Regulation of feeding mode by the sand dollar Dendraster excentricus in a shallow estuarine habitat

The sand dollar Dendraster excentricus is a facultative suspension feeder, which is relatively rare within the Order Clypeasteroida. Field studies of regulation of its feeding mode have been mostly conducted in exposed coastal habitats, where they are typically observed in an inclined position and oriented parallel to each other during suspension feeding. Physical (current speed and direction, reduction of drag and lift) and biological factors (increased efficiency of capture of particulates, and density) have been associated with regulation of its feeding mode in exposed coastal systems. We simultaneously measured the role of physical and biological factors in regulating the selection mode for feeding under varying tidal conditions in a shallow estuary in Baja California, Mexico. We used photographic records and direct sampling in fixed plots to determine the relationship between feeding behavior and environmental conditions. Current direction and speed, tidal level, density and content of organic matter in the water column and sediments were measured with respect to feeding mode (prone or inclined) and orientation relative to prevailing currents during spring and neap tides. Multiple regression analysis indicated that the percentage of inclined sand dollars was strongly and positively correlated only with tidal level at the densities found in the estuary (mean < 180 individuals m^− 2); there was no relationship with current velocity, density, and organic matter content of the water. The prone position, indicative of deposit feeding, was largely limited to low tidal levels. We used circular statistics to determine whether the orientation of inclined sand dollars was correlated with current direction and speed. Sand dollars were only oriented parallel to the prevailing currents during the strongest currents of spring tides (> 20 cm s^− 1). We did not observe the predominant oral:aboral configuration found in exposed coastal systems, which may be attributed to the relatively low densities of sand dollars in the exposed coastal environment (30-180 individuals m^− 2), compared to protected coastal habitats (up to 1000 individuals m^− 2). Our results suggest that regulation of the feeding mode of sand dollars in shallow and hydrodynamically complex estuarine systems differs from the feeding mode found in exposed coastal environments.     

Differential susceptibility of marine invertebrate larvae: Laboratory predation of sand dollar, Dendrasterexcentricus (Eschscholtz), embryos and larvae by zoeae of the red crab, Cancer productus Randall

Predation on eggs, embryos, and larvae of the sand dollar, Dendraster excentricus (Eschscholtz) was investigated in a series of laboratory feeding experiments. Dendraster susceptibility to predation by zoea larvae of the red crab, Cancer productus Randall was strongly dependent on developmental stage and ontogenetic differences in motility. Clearance rates by C. productus were highest for eggs and averaged 0.551·zoea⁻¹·day⁻¹. Embryos and prism larvae of Dendraster were consumed at an intermediate rate, while pluteus larvae were captured at a relatively low rate. Clearance rates decreased from 0.18 to 0.031·zoea⁻¹·day⁻¹ during the transition of prism larvae into echinoplutei. Differences in Dendraster susceptibility to predation cannot be attributed to increasing prey body size because dwarf plutei were captured at the same rate as normal plutei. Reduced capture rates by Cancer productus zoeae are dependent on the development of Dendraster swimming behavior. Periodic reversals in the direction of ciliary beating and backwards swimming effectively remove Dendraster plutei from the immediate capture sphere of Cancer productus. Reversed swimming appears to function as a post-contact encounter response that reduces the mortality rates of Dendraster plutei.     

Causes of sulfur isotope variability in the seagrass, Zostera capricorni

Sulfur has been proposed as a useful element to employ in addition to carbon and nitrogen in stable isotope studies of marine food webs, but variability in δ³⁴S of primary producers may prevent food web resolution. δ³⁴S values in green leaves of the seagrass, Zostera capricorni, showed considerable variability (12.7-17.6‰) in a survey in Moreton Bay, Australia. We demonstrated that δ³⁴S values were correlated with sediment organic matter (OM) content and height of seagrass on the tidal gradient, but these relationships were opposite to those expected from work elsewhere. In our survey, δ³⁴S values were relatively depleted at sites higher on the shore and with lower OM content. We did find the expected relationship of depleted δ³⁴S values where sediment porewater sulfide concentrations were higher. Any influence of OM content on δ³⁴S values would have been confounded in the survey by the relationship between height on shore and OM content itself. We separated the effects of height and OM content by creating the following treatments at one height on the shore: (1) OM added, (2) procedural control, and (3) untouched control. δ³⁴S values of seagrass in OM added plots were significantly depleted (5.6‰) relative to procedural (10.1‰) and untouched (11.0‰) controls 8 weeks after the manipulation. This demonstrated that OM content on its own does have the expected effect on δ³⁴S values of seagrass, so in the initial survey another factor, probably related to height on shore, must have overridden the influence of OM content. Seagrass roots are able to exude excess oxygen produced during photosynthesis, reoxidising sulfides in surrounding porewater. We demonstrated that the above and below-ground biomass of seagrass was higher low on the shore, and contend that higher seagrass productivity low on the shore results in greater reoxidation of sulfides and leads to more enriched δ³⁴S values of seagrass.     

The role of sediment organic matter composition in larval habitat selection by the polychaete Capitella sp. I

The role of sediment organic matter composition in larval habitat selection by the polychaete Capitella sp. I was investigated using a still water multiple-choice experiment. Specifically, the following question was addressed: what are the roles of sediment labile organic compounds in larval habitat selection? A replicated Latin-square design was used to test larval preferences among sediments collected from five sites (TLC—Tung Lung Chau; VHE—Victoria Harbor east; VH—Victoria Harbor; VHW—Victoria Harbor west; PC—Peng Chau) in the vicinity of Victoria Harbor, Hong Kong. Although mean grain size was not different among sites, water content and the total organic matter were higher in VH Mud but were similar among other sites. Carbohydrate, lipid, and biopolymeric fractions followed the same trend. According to protein content, sites were ranked as: VH > VHW > TLC = VHE > PC. PC Mud had the highest carbohydrate/protein ratio, due to low concentration of protein. According to larval preference, sites were ranked as: TLC = VH = VHW = VHE > PC. Our study shows a precise relationship between sediment biochemical properties and larval habitat selection. Larvae tend to choose sediments with low carbohydrate/protein ratio. Results suggest that types of sediment organic matter may play more important role in the larval habitat selection process of Capitella sp. I than concentration of organic matter alone.     

Diet and physiological responses of Spondyliosoma cantharus (Linnaeus, 1758) to the Caulerpa racemosa var. cylindracea invasion

Marine invasions are a worldwide problem that involves changes in communities and the acclimation of organisms to them. The invasive Chlorophyte Caulerpa racemosa var. cylindracea is widespread in the Mediterranean and colonises large areas from 0 to 70 m in depth. The omnivorous fish Spondyliosoma cantharus presents a high frequency of occurrence of C. racemosa in the stomach contents at invaded areas (76.3%) while no presence of C. racemosa was detected in control areas. The isotopic composition of muscle differed significantly between invaded and non-invaded sites for δ¹³C (− 16.67‰ ± 0.09 and − 17.67‰ ± 0.08, respectively), δ¹⁵N (10.22‰ ± 0.22 and 9.32‰ ± 0.18, respectively) and the C:N ratio (2.01 ± 0.0002 and 1.96 ± 0.009, respectively). Despite the high frequency of occurrence of C. racemosa in the stomach contents of S. cantharus and its important contribution to the δ¹³C source (20.7% ± 16.2), the contribution of C. racemosa to the δ¹⁵N in S. cantharus food sources was very low (6.6% ± 5.8). Other invertebrate prey such as decapods and polychaetes were more important contributors to the δ¹⁵N source at both invaded and non-invaded sites. Activation of enzymatic pathways (catalase, superoxide dismutase, glutathione-s-tranferase, 7-ethoxy resorufin O-de-ethylase) but not a significant increase in lipid peroxidation MDA (0.49 ± 0.01 nmol/mg prot at non-invaded and 0.53 ± 0.01 nmol/mg prot at invaded sites) was observed in S. cantharus individuals living in C. racemosa-invaded sites compared with control specimens. The low δ¹⁵N contribution values of C. racemosa by S. cantharus together with the toxicity demonstrated by the activation of the antioxidant defences and the important contribution of invertebrate prey to the δ¹⁵N could mean that the ingestion of C. racemosa by S. cantharus might be unintentional during the predation of invertebrate preys living underneath the entanglement of the C. racemosa fronds and stolons mats.     

An insight into the feeding ecology of deep-sea canyon nematodes — Results from field observations and the first in-situC feeding experiment in the Nazaré Canyon

Submarine canyon systems provide a heterogeneous habitat for deep-sea benthos in terms of topography, hydrography, and the quality and quantity of organic matter present. Enhanced meiofauna densities as found in organically enriched canyon sediments suggest that nematodes, as the dominant metazoan meiobenthic taxon, may play an important role in the benthic food web of these sediments. Very little is known about the natural diets and trophic biology of deep-sea nematodes, but enrichment experiments can shed light on nematode feeding selectivity and trophic position. An in-situ pulse-chase experiment (Feedex) was performed in the Nazaré Canyon on the Portuguese margin in summer 2007 to study nematode feeding behaviour. ¹³C-labelled diatoms and bacteria were added to sediment cores which were then sampled over a 14-day period. There was differential uptake by the nematode community of the food sources provided, indicating selective feeding processes. ¹³C isotope results revealed that selective feeding was less pronounced at the surface, compared to the sediment subsurface. This was supported by a higher trophic diversity in surface sediments (Θ⁻¹ = 3.50 ± 0.2) compared to the subsurface (2.78 ± 0.6), implying that more food items may be used by the nematode community at the sediment surface. Predatory and scavenging nematodes contributed relatively more to biomass than other feeding types and can be seen as key contributors to the nematode food web at the canyon site. Non-selective deposit feeding nematodes were the dominant trophic group in terms of abundance and contributed substantially to total nematode biomass. The high levels of ‘fresh’ (bioavailable) organic matter input and moderate hydrodynamic disturbance of the canyon environment lead to a more complex trophic structure in canyon nematode communities than that found on the open continental slope, and favours predator/scavengers and non-selective deposit feeders.     

Effects of density dependent sex allocation on the dynamics of a simultaneous hermaphroditic population: Modelling and analysis

In this work we present a mathematical model describing the dynamics of a population where sex allocation remains flexible throughout adult life and so can be adjusted to current environmental conditions. We consider that the fractions of immature individuals acquiring male and female sexual roles are density dependent through nonlinear functions of a weighted total population size. The main goal of this work is to understand the role of life-history parameters on the stabilization or destabilization of the population dynamics.The model turns out to be a nonlinear discrete model which is analysed by studying the existence of fixed points as well as their stability conditions in terms of model parameters. The existence of more complex asymptotic behaviours of system solutions is shown by means of numerical simulations.Females have larger fertility rate than males. On the other hand, increasing population density favours immature individuals adopting the male role. A positive equilibrium of the system exists whenever fertility and survival rates of one of the sexual roles, if shared by all adults, allow population growing while the opposite happens with the other sexual role. In terms of the female inherent net reproductive number, η_F, it is shown that the positive equilibria are stable when η_F is larger and closed to 1 while for larger values of η_F a certain asymptotic assumption on the investment rate in the female function implies that the population density is permanent. Depending on the other parameters values, the asymptotic behaviour of solutions becomes more complex, even chaotic. In this setting the stabilization/destabilization effects of the abruptness rate in density dependence, of the survival rates and of the competition coefficients are analysed.     

Stable carbon and nitrogen isotopic discrimination in whole blood of red-throated ant tanagers Habia fuscicauda

Analysis of stable isotope ratios is increasingly used to reconstruct diets in passerine birds, but studies of diet–tissue isotopic discrimination for this avian group are scarce. We determined ¹⁵N and ¹³C diet–tissue discrimination factors on whole blood in the red-throated ant tanager (Habia fuscicauda), an insectivorous–frugivorous passerine. Birds were fed an isotopically uniform, semi-synthetic diet of dog puppy dry food, soy protein isolate, wheat germ, and other ingredients, during 92 days. Average (± SD) diet–tissue discrimination was 2.6 ± 0.2‰ for N and 2.2 ± 0.1‰ for C. Nitrogen diet-tissue discrimination was similar to the values found previously in other passerines fed animal protein and it can probably be used to accurately reconstruct protein dietary origin in passerines feeding on animal protein (e.g., insects). In the case of C, diet reconstruction might be affected by metabolic routing of dietary nutrients.     

Effect of prey density on behaviour and development of the predatory mosquito,Toxorhynchites towadensis

The effect of prey density on feeding behaviour, killing behaviour, and development of the predatory mosquito,Toxorhynchites towadensis, was investigated in the laboratory. The number of prey consumed per larva increased toward an upper asymptote as prey density increased. Prey consumption curves during fourth instar were concave at low prey densities but convex at high prey densities. This phenomenon was not observed during other instars. Killing without consuming any part of prey occurred at prey densities of 20 per container and over. The number of prey killed but not consumed increased linearly with the number of unconsumed prey in the container. Prey acquisition behaviour was not affected by prey densities during the prepupal period. Developmental time from first instar to adult emergence decreased with increasing prey densities, but remained constant at densities of 10 per container and over. Adult size increased with increasing prey densities but there was no effect at prey densities of 20 and over.     

Interannual variations of reproductive parameters in a booted eagle (Hieraaetus pennatus) population: the influence of density and laying date

A population of booted eagles (Hieraaetus pennatus) consisting of 21–29 pairs was studied for 7 years (1998–2004) in a forested mountainous region of southeastern Spain. The average egg-laying time was 25 April (earliest 31 March, latest 26 May). Sixty-eight percent of territorial pairs produced eggs, and 81% of reproductive pairs successfully raised at least one offspring. Mean clutch size was 1.91 eggs, and 84% hatched. The survival rate of chicks was high, with only 17% dying. The major known cause of nestling mortality was predation, mainly by the Eurasian eagle owl (Bubo bubo). The mean number of flying offspring was 0.91 per territorial pair, 1.29 per reproductive pair and 1.65 per successful pair. All population breeding parameters analysed were density independent, but most showed a significantly negative relationship with mean laying date.     

On the embryology and mode of reproduction of selected diploid species of Hieracium s.l. (Asteraceae) from Bulgaria

Three diploid taxa – Hieracium transylvanicum (subgenus Hieracium), Hieracium caespitosum subsp. brevipilum and Hieracium pavichii (subgenus Pilosella) – from five natural Bulgarian populations were investigated embryologically. The peculiarities of the male and female gametophytes, embryo- and endospermogenesis were established in each species. The results suggest that the species propagate sexually as expected from their diploid chromosome number. However, some forms of apomixis have also been observed, e.g. somatic apospory and integumental embryony in H. pavichii. The presence of apomixis in a diploid taxon shows that polyploidy is not an obligatory prerequisite for apomixis and the two phenomena are independent. The embryological plasticity detected in H. pavichii and H. transylvanicum suggests they may have higher opportunities for adaptation and successful reproduction.     

Ecology of phytoplankton communities dominated by Aureococcus anophagefferens: the role of viruses, nutrients, and microzooplankton grazing

We investigated the impact of viruses, nutrient loading, and microzooplankon grazing on phytoplankton communities in two New York estuaries that hosted blooms of the brown tide alga Aureococcus anophagefferens during 2000 and 2002. The absence of a bloom at one location during 2002 allowed for the fortuitous comparison of a bloom and non-bloom year at the same location as well as a comparison of two sites experiencing bloom and non-bloom conditions during the same year. During the study, blooms were found at locations with high levels of dissolved organic nitrogen and lower nitrate concentrations compared to a non-bloom location. Experimental additions of inorganic nitrogen and phosphorus yielded growth rates within the total phytoplankton community which significantly exceeded control treatments in 83% of experiments, while A. anophagefferens experienced significantly increased growth during only 20% of experimental inorganic nutrient additions. Consistent with prior research, these results suggest brown tides are not caused by eutrophication, but instead are more likely to occur when sources of labile DOM are readily available. Microzooplankton grazing rates on the total phytoplankton community during a bloom were lower than grazing rates at a non-bloom site, and grazing rates on A. anophagefferens were lower than grazing rates on the total community on some dates, suggesting that reduced grazing mortality may also promote brown tides. Mean densities of viruses during blooms (3 × 10⁸ ml⁻¹) were elevated compared to most estuarine environments and were twice the levels found at a non-bloom site. Experimental enrichment of the natural viral densities yielded a significant increase in A. anophagefferens growth rates relative to control treatments when background levels of viruses were low (<1.7 × 10⁸ ml⁻¹), suggesting that viruses may promote bloom occurrence by regenerating DOM or altering the composition of microbial communities.     

Habitats, abundance patterns and isotopic signals of morphotypes of the planktonic foraminifer Globigerinoides ruber (d'Orbigny) in the eastern Mediterranean Sea since the Marine Isotopic Stage 12

The chemical composition of shells of the planktonic foraminifer Globigerinoides ruber (white) is frequently used to determine past sea surface conditions. Recently, it has been shown that arbitrarily defined morphotypes within this species exhibit different chemical and isotopic signatures. Here, we investigate the occurrence through time and in space of morphological types of G. ruber (white) in late Quaternary and Holocene sediments of the central and the eastern Mediterranean Sea. In 115 samples representing two distinct time intervals (MIS 1–2 and MIS 9–12) at ODP Site 964 and the piston core GeoTü-SL96, we have defined three morphological types within this species and determined their relative abundances and stable isotopic composition. A quantitative analysis of morphological variation within G. ruber (white) in four samples revealed that the subjectively defined morphotypes occupy separate segments of a continuous and homogenous morphospace. We further show that the abundance of the morphotypes changes significantly between glacials and interglacials and that the three morphotypes of G. ruber show significant offsets in their stable isotopic composition. These offsets are consistent within glacial and interglacial stages but their sign is systematically reversed between the two Sites. Since the isotopic shifts among the three G. ruber morphotypes are systematic and often exceed 1‰, their understanding is essential for the interpretation of all G. ruber-based proxy records for the paleoceanographic development of the Mediterranean during the late Quaternary.     

Effect of salinity, temperature, organic and inorganic nutrients on growth of cultured Fibrocapsa japonica (Raphidophyceae) from the northern Adriatic Sea

The effects of temperature, salinity, inorganic and organic nutrients on the growth and physiological performance of Fibrocapsa japonica have been investigated in strains isolated from the northern Adriatic Sea, where intense and regular blooms have occurred since 1997 in coastal waters during summer. Strains isolated at different times and from different locations appeared homogeneous in terms of both physiological responses and molecular (ITS-5.8S rDNA) characteristics. Growth rates were higher at temperatures between 20 and 26 °C and in a salinity range of 30–35 (0.7 div day⁻¹). The temperature of 16 °C inhibited growth, more markedly at the lowest and highest salinity values, a result also confirmed by a lower photosynthetic efficiency and by an increase in cell volume due to impaired division. Higher cell concentrations were obtained with macronutrients at f/2 levels than in a fivefold diluted medium. Comparing the utilization of 200 μM nitrate to that of different N sources (inorganic and organic), F. japonica showed an efficient growth with equivalent amounts of ammonia, urea and amino acids, such as glycine and tryptophan; glutamate was less effective, while methionine had toxic effects. Organic phosphate, administered as glycerophosphate, could also sustain F. japonica growth, probably on account of an alkaline phosphatase whose activity was enhanced in the presence of the organic form. Vitamins were necessary for growth, though no further stimulation was observed when a surplus of vitamin B12 (3 nM) was added. The addition of 11.7 μM iron instead of 2.3 μM, as well as that of humic acid, with or without macronutrients, did not enhance algal growth either. These results led us to hypothesise that F. japonica blooms became more frequent due to a general seawater temperature increase and to the availability of organic forms which, in coastal anthropized areas, are especially abundant in summer periods.