Stability of eelgrass (<Emphasis Type="Italic">Zostera marina</Emphasis> L.) depth limits: influence of habitat type

Seagrass meadows are generally considered stable although few studies have specified and tested this statement. On the basis of a large monitoring dataset from Danish coastal waters, we aimed to test whether the stability of deep eelgrass populations changes along a eutrophication gradient extending from inner bays over outer bays to open coastal waters. We defined stability in terms of the stability properties of constancy, resilience, and/or persistence. Our data allowed us to investigate the stability property constancy expressed as temporal variability in eelgrass depth limits over a time scale of 10 years. We hypothesised that colonisation depths were large and relatively constant along open coasts because of low and relatively constant levels of nutrients and turbidity. Conversely, colonisation depths were hypothesised to be low and variable in protected bays due to higher and more variable levels of nutrients and turbidity. We found that depth limits increased from inner bays towards open coastal waters, matching declines in nutrient concentration and increases in water clarity and oxygen concentration. Stability expressed as constancy of depth limits did not differ significantly between habitat types, and neither did stability of physicochemical variables. However, when data from all habitat types were analysed together, they showed that eelgrass populations at the depth limit were significantly more constant and thus, in this respect, more stable when occurring in deep waters as compared to shallow waters. Areas of good water quality may thus obtain the double benefit of deeper-growing and more stable eelgrass populations. The most likely reason why this pattern did not appear at habitat-type level is that the habitat types studied represented wide spatial variation in water quality and depth limit. We conclude that the question of whether eelgrass populations are stable depends on the stability property and the ecological situation in question. Populations may be considered unstable in terms of inter-annual variation but stable in terms of long-term persistence. Therefore, the common statement that eelgrass populations are stable is not universally true.Communicated by M. Kühl, Helsingør     

Is benthic trophic structure affected by food quality? The Dogger Bank example

The aim of this study was to evaluate the relationship between food quality and trophic structure of macrobenthic communities of the Dogger Bank. Therefore, 21 stations along different depth gradients (a transect across the central Dogger Bank and at a station cross in the north-eastern part) were sampled and analysed for macrofauna trophic structure and food supply, indicated by phytopigment composition and organic matter contents of the sediment. The study revealed differences in the trophic structure of the benthic community in areas with contrasting food supply. The trophic structure reflected differences in relative quality of organic matter. The sandy sediments on top of the bank receiving intermediate quantities of high food quality were inhabited by high abundances of interface feeders and sand-licking amphipods of the genus Bathyporeia. The southern stations characterised by a comparatively high amount of intermediate food quality were dominated by interface feeders such as Amphiura filiformis. The northern and north-eastern stations showing the highest diversity of feeding types were characterised by a comparatively low, but highly variable input of intermediate-quality organic matter consisting of both fresh and degraded organic matter. Results are discussed with a special emphasis on specific Dogger Bank features such as benthic primary production and frontal system characteristics.Communicated by O. Kinne, Oldendorf/Luhe     

Crinoid phylogeny: a preliminary analysis (Echinodermata: Crinoidea)

We describe the first molecular and morphological analysis of extant crinoid high-level inter-relationships. Nuclear and mitochondrial gene sequences and a cladistically coded matrix of 30 morphological characters are presented, and analysed by phylogenetic methods. The molecular data were compiled from concatenated nuclear-encoded 18S rDNA, internal transcribed spacer 1, 5.8S rDNA, and internal transcribed spacer 2, together with part of mitochondrial 16S rDNA, and comprised 3,593 sites, of which 313 were parsimony-informative. The molecular and morphological analyses include data from the bourgueticrinid Bathycrinus; the antedonid comatulids Dorometra and Florometra; the cyrtocrinids Cyathidium, Gymnocrinus, and Holopus; the isocrinids Endoxocrinus, and two species of Metacrinus; as well as from Guillecrinus and Caledonicrinus, whose ordinal relationships are uncertain, together with morphological data from Proisocrinus. Because the molecular data include indel-rich regions, special attention was given to alignment procedure, and it was found that relatively low, gene-specific, gap penalties gave alignments from which congruent phylogenetic information was obtained from both well-aligned, indel-poor and potentially misaligned, indel-rich regions. The different sequence data partitions also gave essentially congruent results. The overall direction of evolution in the gene trees remains uncertain: an asteroid outgroup places the root on the branch adjacent to the slowly evolving isocrinids (consistent with palaeontological order of first appearances), but maximum likelihood analysis with a molecular clock places it elsewhere. Despite lineage-specific rate differences, the clock model was not excluded by a likelihood ratio test. Morphological analyses were unrooted. All analyses identified three clades, two of them generally well-supported. One well-supported clade (BCG) unites Bathycrinus and Guillecrinus with the representative (chimaeric) comatulid in a derived position, suggesting that comatulids originated from a sessile, stalked ancestor. In this connection it is noted that because the comatulid centrodorsal ossicle originates ontogenetically from the column, it is not strictly correct to describe comatulids as unstalked crinoids. A second, uniformly well-supported clade contains members of the Isocrinida, while the third clade contains Gymnocrinus, a well-established member of the Cyrtocrinida, together with the problematic taxon Caledonicrinus, currently classified as a bourgueticrinid. Another cyrtocrinid, Holopus, joins this clade with only weak molecular, but strong morphological support. In one morphological analysis Proisocrinus is weakly attached to the isocrinid clade. Only an unusual, divergent 18S rDNA sequence was obtained from the morphologically strange cyrtocrinid Cyathidium. Although not analysed in detail, features of this sequence suggested that it may be a PCR artefact, so that the apparently basal position of this taxon requires confirmation. If not an artefact, Cyathidium either diverged from the crinoid stem much earlier than has been recognised hitherto (i.e., it may be a Palaeozoic relic), or it has an atypically high rate of molecular evolution.Electronic Supplementary Material Supplementary material is available in the online version of this article at Communicated by J.P. Thorpe, Port Erin     

The effect of feeding behavior of the gastropods<Emphasis Type="Italic"> Batillaria zonalis</Emphasis> and<Emphasis Type="Italic"> Cerithideopsilla cingulata</Emphasis> on their ambient environment

Feeding behaviors of the gastropods Batillaria zonalis, a suspension and deposit feeder, and Cerithideopsilla cingulata, an obligate deposit feeder, were studied to examine their effect on dynamics of suspended materials, total nitrogen (TN) and total organic carbon (TOC) in sediments. Suspension feeding in B. zonalis was observed in detail visually, as it had been previously unreported. An experimental system where B. zonalis and C. cingulata were cultured for 10 weeks, using previously frozen microalgae Nannochloropsis oculata as food, was then constructed. During feeding observations, the suspension-feeding B. zonalis formed a mucus food cord to entangle particulate materials, which were subsequently ingested. The feeding mode of B. zonalis is hence categorized as ctenidial filter feeding. For the culture experiments, decreases in suspended materials were seen only in the B. zonalis cultures, while the control (no gastropods) and C. cingulata cultures remained nearly unchanged. Sediment TN and TOC showed no significant differences between B. zonalis (with mean TN at 0.0345% and mean TOC at 0.261%) and control cultures (with TN at 0.0389% and TOC at 0.331%), but the sediments in C. cingulata cultures had lower levels (with TN at 0.0204% and TOC at 0.156%). The C/N ratios were similar for both B. zonalis (7.55) and C. cingulata (7.68) cultures, and both were lower than the control cultures (8.55). The filtration rate for B. zonalis was lower than that previously observed in bivalves inhabiting the same intertidal flat (e.g. Cyclina sinensis, Grafrarium tumidum and Barbatia virescens). However, Batillaria zonalis occurs at higher abundances than these bivalves. Therefore, it is expected that this species has a large affect upon the transport of suspended materials to the sediments. The addition of TN and TOC to sediments in B. zonalis cultures was probably caused by biodeposition, but deposit feeding by B. zonalis may have restrained the accumulation of those components. The impact of deposit feeding in Cerithideopsilla cingulata cultures was most probably stronger than sedimentation and biodeposition, because of the lower sediment TN and TOC. Bioturbation by both B. zonalis and C. cingulata yields the same effect on sediment quality, as indicated by the low C/N in the culture sediment of both treatments, despite difference in feeding modes. This paper demonstrates, for the first time, the importance of gastropods in bioturbation and removal of suspended materials in subtropical tidal flat habitats.Communicated by T. Ikeda, Hakodate     

What attracts juvenile coral reef fish to mangroves: habitat complexity or shade?

The value of mangroves for fish species is usually explained in terms of high food abundance or shelter against predators as a result of high turbidity and structural complexity. In a field experiment, artificial mangrove units (AMUs) were designed as open cages, each of which was provided with a different degree of structural complexity and shade. Fish species that were attracted to the AMUs were identified and counted and the effects of shade and structural complexity, as well as the interaction between the two factors, were tested. Diurnal fish showed a preference for the greatest structured complexity and for a moderate increase in shade. Two nocturnal species common in local mangroves as well as seagrass beds showed statistically significant effects: densities of Haemulon sciurus were positively related to both shade and structural complexity, whereas only shade had a significant positive effect on densities of Ocyurus chrysurus. The experiment indicated that the attractiveness of mangrove vegetation for H. sciurus may be influenced by the structural complexity of the habitat as well as by the degree of shade, and that both factors are equally and separately important. Individuals of O. chrysurus that are attracted to mangroves are more likely to be influenced by the presence of shade than by the degree of structural complexity. The data thus indicated that the positive relationship between fish densities in mangrove habitats and the degree of shade and structural complexity, or both, is species-specific.Communicated by O. Kinne, Oldendorf/Luhe     

Strategic investment in reputation

Although collective efforts are common in both animal and human societies, many human and probably animal social dilemmas have no obvious cooperative solution, which is a challenge for evolutionary biologists. In public goods games, i.e. the experimental paradigm for studying the sustainability of a public resource with human subjects, initial cooperation usually declines quickly. Recently, it has been shown that the interaction with another social game in which good reputation attracts help, can maintain a high level of cooperation in the public goods game. Here we show experimentally that humans use different strategies in the public goods game conditional on whether the player knows that his decisions will be either known or unknown in another social game. The knowledge of being recognized as the same individual in both scenarios motivates players to invest in their reputation and thus sustain the public resource. However, cooperation declines immediately when individual identities switch from being recognizable to being unrecognizable between the two interacting games.Communicated by M. Borgerhoff-Mulder     

Responses to threat by female bobucks,<Emphasis Type="Italic"> Trichosurus caninus</Emphasis>, during different stages of offspring development

The costs of reproduction are widely recognised as a major selective force in the evolution of various behavioural and life-history characteristics. In particular, the behaviour of reproductively active animals is likely to change when breeding increases risk of predation. We investigated the effect of an experimentally derived threat on the vigilance and escape behaviour of female Trichosurus caninus with and without dependent offspring and at different stages of offspring development. Females with offspring showed a heightened response to the threat in comparison to females without offspring. In addition, females with offspring displayed a stronger response at earlier stages of their offsprings development.Communicated by P. Bednekoff     

Distribution normality of pH and H<Superscript>+</Superscript> activity in soil

Soil pH, a log-transformed form of hydrogen ion (H⁺) activity, is often used for various statistics. However whether soil pH meets the requirement of distribution normality for statistical analyses is unknown. We studied the statistical suitability of both soil pH and H⁺ activity by examining their distribution normalities. Results show that the variability of H⁺ activity is higher and its distribution is skewer, which is significantly different from the normal and soil pH, regardless of soil conditions and analytical methods. Here we demonstrate that soil pH is more appropriate than the H⁺ activity for performing statistical analysis in terms of distribution normality.     

The effects of light and thallus scour from<Emphasis Type="Italic"> Ecklonia radiata</Emphasis> canopy on an associated foliose algal assemblage: the importance of photoacclimation

Kelp canopies have long been recognised for their influence on the structure of algal assemblages on sublittoral reefs. In Marmion Lagoon, Western Australia, we investigated how a canopy of the small kelp Ecklonia radiata affected the associated foliose algal assemblage and what potential mechanisms were responsible for the effects we observed. Light levels and physical abrasion (thallus scour) by an E. radiata canopy were manipulated in an orthogonal fixed-factor experiment that revealed that changes in light, but not thallus scour, had a significant effect on the composition of the algal assemblage. Reduced light levels were associated with a decrease in the Shannon-Wiener diversity index and an increased dominance of the foliose algae Pterocladia lucida and Rhodymenia sonderi. Photobiological investigations of three foliose species, P. lucida, R. sonderi and Chauviniella coriifolia indicated that they were able to photoacclimate to low light levels by increasing thallus absorption by up to 11%. Photoacclimation was also evident by an increase in the maximum rate of electron transport under low-light conditions. We conclude that the E. radiata canopy in Marmion Lagoon structures the foliose algal assemblage through the modification of the light environment and that this effect may be mediated by differences in the ability of different species of foliose algae to photoacclimate.Communicated by M.S. Johnson, Crawley