Genetic differentiation in a pelagic crustacean (Meganyctiphanes norvegica: Euphausiacea) from the North East Atlantic and the Mediterranean Sea

Meganyctiphanes norvegica (M. Sars)is a pelagic crustacean that plays a key role in marine food webs of North Atlantic Ocean and marginal seas. We studied eight population samples collected in the European Atlantic and Mediterranean Sea. By means of single strand conformation polymorphism analysis (SSCP) and direct sequencing, we investigated a segment of 158 base pairs of the mitochondrial gene coding for the subunit 1 of NADH dehydrogenase. We found 12 sequence variants among the 385 individuals studied. Analysis of molecular variance (AMOVA) showed that 14.75% of the total genetic variability was explained by differences between populations, thus indicating absence of panmixia for these populations. Pairwise comparisons revealed three distinct genetic pools: the first one represented by Cadiz Bay, the second one by the Ligurian Sea, and the third one included all the NE Atlantic samples. We also investigated one population from the Alboran Sea (within the Mediterranean basin, east of the Strait of Gibraltar). This population was found to be genetically intermediate between the NE Atlantic samples and the Ligurian sample, suggesting that the restriction to the gene flow is not associated with the Strait of Gibraltar, but possibly with the Oran–Almeria oceanographic front. The present work indicates that M. norvegica, although endowed with a high dispersal capacity because of its pelagic habit, can develop separate breeding units inside the same oceanic basin (the Atlantic). Furthermore, the Ligurian sample should be considered as a distinct evolutionary entity, separated from the Atlantic population. Received: 2 May 1999 / Accepted: 26 November 1999     

Mating behaviour of Macrophthalmus hirtipes (Brachyura: Ocypodidae)

 We examined the mating behaviour of the New Zealand ocypodid crab Macrophthalmus hirtipes in the laboratory between February and June 1998. This species has a discrete breeding season. Mating and moulting were not linked and only intermoult females with mobile gonopore opercula were attractive to males. Allometry and compatibility of gonopods and gonopores of different-sized crabs was investigated. Under laboratory conditions, the opercula of intermoult females remained mobile on average for 11.4 d, but the duration of receptivity did not appear to be under female control. The operational sex ratio in the laboratory fluctuated greatly, but was always male-dominated. During the period of opercular mobility, females mated many times with several different males. Matings in the absence of burrows were relatively short (mean duration = 23 min, max. = 122 min) and the mating behaviour of M. hirtipes lacked courtship and mate-guarding. Males used a search-intercept method to acquire mates, with very low levels of intrasexual competition. There was no evidence of mate preference in M. hirtipes, and males spent just as long mating with ovigerous females as with non-ovigerous ones. Although M. hirtipes has ventral-type spermathecae, as do several other ocypodid crabs, it is unclear whether this promotes last-male sperm precedence. The role of burrows in modifying the mating behaviour of M. hirtipes in the field remains to be established. Received: 7 January 2000 / Accepted: 5 June 2000     

Phylogeny and biogeographic history of hake (genus Merluccius), inferred from mitochondrial DNA control-region sequences

 Phylogenetic analyses of the left domain of the mitochondrial DNA control-region sequence have been used to examine the relationships among species of the genus Merluccius (Rafinesque, 1810), and to compare these with hypotheses based on morphological, meristic and allozyme characters. Analysis of aligned sequences revealed that transition bias was much lower than in mammalian mtDNA, and that nucleotide composition of control-region sequences was biased toward A and T. We have roughly calibrated a molecular clock for the genus, based on the rise of the Isthmus of Panamá, which is believed to have created a barrier to dispersal between marine species of the Atlantic and Pacific Oceans. Our mtDNA-based phylogeny was highly congruent with allozyme-based phylogenies, but poorly so with a previously described phylogeny based on morphology. Specifically, our phylogeny resolved two well-supported principal clades, one of American (west Atlantic and east Pacific) species and the other of Euro–African (east Atlantic) species. This suggests an evolutionary history during which the ancestral lineage of Merluccius was divided between two geographic regions, with subsequent dispersal and vicariant events resulting in the evolution and distribution of extant taxa. However, the relationships between some taxa within the American clade could not be resolved. We suggest that this is consistent with an hypothesis of a rapid origin and radiation of these taxa. Received: 12 December 1998 / Accepted: 15 October 1999     

Grazing rates of organic matter and living fungal biomass of decaying Spartina alterniflora by three species of salt-marsh invertebrates

 The pathway for the flow of salt-marsh grass production into marsh food-webs is still not well defined. We compared the abilities of three marsh macroinvertebrates [salt marsh periwinkles, Littoraria irrorata (Say) (=Littorina irrorata), salt-marsh coffee-bean snails, Melampus bidentatus (Say); and a talitrid amphipod, Uhlorchestia spartinophila Bounsfield and Heard] to access standing-dead leaves of smooth cordgrass (Spartina alterniflora Loisel). The invertebrates were incubated with naturally-decaying leaves, and the rates of removal of organic matter and living fungal biomass (ergosterol) were measured. The impact of invertebrate activity upon fungal growth rates was measured as rates of fungal-membrane synthesis (incorporation of radioacetate into ergosterol). The removal rates of organic leaf biomass per mg individual biomass were highest for amphipods (700 μg mg⁻¹ d⁻¹) and lowest for periwinkles (90 μg mg⁻¹ d⁻¹), but the relatively large biomass of the snails made their removal rates per individual greater than those of amphipods. Net removal of ergosterol by all three invertebrates was >50% for yellow-brown (early-decay) leaf blades. For fully-brown (advanced-decay) blades, >50% removal of ergosterol was found only for periwinkles; exposure to coffee-bean snails and amphipods resulted in a net ergosterol reduction of ≤20%. The lower net reduction of living fungal biomass by coffee-bean snails and amphipods may have been due to fungal-growth stimulation (2.3-fold stimulation in coffee-bean snails and 1.5-fold stimulation in amphipods). Grazing by periwinkles did not stimulate fungal growth, possibly because of its high intensity. Grazing by these three salt-marsh shredders may affect marsh-grass shoot-decay in different ways. Periwinkles may abbreviate the period of fungal production, and incorporate the decaying material relatively quickly into snail biomass and fecal-pellet rain to the sediments. Coffee-bean snails and amphipods may enhance and prolong fungal production, along with the formation of fecal-pellet rain. All three invertebrates fed preferentially on leaf blades rather than leaf sheaths, and feeding rates of gastropods were higher during the night than during the day. Received: 25 November 1998 / Accepted: 4 November 1999     

Has Economic Research Answered the Needs of Environmental Policy?

Efficient environmental regulation requires empirical estimates of the benefits and costs of environmental standards and policies. In the last 25 years, economists have made considerable progress in developing methods for measuring environmental benefits and in providing empirical estimates of costs and benefits. This paper reviews this progress. It examines by category of benefit the advancements made in providing estimates that can be used in benefit-cost analyses and identifies areas where additional research is required.     

Effect of temperature on statolith growth of the European squid Loligo vulgaris during early life

Over the past decade, statolith interpretation has resulted in a major advance in our knowledge of squid population-dynamics, but the way in which environmental conditions affect the statolith increment-deposition ratio remains virtually unknown. The object of the present study was to determine the effect of temperature on this process, using tetracycline marks to validate statolith growth in Loligo vulgaris Lamarck, 1798 under rearing conditions equivalent to severe winter (11 °C) and summer (19 °C) temperature regimes. Tetracycline marking was performed every 10 d (at 10, 20, 30, 40, 50 and 60 d of age). The newly hatched squid paralarvae were slightly smaller in summer than those hatched in winter. Survival rates were similar in both cultures, but growth rates (wet mass) of summer squids were double those in winter. At hatching, statoliths were already longer in the summer squids, and growth rates were 2% d⁻¹ as opposed to 0.9% d⁻¹ for winter statoliths. For the dorsal dome area of the statolith, where more increment counts were made, statolith growth was of 3.25 μm d⁻¹ in summer, and daily increment deposition was confirmed in 87% of the statoliths. The slow growth of statoliths at winter temperatures yielded a mean growth of 1.1 μm d⁻¹– insufficient to discern the increments using light microscopy. Subsequent SEM observation enabled only 21% of the winter statoliths to be read; these also indicated a deposition rate of one increment d⁻¹. Since the life span of L. vulgaris is ≃1 yr, squids will experience at least one winter during their life cycle, and this might be visible on the statolith. Received: 28 June 1999 / Accepted: 20 December 1999     

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     

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