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     

Cadmium accumulation in the female shore crab Carcinus maenas during the moult cycle and ovarian maturation

 The influence of moulting and ovarian maturation on cadmium accumulation in the tissues of female shore crabs Carcinus maenas exposed to 1 mg Cd l⁻¹ in the water was investigated. Cadmium accumulation in all tissues examined was markedly increased in crabs in the postmoult stages (A and B) compared to crabs in all other moult stages. During the moult cycle, average cadmium accumulation in the midgut gland ranged from 29 μg Cd g⁻¹ dw at premoult stage (D₂) to 589 μg Cd g⁻¹ dw at postmoult stage (A). Average cadmium concentrations in the haemolymph ranged from 0.56 μg Cd ml⁻¹ at intermoult stage (C₄) to 4.6 μg Cd ml⁻¹ at postmoult stage (A), while the gills accumulated from 103 μg Cd g⁻¹ dw in intermoult stage (C₃) to 352 μg Cd g⁻¹ dw in postmoult stage (A). Cadmium concentration in gills and haemolymph was also significantly higher in crabs in late premoult stage (D₃) compared to C₄-crabs, while midgut gland cadmium concentration remained elevated in C₁- and C₃- intermoult stages relative to C₄. During ovarian maturation the cadmium accumulation in midgut gland, gills, ovaries and haemolymph decreased. Average cadmium concentration in the midgut gland decreased from 63 μg g⁻¹ dw in ovarian Stage I to 19 μg g⁻¹ dw in ovarian Stage VI. The same pattern was observed for gills, haemolymph and ovaries. The present study demonstrates that cadmium accumulation in the female shore crab strongly depends on the physiological status of the animal. A possible association between physiological calcium requirements and cadmium accumulation during moulting is discussed. Received: 20 January 2000 / Accepted: 20 July 2000     

A comparative study of feeding and growth in two coexisting species of carnivorous gastropods

The effects of variable food supply on growth rate and feeding behaviour in two coexisting intertidal gastropods were examined through field and laboratory experiments involving mark and recapture procedures, growth experiments under controlled food supply and video observations of feeding. All cohorts of both Japeuthria species grew most in summer, with the highest growth rate being achieved by the youngest cohort of both sexes. In contrast, almost all cohorts (except the youngest one) showed zero to negative growth in winter. The growth rates of Japeuthria species clearly varied depending on feeding frequencies and the initial size of individuals. Females of both species had a significantly higher growth rate than males, though the difference became less marked with decreasing food supply. The total time spent on feeding and the length of the first feeding bout generally increased as the duration of the interval between meals was increased across treatments. From both the field growth experiment and the observations of feeding behaviour it is clear that J. ferrea had a higher frequency of feeding than J. cingulata in the field; 1- to 3-d intervals of feeding frequency in the former and 3- to 6-d intervals in the latter. Most notably, the two species showed a reversal in growth rates with changing conditions of food supply. When food was relatively abundant J. ferrea achieved better growth than J. cingulata, but J. cingulata grew better than J. ferrea under low food supply. In view of the fact that J. ferrea seems to be competitively superior to J. cingulata under normal levels, the present observation that J. cingulata can perform better than J. ferrea when food availability is low gives an important advantage to the latter in terms of its coexistence with the former. Received: 9 March 1999 / Accepted: 8 October 1999     

Size-related aspects of arm damage, tissue mechanics, and autotomy in the starfish Asterias rubens

 Arm damage is a widely reported but superficially investigated aspect of the biology of the starfish Asterias rubens L. In the present study, the incidence of arm damage was surveyed in populations of A.  rubens at two sites in the Firth of Clyde, Scotland, and three sites in Gullmarsfjorden, Sweden. The mean (±SD) incidence across all sites of individuals with basal arm damage (resulting from detachment at the basal autotomy plane) was 19.69 ± 8.86%, the incidence of those with distal arm damage (resulting from amputation at more distal levels) was 7.74 ± 10.01%. The mean incidence of arms with basal damage was 5.28 ± 4.12%, of those with distal damage 1.83 ± 2.45%. There was a significant negative correlation between size and the incidence of basal damage at all but one site, but no significant correlation between size and distal damage at any site. Mechanical tests on specimens of the aboral body wall from the basal region of the arm (which included the autotomy plane) and from a more distal region revealed that with increasing body size there was a significant increase in yield stress, ultimate stress and Young's modulus (stiffness) but no significant change in yield strain and ultimate strain. There was no significant difference between the relationships for basal and distal specimens. It is hypothesised that in larger individuals increased mechanical toughness replaces autotomy as an effective antipredator strategy. Using two methods to induce autotomy, a significant positive correlation between size and the delay between the onset of stimulation and arm detachment was found; this may represent a size-related decline in the efficiency of the autotomy mechanism through the relaxation of selection pressure. Since size is an unreliable indicator of age in A.  rubens, the trends identified herein can be interpreted only tentatively as age-associated phenomena. Received: 4 September 1999 / Accepted: 17 February 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     

Manipulation of food and photoperiod promotes out-of-season gametogenesis in the green sea urchin, Strongylocentrotus droebachiensis: implications for aquaculture

Green sea urchins, Strongylocentrotus droebachiensis OF Müller, collected off the coast of New Hampshire, USA, in late February 1995, were brought into the laboratory and fed an artificial diet ad libitum, and subjected to a photoperiod advanced by 4 months. During this study, temperatures and salinities for experimental urchins mirrored those recorded at the collection site. We examined the effects of changes in feeding regime and photoperiod on gametogenesis and compared the experimental urchins with those from the source population. During the 7-month period, experimental urchins showed no detectable changes in mean test height or diameter. Experimental urchins had a significantly higher gonad index (GI) in March, April and May (18 ± 6%) compared with field urchins in March (11 ± 3%). Subsequently, experimental urchins had a mean monthly GI of 25 to 30%, while the mean GI for field urchins was 11 to 13%. Gonial cell mitosis and gametogenesis occurred earlier in experimental male and female urchins compared with field urchins. Stereological and histological observations and stage–frequency data showed that the ovaries of experimental urchins were large because of the accelerated development of nutritive phagocytes, the volume fraction (V _v) of which was 89 to 90% of the gonad, while new vitellogenic primary oocytes occupied <1% V _v. In males, stereological and histological observations and stage–frequency data suggested a mobilization of materials from the nutritive phagocytes beginning between June and August, i.e. earlier than in females, and, by September, new gametes occupied a V _v of 49 ± 3% of the testes. Oocyte size–frequency distributions demonstrated that most primary oocytes were <80 μm in diameter between March and September, suggesting that cold temperatures may be needed for completion of vitellogenesis. We describe changes in the two principal cell types in the germinal epithelium of urchin gonads and indicate how knowledge of their population dynamics may be useful in aquaculture applications. Received: 21 February 1997 / Accepted: 3 June 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     

Effects of Trichodesmium spp. blooms on penaeid prawn larvae

The effects of blooms of the cyanobacterium Trichodesmium spp. on penaeid prawn larvae were examined using in situ and laboratory rearing experiments and plankton surveys in Albatross Bay, Gulf of Carpentaria, Australia. The in situ experiments demonstrated that, during a bloom of Trichodesmium spp., larvae of the prawn Penaeus merguiensis did not develop beyond the first protozoea stage, and survival was low compared with times when diatoms were dominant in the same study area. Laboratory experiments confirmed the in situ results. None of the prawn larvae fed Trichodesmium sp. in laboratory experiments developed beyond the first protozoeal stage. In contrast, 94% of prawn larvae fed the green flagellate Tetraselmis suecica successfully developed to the second protozoea stage. Electron microscopy of larvae gut-contents revealed that Trichodesmium spp. were ingested by larvae but were of no nutritional value, resulting in starvation. A 7 yr plankton survey, from 1985 to 1992, showed that minimum abundance of prawn larvae occurs during the annual summer blooms of Trichodesmium spp. and that maximum abundance of prawn larvae generally occurs just after the bloom. There was a negative correlation between the abundance of larvae and the abundance of Trichodesmium at individual sites, one offshore and one inshore, indicating that the blooms affect the survival of larvae. We conclude that variations in both timing and magnitude of Trichodesmium blooms are important determinants of prawn larvae abundance in Albatross Bay. Received: 28 April 1997 / Accepted: 2 April 1998     

Population biology of Dyopedos monacanthus (Crustacea: Amphipoda) on estuarine soft-bottoms: importance of extended parental care and pelagic movements

The suspension-feeding amphipod Dyopedos monacanthus (Metzger, 1875) is a common epibenthic amphipod that lives on self-constructed “mud whips” (built from filamentous algae, detritus and sediment particles) in estuaries of the northern North Atlantic Ocean. The population biology of D. monacanthus at a shallow subtidal site in the Damariscotta River Estuary (Maine, USA) was examined between July 1995 and July 1997. The resident population at the study site was dominated by adult females during most months of the year. High percentages of subadults were found in late summer/early fall. Often, between 10 and 20% of the adult females were paired with males, and the percentage of ovigerous females varied between 40 and 100%, indicating continuous reproduction. The percentage of parental females varied between 40 and 80% during most months, but dropped to levels below 20% during summer/early fall. The average size of amphipods on their own mud whips was ∼4 mm during the summer/early fall, after which it increased continuously to >7.0 mm in March or April, and then dropped again. In March and April, the average number of eggs and juveniles female⁻¹ was ∼100 eggs and 55 juveniles, while during the summer/early fall the average number of eggs female⁻¹ was <20 and that of juveniles female⁻¹ was <10. Many juveniles grew to large sizes (>1.4 mm) on their mothers' whips in winter/early spring but not in the summer/fall. The average number of amphipods at the study site was low in late summer/early fall (<50 individuals m⁻²), increased steadily during the winter, and reached peak densities of >3000 individuals m⁻² in April 1996 (>1600 individuals m⁻² in May 1997), after which densities decreased again. The decrease of the D.␣monacanthus population at the study site coincided with a strong increase of amphipods found pelagic in the water column. This behavioural shift occurred when temperatures increased and benthic predators became more abundant and active on shallow soft-bottoms, suggesting that D. monacanthus at the study site is strongly affected by predation. The effects are direct (by predation on amphipods) and indirect (by reducing duration of extended parental care and enhancing pelagic movements). Both extended parental care and pelagic movements are important behavioural traits of D.␣monacanthus (and other marine amphipods), and significantly affect its population dynamics. Received: 18 January 1998 / Accepted: 27 May 1998