Biomass partitioning of benthic microbes in a Baltic inlet: relationships between bacteria, algae, heterotrophic flagellates and ciliates

 The structure of a benthic microbial food web and its seasonal changes were studied in the shallow brackish waters of the island of Hiddensee, northeastern Germany, at two sites in close proximity by monthly or bimonthly sampling from July 1995 to June 1996. Abundance and biomass of phototrophic and non-phototrophic bacteria, heterotrophic flagellates (HF) and ciliates as well as the biomass of microphytobenthos were determined in the upper 0.3 cm sediment layer. Abundance of organisms showed strong positive correlation with water temperature, with the exception of the bacteria. Non-phototrophic bacterial numbers ranged from 7 × 10⁸ to 6.7 × 10⁹ cells cm⁻³ and phototrophic bacterial abundance from 4 × 10⁷ to 2.7 × 10⁸. Heterotrophic protist abundance ranged from 8 × 10³ to 104 × 10³ ind cm⁻³ for HF and from 39 to 747 ind cm⁻³ for ciliates. The biomass partitioning demonstrated the primary importance of non-phototrophic bacteria (min. 0.83, max. 84.87 μg C cm⁻³), followed by the microphytobenthos (min. 1.32, max. 50.93 μg C cm⁻³). The heterotrophic protists contributed roughly the same fraction to the total microbial biomass, with the biomass of the HF being slightly higher (HF 0.23 to 1.76 μg C cm⁻³, ciliates 0.04 to 1.17 μg C cm⁻³). Taxonomic classification of the benthic HF revealed the euglenids to be the most important group in terms of abundance and biomass, followed by thaumatomastigids and kinetoplastids. Other important groups were apusomonads, cercomonads, pedinellids and cryptomonads. The structure of the HF assemblage showed strong seasonal changes with euglenids being the most abundant taxa in summer, while apusomonads and thaumatomastigids were predominant in winter. Similar to the pelagic microbial food web, benthic picophototrophic bacteria were occasionally abundant, and the feeding modes of heterotrophic protists exhibited a great variety (predominantly omnivores, bacterivores, herbivores or predators). Filter-feeding HF were of little importance. Contrary to the pelagic environment, a top-down control on total benthic bacterial numbers by HF seemed unlikely at the studied stations which were characterised by muddy sand. Received: 6 January 1999 / Accepted: 21 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     

Functional mouthpart morphology of the squat lobster Munida sarsi, with comparison to other anomurans

 The mouthparts of Munida sarsi Huus, 1935, including the labrum and paragnaths, are described using LM and SEM, with special attention to the setal composition and its function. We also used high-resolution macrovideo equipment to record the feeding processes and monitor setae in action. This combination of morphological and preliminary behavioral data enabled us to arrive at a more comprehensive understanding of the functional morphology. M. sarsi was found to be an opportunist and capable of handling and eating most kinds of animal tissue. The species can also be a selective deposit feeder, sorting the sediment by quality and not by size. This sorting is performed by the setae on the medial rim of the bases of maxilla 1, maxilla 2 and maxilliped 1. Many of the other mouthpart functions ranging from food detection to ingestion are mapped with comparison to what has been found for other anomurans. Grooming was shown to be a significant function of many mouthparts, especially the endopod of maxilliped 3. In general the mouthparts closest to the mouth have the least complex movements and functional scheme, with the mandibles being able to perform only one movement, i.e. cutting or crunching food objects. Some functions are attributed to the paragnath, the endopod of maxilla 2 and the epipod of maxilliped 1. Received: 3 October 1999 / Accepted: 22 March 2000     

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     

Allozyme electrophoresis indicates high gene flow between populations of Holothuria (Microthele) nobilis (Holothuroidea: Aspidochirotida) on the Great Barrier Reef

 Genetic variation in 15 Holothuria (Microthele) nobilis (Selenka, 1867) populations on the Great Barrier Reef was studied at seven polymorphic allozyme loci. Although populations were separated by distances up to 1300 km, there were no apparent restrictions to gene flow (F _ST values were not significantly different from 0) and the maximum Nei's unbiased genetic distance was 0.003. Populations were in Hardy–Weinberg equilibrium at all loci. The estimated maximum sexual input [this is the ratio of the number of sexually produced individuals (N*) to the sample size (N _i )] and the minimum sexual input [this is the ratio of the number of genotypes (N _go ) over the sample size (N _i )] were used as estimators for the amount of asexual reproduction. Both parameters suggested that H. nobilis reproduces solely by sexual means (N*:N _i : = 1; N _go :N _i  = 0.74 to 1). The allozyme data indicated high gene flow between populations, but the possibility that allozyme frequencies may not be at equilibrium means that it was not possible to distinguish whether the patterns reflect present-day dispersal or dispersal that occurred in the past. Received: 27 March 2000 / Accepted: 29 June 2000     

Patterns of time and space utilisation in coelacanths (Latimeria chalumnae), determined by ultrasonic telemetry

Acoustic telemetry was used to examine patterns of activity and space utilisation of coelacanths, nocturnal predators which spend the day in submarine caves. Nine coelacanths (Latimeria chalumnae) were tracked, each for a period of 1 to 16 nights at Grande Comore, West Indian Ocean. Activities lasted on average 9 h, usually starting shortly after sunset and ending before sunrise. Vertically, coelacanths moved up and down at and below cave level by following the bottom contour, mainly between 180 and 400 m depth. The deepest record was 698 m, the shallowest 133 m. Most time was spent between 200 and 300 m depth. Large individuals performed deep excursions to depths below 400 m, usually once per night. The fish spent most time in water temperatures of 15 to 19 °C; they rarely ventured into waters warmer than 22 °C measured at depths shallower than 160 m depth. Horizontally, coelacanths stayed in narrow areas ranging from <1 to 10 km of coastline. Coelacanths are extremely slow drift-hunters with an estimated average swimming speed of 3.2 m min⁻¹, often travelling not more than 3 km per night. They probably take advantage of local upwelling and downwelling and slow currents occurring parallel to the steep slopes. This study shows that coelacanths are inhabitants of the subphotic zone, where they are active mainly below the depth of their daytime refuges. Received: 7 July 1999 / Accepted: 11 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