An experimental study on recolonization and succession of marine macrobenthos in defaunated sediment

Hypoxia/anoxia in coastal waters is a world wide problem which often results in mass mortality and defaunation of benthos. In this study, field experiments were carried out to examine recolonization and succession of macrobenthic infauna in defaunated sediments, and the time required for recovery from complete defaunation to a stable community. Trays (33 cm length × 25.5 cm width × 11 cm depth) of defaunated sediment were exposed at the subtidal of a pristine site in subtropical Hong Kong. Temporal changes of macrobenthic communities in defaunated sediment were analyzed by univariate and multivariate statistics, and compared with those in undisturbed natural sediment at the same site. Initial colonization of macrobenthos occurred rapidly. A total of 42 species was found, with an average of 258 animals per tray and 24 species per tray recorded in the first month. Abundance showed a small peak (496 animals per tray) after 3 months, reached a sharp peak (1154 animals per tray) after 6 months, and declined thereafter. Species number increased gradually, reached a maximum (68 species per tray) after 9 months, and then decreased. Recolonization was predominantly contributed by larval settlement rather than adult migration. Temporal changes in abundance, species number and diversity of the macrobenthic community in defaunated sediment resemble the spatial changes along a decreasing pollution gradient previously defined by other authors. Results of this experiment suggest that newly available sediment may allow more species to colonize (or coexist) than sediment pre-occupied by an established community. This is probably due to less interspecific competition in the former habitat. No significant difference in abundance or species richness was observed between defaunated and natural sediments after 15 months, suggesting that a stable community had been achieved, although minor variations in species composition were still discernible between defaunated and natural sediments. Received: 24 May 1999 / Accepted: 20 October 1999     

Metabolic differences between “demersal” and “pelagic” development of the Antarctic sea urchin Sterechinus neumayeri

 Early development of the Antarctic sea urchin Sterechinus neumayeri was examined under two differ-ent culture regimes: one to simulate development near-bottom (“demersal development”) and the other to simulate the development of embryos in the water column (“pelagic development”). When embryos of both treatments reached the hatching blastula stage at 5 d post-fertilization (−1.5 °C), the blastulae that had undergone demersal development evidenced significant differences (by ANOVA or suitable non-parametric comparison) in the following: a thicker blastoderm layer (12%, P < 0.001), higher ash-free dry weights (19%, P < 0.01), lower mass-specific respiration rates (50%, P < 0.001), higher incorporation rates of ³⁵S-methionine into protein (23%, P < 0.003), and a differential pattern of protein synthesis. When embryos developed demersally, they remained in the jelly-coat material released with the eggs at spawning. Quantitative isolation of this jelly-coat material in S. neumayeri demonstrated that it contained a significant amount of organic matter, 115 ng ash-free dry mass per egg, equivalent to 17% of the egg's initial organic mass. Uptake of external nutrients during embryogenesis may be a significant component of the physiological energetics of this polar invertebrate by allowing the utilization of jelly-coat material released by a female during spawning. Received: 21 April 1999 / Accepted: 5 June 2000     

Short retention times of stomach temperature loggers in free-living seabirds: is there hope in the spring?

Stomach temperature loggers have proved useful for the study of feeding activity in free-living seabirds, but their usage has been restricted because they are frequently dislodged and lost during the seabirds' regular regurgitation of indigestible prey remains. In the present study we examine the incidence of spontaneous regurgitation (pellet production) in free-living seabirds, consider the effect this has on the likely retrieval of stomach temperature loggers and present a structural modification of the logger housings which leads to a much lower incidence of regurgitation. Systems were tested on albatrosses, cormorants, gannets and penguins. Received: 15 July 1997 / Accepted: 31 October 1997     

Simultaneous hermaphroditism in the marine shrimp Lysmata wurdemanni (Caridea: Hippolytidae): an undescribed sexual system in the decapod Crustacea

Simultaneous hermaphroditism with outcrossing, a previously unconfirmed sexual system in decapod crustaceans, is documented in the shrimp Lysmata wurdemanni (Gibbes), using time-lapse video observations on mating in pairs of “female-phase” (FP) individuals. Copulations between FPs resulted in successful spawning and development of embryos. However, female-phase hermaphrodites maintained in isolation were unable to self-fertilize spawned eggs. All smaller individuals possessed characters typical of caridean males, including male gonopores, appendices masculinae, ejaculatory ducts, and cincinnuli on the first pleopods. However, the gonad of these male-phase (MP) individuals was an ovotestis with an undeveloped ovarian portion. FPs, which spawn eggs and incubate embryos, also had male gonopores and an ovotestis terminating in ejaculatory ducts containing sperm. In FPs, male pleopod characters were absent or reduced, and a female incubatory character, expanded pleopod flanges, showed a greater relative size. Smaller size classes of the population were composed only of MPs while larger size classes consisted of both MPs and FPs. “Transitionals,” individuals with MP characters but an ovotestis containing vitellogenic oocytes, were rare and overlapped completely in size with FP and larger MP individuals. It is suggested that not all MPs transform into FP hermaphrodites. Social interactions may mediate MP change into FP hermaphrodites in L. wurdemanni. The social system of L. wurdemanni and other Lysmata spp. may determine the exact form of the sexual system in these species. Received: 26 October 1997 / Accepted: 21 April 1998     

Gender- and oxygen-related irrigation behaviour of the decapod Nephrops norvegicus

The Norway lobster Nephrops norvegicus (L.) inhabits burrows in muddy clay sediments (e.g. on the Swedish west coast), where an autumnal oxygen deficiency in the bottom water can occur. Our experiments investigated whether the irrigation of the burrows would reflect a behavioural adaptation to hypoxia, and whether any gender differences of such behaviour exist. Irrigation is performed by the pleopods which may compensate for a decreasing oxygen tension. Pleopod activity (total number of strokes per sampling time), associated with oxygen concentration and gender, was studied in N. norvegicus kept in artificial burrows resembling their natural habitat. Male and female lobsters were separately exposed to either normoxia (70% oxygen saturation) or hypoxia (30% oxygen saturation). A sexual difference in behaviour was found, where females irrigated the burrow less than males during normoxia. Females showed a significant increase of pleopod activity in hypoxia compared with normoxic conditions, which was not displayed by the males probably due to the degree of individual variation found. However, when only males were studied during progressive hypoxia (from 60 to 5% oxygen saturation), following any changes of irrigational behaviour, a significant increase of accumulated pleopod activity occurred. A major increase of pleopod activity appeared between 60 and 50% oxygen saturation, below which the activity remained high until a critical point (<10% saturation, 11 °C, 33 psu) where irrigation dropped to a level close to that of normoxic values. Activity sessions during hypoxia were longer and had a higher stroke rate than during normoxia. Received: 22 October 1997 / Accepted: 26 February 1998     

Distribution and relative abundance of Pseudocalanus moultoni and P. newmani (Copepoda: Calanoida) on Georges Bank using molecular identification of sibling species

The sibling species, Pseudocalanus moultoni (Frost, 1989) and P. newmani (Frost, 1989), occur sympatrically on Georges Bank. Taxonomic discrimination of the species relies on subtle morphological characteristics, making routine identification of the species very difficult. DNA sequence variation of two mitochondrial genes, 16S rRNA and cytochrome oxidase I (COI), reliably discriminated P. moultoni and P.␣newmani. Levels of DNA sequence variation for both genes were consistent with those between species of calanoid copepods. A molecular systematic protocol (based on allele-specific PCR amplification) was designed from the COI sequences and used to discriminate females of the two species. The distributions and relative abundances of the two species were mapped for April 1996 based on samples of 15 to 30 females from 12 collections across Georges Bank. The results of this study indicated that P. moultoni females predominated along the northern flank of Georges Bank, while P. newmani females were common on the southern flank, deeper than the 60-m isobath. Received: 16 May 1997 / Accepted: 26 April 1998     

Recovery of benthic communities at Macquarie Island (sub-Antarctic) following a small oil spill

Intertidal communities at Macquarie Island (sub-Antarctic) were re-evaluated seven years after a small oil spill resulting from the grounding of the “Nella Dan”, and six years after the first assessment of biological impact. Sampling was conducted to evaluate community structure in three zones of the exposed rocky shore (upper red, kelp, and lower red) and in samples of Durvillaea antarctica holdfasts. There were no significant differences between the community structure in oiled and control locations in any of the three shore zones, but holdfast macrofaunal communities at oiled sites still showed evidence of impact. Holdfast community-structure in samples from heavily oiled sites showed moderate levels of recovery, with increased abundances of species which were considered sensitive to the oiling in the first post-impact studies and decreased abundances of opportunistic polychaete and oligochaete worms. In contrast, samples collected from the moderately oiled location at Secluded Bay showed little evidence of recovery. Holdfasts at this site were filled with sediment containing traces of diesel oil, and the macrofaunal community was dominated by opportunistic worms. This study clearly indicates that even small incidents of anthropogenic perturbation can have long-lasting consequences for marine communities at Macquarie Island. Received: 23 July 1997 / Accepted: 5 February 1998     

Factors influencing food choice by the seaweed-eating marine snail Norrisianorrisi (Trochidae)

Through two-choice gustatory experiments, a␣preference hierarchy was established␣for␣the␣herbivorous gastropod Norrisianorrisi Sowerby, with laminarialean kelps preferred over all other seaweeds. Among the kelps, laminae of Macrocystispyrifera were slightly preferred over Egregiamenziesii, and both were strongly preferred over sporophylls of Eiseniaarborea. E.arborea, the least preferred kelp, was consistently chosen over other algae common in the snail's habitat (Halidrysdioica, Dictyotaflabellata, and Pterocladiacapillacea) and over seaweeds believed to be edible and palatable based on their morphology, structure, and secondary chemistry (Endarachnebinghamiae, Mazzaellaflaccida, and Ulvalobata). The morphologies and structural toughness of tested seaweeds varied significantly as did their nutritional (% carbon, % nitrogen, C:N ratio, and % ash) contents and phlorotannin concentrations; however, snails preferred to feed on kelps regardless of nutritional content, toughness or phlorotannin concentration; and among kelps preferred to feed on the least tough species (based on penetrometer measurements), which also were those containing the lowest phlorotannin concentrations. Preference for kelp was not upheld in experiments using agarose thalli to which freeze-dried powder, of either the kelp Eiseniaarborea or non-kelp Endarachnebinghamiae was added, suggesting the destruction of attractant chemicals during the making of the artificial foods. Our data suggest that the preference of N. norrisi for kelps over other potentially edible and palatable seaweeds may not be related to nutritional content, but instead may have evolved in response to factors such as availability, habitat provision, or refuge from predation. Received: 27 September 1996 / Accepted: 7 October 1997     

Effects of UV radiation of different wavebands on pigmentation, 15N-ammonium uptake, amino acid pools and adenylate contents of marine diatoms

Three marine diatoms Lauderia annulata Cleve, Odontella sinensis (Greville) Grunow and Thalassiosira rotula Meunier were exposed to ultraviolet (UV) radiation of different wavebands under controlled laboratory conditions (0.035 vol% CO₂, 18 °C). Several changes in the patterns of pigments in these organisms were seen depending on the waveband of UV radiation and species examined. UV-B and UV-B plus UV-A radiation led to a reduction in the overall pigment content of all three diatoms. The uptake of ¹⁵N-ammonium was less affected by 5-h UV-A (WG 320) but significantly reduced after UV-B and UV-B plus UV-A exposure. The pattern of free amino acid pools varied depending on the applied UV wavebands and the tested diatom. The main protein-bound amino acids of T. rotula decreased after 5-h UV irradiance except leucine. Contents of adenosine 5′-mono-, di-, and triphosphate (AMP, ADP and ATP) were affected differently by UV radiation; ATP values increased at the end of UV-B and UV-B plus UV-A exposure. These results have been discussed with reference to the impact of the different UV sources and the influence on the nitrogen metabolism in connection to pigments and supply with energy. Received: 13 May 1997 / Accepted: 11 October 1997     

Instrument Choice When Regulators and Firms Bargain

We compare outcomes with an emissions tax and an emissions standard when a firm and regulator engage in cooperative bargaining over the stringency of the regulation. Bargaining is motivated by giving the firm a choice of abatement technologies. If the firm's preferred technology differs from the regulator's, the first-best outcome is not an equilibrium of the traditional noncooperative game in which the regulator is a Stackelberg leader. The regulator may therefore choose to offer the firm a more lenient regulation if it agrees to switch technologies. We find that the resulting bargaining outcomes differ for a tax and a standard even though information is symmetric, and we identify conditions under which each instrument yields lower social costs.