮用水水源地之一,从2000年起每年都周期性爆发蓝藻门微囊藻属(Microcystis)水华现象,特别是在温暖的季节。对于这个富营养化水库,是否在沉积物中存在蓝藻门微囊藻的"种源"？假设山仔水库底泥中存在蓝藻门微囊藻休眠体,一定的环境条件能够促进蓝藻门微囊藻的复苏。研究于2009年12月采集水库大坝断面5根柱状沉积物,采用正交试验的方法,模拟了温度、光照、pH值、营养盐、物理扰动和浮游动物(膨大肾形虫)等环境因子对山仔水库沉积物中蓝藻门微囊藻的复苏响应。结果表明,底泥中存在着一定数量的底栖动物和硅藻、蓝藻和绿藻等微藻,从实验结束后沉积物中微囊藻数量的减少和上覆水体中微囊藻数量的增加,可以判断在适宜的环境条件下,蓝藻门微囊藻能够复苏并上浮到上覆水体中。正交实验显著性分析表明,温度是沉积物蓝藻门微囊藻复苏的重要影响因子,光照次之,上覆水体的pH值、营养盐、物理扰动和浮游动物干扰对沉积物蓝藻门微囊藻的复苏影响作用不显著,升温有利于沉积物中微囊藻的复苏。     

Effects of seagrass habitat fragmentation on juvenile blue crab survival and abundance

Seagrasses form temporally dynamic, fragmented subtidal landscapes in which both large- and small-scale habitat structure may influence faunal survival and abundance. We compared the relative influences of seagrass (Zostera marina L.) habitat fragmentation (patch size and isolation) and structural complexity (shoot density) on juvenile blue crab (Callinectes sapidus Rathbun) survival and density in a Chesapeake Bay seagrass meadow. We tethered crabs to measure relative survival, suction sampled for crabs to measure density, and took seagrass cores to measure shoot density in patches spanning six orders of magnitude (ca. 0.25-30,000 m²) both before (June) and after (September) seasonally predictable decreases in seagrass structural complexity and increases in seagrass fragmentation. We also determined if juvenile blue crab density and seagrass shoot density varied between the edge and the interior of patches. In June, juvenile blue crab survival was not linearly related to seagrass patch size or to shoot density, but was significantly lower in patches separated by large expanses of unvegetated sediment (isolated patches) than in patches separated by <1 m of unvegetated sediment (connected patches). In September, crab survival was inversely correlated with seagrass shoot density. This inverse correlation was likely due to density-dependent predation by juvenile conspecifics (i.e. cannibalism); juvenile blue crab density increased with seagrass shoot density, was inversely correlated with crab survival, and was greater in September than in June. Shoot density effects on predator behavior and on conspecific density also likely caused crab survival to be lower in isolated patches than in connected patches in June. Isolated patches were either large (patch area >3000 m²) or very small (<1 m²). Large isolated patches had the lowest shoot densities, which may have allowed predators to easily find tethered crabs. Very small isolated patches had the highest shoot densities and consequently a high abundance of predators (=juvenile conspecifics). Though shoot density did not differ between the edge and the interior of patches, crabs were more abundant in the interior of patches than at the edge. These results indicate that seagrass fragmentation does not have an overriding influence on juvenile blue crab survival and density, and that crab cannibalism and seasonal changes in landscape structure may influence relationships between crab survival and seagrass habitat structure. Habitat fragmentation, structural complexity, faunal density, and time all must be incorporated into future studies on faunal survival in seagrass landscapes.     

Would adding scallop shells (Chlamys islandica) to the sea bottom enhance recruitment of commercial species?

We examined the impact of adding scallop shells (byproduct of the fisheries) to sandy and rocky sea bottoms in the northern Gulf of St. Lawrence. The effect of adding shells was greatest on sandy bottoms where species richness increased 3.7-fold and species diversity 1.9-fold. The increase in most species was due to immigration rather than new settlement. Trials examining the effect of different densities of shells in plots of the same size (4 m²) showed that species diversity increased rapidly with shell abundance and levelled off when shells covered half of the bottom, whereas species richness only levelled off when shells almost completely covered the bottom. Trials examining the effect of the size of the shell patches (shell density being kept constant) showed that species diversity was already maximal in 1-m² plots, whereas species richness only attained a plateau at 4 m². Our small-scale trials indicate that the addition of shells would have a positive impact, increasing numerous invertebrates, including commercial species (scallops, whelks and urchins).     

Fate of resting cysts of Alexandrium spp. ingested by Perinereis nuntia (Polychaeta) and Theola fragilis (Mollusca)

The ingestion of resting cysts of Alexandrium spp. by Perinereis nuntia (Polychaeta) and Theola fragilis (Mollusca) was experimentally examined in the laboratory. P. nuntia and T. fragilis were cultured in bottom sediment containing a high density of Alexandrium cysts under dark conditions. Moreover, to evaluate the degree and consequence of being ingested, the density of cysts in the control sediment (no macrobenthic organisms) and the germination capability of the cysts in the faecal pellets of the two species of macrobenthos were examined.Cysts in the culture sediment were found to be ingested by both P. nuntia and T. fragilis. No difference in the density of cysts between the sediments cultured with and without P. nuntia was observed. However, the density of cysts in the sediments with T. fragilis decreased by 24% compared to the density in the control sediment. It is possible that most of the cysts ingested were digested by T. fragilis. The rate of Alexandrium cyst digestion by this species is estimated 594 cysts/individual/day. It is estimated that 91% of the cysts ingested by T. fragilis were partially or totally digested and only 9% were excreted in a viable state during the experiment. Thus, T. fragilis has a stronger affect on the abundance of Alexandrium cysts compared with P. nuntia.No significant difference was observed between the germination success of the cysts from faecal pellets of P. nuntia and T. fragilis compared to the cysts in the control sediment. If, however, the necessary light for the cysts to germinate is cut off by being enclosed within the faecal pellet, the germination rate of cysts from the faecal pellets may be suppressed.     

Sediment-associated cues for larval settlement of the infaunal spionid polychaetes Polydora cornuta and Streblospio benedicti

Do patchy distribution patterns of infaunal polychaetes result from active site selection of larvae influenced by sediment-associated microbial cues? This hypothesis was tested with still-water laboratory settlement assays revealing the acceptance or rejection of polychaete larvae to qualitatively different sediments. Laboratory brood cultures of the spionid polychaetes Polydora cornuta and Streblospio benedicti yielded a sufficient number of larvae with planktotrophic development for bioassays. High settlement rates (75-95%) of test larvae were observed in response to natural sediment. Sterilization of natural sediment significantly decreased settlement of P. cornuta (25-55%) while combustion of sediment significantly decreased the settlement rate in both species (5-50%). Differences in settlement responses to sediments treated by sterilization or combustion most likely resulted from a variety of factors such as modified sediment fabric, grain size distribution and quantity of adsorbed organic matter. To experimentally address the potential role of microorganisms and microbial metabolites as mediators of larval settlement, ashed sediment was inoculated with viable microorganisms obtained from natural sediment. In both polychaete species, this treatment significantly increased larval settlement in comparison to the control of ashed sediment indicating that larval settlement was at least partially mediated by the presence of microorganisms associated with sediment.     

Post-fire spread of alien plant species in a mixed broad-leaved forest of the Insubric region

How do tree species regenerate and which ecological conditions are required after forest fire in the Insubric region of the Alps? Are indigenous stand-forming tree species resistant over the invasion of alien plant species after such a disturbance? We addressed these questions in a case study in the Swiss canton of Ticino. In April 2006, a surface fire with severe intensity burnt a forest area of 55 ha on a south-facing slope (400-800 m.a.s.l.). The dominant trees in the investigated area were chestnut (Castanea sativa Mill.), beech (Fagus sylvatica L.) and deciduous oaks (Quercus spp.) mixed with tree species of intermediate height. Vegetation data were collected in May and August 2009 by systematic sampling. Resprouting of the survived trees and generative regeneration were analysed by counting vegetative shoots from sprouting stools, of seedling age, height and damage rate, respectively. Different vegetation structures related to low or high fire intensity were clearly visible three years after the forest fire, creating various habitats for both new invaders and seedlings of the stand-forming trees. The dominant chestnut was the only tree species that regenerated effectively by sprouting from stools. Seedlings of the stand-forming trees grew in high abundance under shadow conditions close to their mother trees which provided the seed source. In contrast, pioneer trees invaded patches where full light was available. Under such conditions the two main woody alien plant species, Ailanthus altissima and Robinia pseudoacacia, grew in high abundance. Due to the different ecological requirements of indigenous and alien tree seedlings, not any interaction between the two groups was detected.     

Restoration of Aegiceras corniculatum mangroves in Jiulongjiang Estuary changed macro-benthic faunal community

Macro-benthic faunal communities were compared between non-vegetation mudflat and Aegiceras corniculatum mangroves with different ages in Jiulongjiang Estuary, China. Faunal species number was highest in the mature mangrove and was higher in mangroves than in the mudflat, as snails and some crustaceans species were only collected in mangroves. The 5-year-old mangrove had the highest infaunal abundance and crustacean biomass. Snails had more abundance in the young mangroves. Uca arcuata was the dominant crab species in the non-vegetation mudflat and 5-year-old mangrove. Mangrove vegetation and sediment characteristics analyses indicated different habitats due to A. corniculatum mangrove restoration. However, overall poor correlations between faunal assemblage and sediment properties indicated that sediment properties were not the major factors influencing faunal distribution.     

Biodiversity of benthic invertebrates and organic matter processing in shallow marine sediments: an experimental study

The main objective of this study was to measure the impact of benthic invertebrate diversity on processes occurring at the water-sediment interface. We analyzed the effects of interactions between three shallow water species (Cerastoderma edule, Corophium volutator, and Nereis diversicolor). The impacts of different species richness treatments were measured on sediment reworking, bacterial characteristics, and biogeochemical processes (bromide fluxes, O₂ uptake, nutrient fluxes, and porewater chemistry) in sediment cores. The results showed that the three species exhibited different bioturbation activities in the experimental system: C. edule acted as a biodiffusor, mixing particles in the top 2 cm of the sediments; C. volutator produced and irrigated U-shaped tubes in the top 2 cm of the sediments; and N. diversicolor produced and irrigated burrow galleries in the whole sediment cores. C. edule had minor effects on biogeochemical processes, whereas the other species, through their irrigation of the burrows, increased the solute exchange between the water column and the sediment two-fold. These impacts on sediment structure and solute transport increased the O₂ consumption and the release of nutrients from sediments. As N. diversicolor burrowed deeper in the sediment than C. volutator, it irrigated a greater volume of sediments, with great impact on the sediment cores.Most treatments with a mixture of species indicated that observed values were often lower than predicted values from the addition of the individual effects of each species, demonstrating a negative interaction among species. This type of negative interaction measured between species on ecosystem processes certainly resulted from an overlap of bioturbation activities among the three species which lived and foraged in the same habitat (water-sediment interface). All treatments with N. diversicolor (in isolation and in mixture) produced similar effect on sediment reworking, water fluxes, nutrient releases, porewater chemistry, and bacterial characteristics. Whichever species associated with N. diversicolor, the bioturbation activities of the worm hid the effect of the other species. The results suggest that, in the presence of several species that use and modify the same sediment space, impact of invertebrates on ecosystem processes was essentially due to the most efficient bioturbator of the community (N. diversicolor). In consequence, the functional traits (mode of bioturbation, depth of burrowing, feeding behaviour) of an individual species in a community could be more important than species richness for some ecosystem processes.