Sediment disturbance and loss of beta diversity on subtidal rocky reefs

How changes in environmental complexity and heterogeneity affect beta diversity is poorly known. We investigated patterns of beta diversity in subtidal assemblages of algae and invertebrates in the northwest Mediterranean in relation to inclination of the substratum and sedimentation. Vertical and horizontal substrata supported distinct assemblages under low, but not under heavy, ambient loads of sediment. To test the hypothesis that sediment reduced the dissimilarity between assemblages, sedimentation was increased experimentally in plots established on vertical and horizontal surfaces at sites experiencing low ambient levels of sedimentation. Patterns were compared to those occurring at unmanipulated sites and at sites exposed to heavy loads of sediment about 2 km apart. After one year, assemblages on vertical substrata were indistinguishable from those occurring on flat surfaces at manipulated sites and both converged toward those occurring at sites exposed to heavy loads of sediment. Control sites still supported distinct assemblages on vertical and horizontal substrata by the end of the experiment. Similar effects of sediment were observed on recovering assemblages in experimental clearings. These results show that sediment increased similarity in assemblages overriding the influence of habitat complexity on beta diversity at small and large spatial scales.     

Settlement, recruitment and potential predators and competitors of juvenile echinoderms in the rocky subtidal zone

Recruitment patterns of marine invertebrates are affected both by settlement and early post-settlement events. This study examined the settlement and recruitment patterns of echinoderms at three sites in the rocky subtidal zone of Bocabec Cove, Bay of Fundy, Canada using artificial turf collectors and quadrats on the natural substrate. Potential predators were quantified at two of the sites along transects and in 1-m² quadrats. Both potential predators and competitors were quantified in 0.0625-m² quadrats. Settlement varied across sites (1.5–3 km apart) and two years of sampling (2004, 2005). The site of most potential settlement differed for the three groups of echinoderms: sea urchins (Strongylocentrotus droebachiensis), sea stars (Asterias spp.) and sea cucumber (Psolus fabricii). Settlement densities on the artificial turf collectors tended to be greater than the densities of settlers on the natural substrate. On the natural substrate, the only significant difference between densities of juveniles over time was that newly settled sea stars were found in July and were not found the following October. Large lobsters and carnivorous worms were potential predators with densities that varied between sites. Potential competitors that differed in abundance between sites were herbivorous gastropods and conspecifics for sea urchins; and carnivorous worms for sea stars. This study suggests that patterns of recruitment are either set up by patterns of settlement or by events during the first few weeks/months on the benthic substrate for these echinoderms.     

Impact of predation-disturbance by large epifauna on sediment-dwelling harpacticoid copepods: Field experiments in a subtidal seagrass bed

The hypothesis was tested that predation-disturbance by epibenthic macrofauna affects the abundance of sediment-dwelling harpacticoid copepods. The copepods inhabited a subtidal seagrass (Zostera marina L.) bed in British Columbia, Canada. The response of the harpacticoid community was observed in controlled field experiments in which epibenthic predators-disturbers were excluded from portions of the seagrass bed. Controlled, exclusion-cage (0.8 m² area, 7-mm mesh) experiments were conducted within the seagrass bed from late March/early April to mid-June in both 1986 and 1987. Sampling was conducted biweekly. Exclusion of large epibenthic predators-disturbers had little effect on sediment-dwelling harpacticoid copepod density. Total harpacticoid numbers and abundances of dominant species generally did not increase in the exclusion treatment relative to the control. Shifts in species composition of the harpacticoid community did not occur. The treatment control was adequate in simulating the exclusion cage structure. It appears that large epibenthic predators-disturbers have little impact on the abundance of harpacticoid copepod populations at this study site.     

Ovarian morphology and oogenesis in Aurelia aurita (Scyphozoa: Semaeostomae): ultrastructural evidence of heterosynthetic yolk formation in a primitive metazoan

In Aurelia aurita, the ovaries arise as horseshoe-shaped evaginations of the gastrodermis in the floor of four interradial gastric pouches. Germ-cell islands arise within endodermally-derived gastrodermal cells. Oocytes grow and gradually bulge into the mesoglea while maintaining physical contact throughout vitellogenesis with specialized cells called trophocytes. Ultrastructural changes suggest that these cells transport yolk precursors from the coelenteron to the oocytes in a manner similar to that reported for the trophonema cells of anthozoan ovaries. Vitellogenesis involves both the autosynthetic activity of the oocyte organelles (Golgi complex and rough endoplasmic reticulum) and the heterosynthetic incorporation of precursors through endocytotic processes involving both coated pits and vesicles and smooth-surfaced tubules. Ultrastructural data suggest that different types or classes of yolk precursors enter the oocyte through the trophocytes and via the surrounding mesoglea. To our knowledge, this is the most primitive animal in which this type of yolk synthesis has been described. The trophocyte-oocyte relationship in oocytes of A. aurita is reminiscent of the trophonema-oocyte relationship in anthozoans and supports the belief that the Anthozoa and Scyphozoa share a close phylogenetic relationship.     

Role of female territoriality in social and mating systems of Canthigaster valentini (Pisces: Tetraodontidae): evidence from field experiments

Sharpnose puffers, Canthigaster valentini (Pisces: Tetraodontidae) at Lizard Island, Australia, live in made-dominated haremic social and mating systems. The hypothesis was that mature females are restricted in their movements and can be monopolized by some males. Field experiments at Lizard Island, Great Barrier Reef, between January and March 1983 showed that mature females were still territorial in the absence of males and movements of females were not controlled by males. Males abandoned their territories when their females were removed. The territorial behavior of those males with access to females (territorial males) restricted the access of other males (bachelor males) to them. Bachelor males took over harems and became territorial males when established territorial males were removed. The results of the experiments thus supported the hypothesis.     

A gradient diver for measurement of respiration in individual organisms from the micro- and meiofauna

A gradient diver method suitable for gasometric studies on aquatic organisms of 1.0 g–1.0 mg wet weight is described. The microgasometer consists of an open glass capillary (diver) which encloses the biological object and an air bubble. It floats in an aqueous density gradient with constant pressure above the gradient. In response to changes in the volume of the air bubble, caused by consumption or release of gas by the biological object, the diver migrates either downwards or upwards in the gradient. On the basis of the rate of migration the rate of gas exchanges is calculated. The method is adaptable to a wide range of laboratory conditions and should also be of use in field laboratories.     

Friendships between males and lactating females in a free-ranging group of olive baboons (Papio hamadryas anubis): evidence from playback experiments

Close association between an anoestrous female at the time of lactation and adult male(s) is relatively rare in mammals, but common in baboons (Papio hamadryas subsp.). The functional significance of these “friendships” remains unclear, however. In chacma baboons (P. h. griseipes), friendships are a counter-strategy to infanticide by immigrant males. Experimental playback of female distress calls in chacma baboons revealed that male friends are more motivated to protect females and infants than are control males. Olive baboons (P. h. anubis) also exhibit friendships, but infanticide is rare, suggesting that friendships provide females with protection from non-lethal aggression (anti-harassment hypothesis) or serve to promote male–infant bonds that later benefit the maturing juvenile (future male caretaker hypothesis). We replicated these playback experiments on a group of olive baboons to test between these hypotheses and to evaluate if the lower costs of non-lethal harassment lessens male protective responsiveness relative to protection from (more costly) infanticide. Spatial data revealed that most lactating females had one to four friend males. Relative to non-friends, friend dyads were characterized by higher rates of allogrooming and infant handling, but less agonism. Female rank was correlated with the number of male friends. Just as in chacma baboons, playback of female screams elicited stronger responses from male friends than control males in support the anti-harassment hypothesis. Compared to the chacma baboon, male olive baboons appeared to exhibit similarly high levels of protective solicitude for female friends although they protect against non-lethal harassment rather than infanticide.     

Trophic relationships of white-chinned petrels from Crozet Islands: combined stomach oil and conventional dietary analyses

The diet of white-chinned petrels Procellaria aequinoctialis breeding at the Crozet Archipelago (southern Indian Ocean) was studied using two complementary methods: lipid analysis of stomach oils as trophic markers together with the conventional dietary approach (i.e., stomach content analysis). Objectives were (1) to investigate the adult diet when they feed for themselves by analyzing stomach oil lipids, and (2) to compare the lipid signature of chick and adult oils. Stomach oils mainly consisted of triacylglycerols (TAG), diacylglycerol-ethers (DAGE) and wax esters (WE) (66, 14 and 11%, respectively). The dietary origin of TAG and WE was evaluated by linear discriminant analyses with fatty acid and fatty alcohol fractions. Analyses evidenced that stomach oils did not originate from Antarctic krill, but instead from myctophid fish, thus demonstrating the importance of mesopelagic fish in the nutrition of adult petrels. This result was consistent with the identification of digested remains of myctophids recovered from adult stomach contents after long foraging trips. Large amounts of a rare lipid class, DAGE (up to 76% of total lipids), were identified in two stomach oils, together with fresh remains of the squid Gonatus antarcticus (99% by mass), suggesting that DAGE could have the potential to be trophic markers of cephalopods. Moreover, six oils probably originated from Patagonian toothfish, thus confirming strong interactions between white-chinned petrels and fisheries. Comparison between chick and adult stomach oils indicated no major differences in their biochemical composition suggesting an identical dietary origin of oils, mainly myctophids. Both adult and chick oils can therefore be used to determine the feeding ecology of adult birds when they feed far away from their breeding grounds. Finally, food analysis of chick samples and adult samples collected after short and long trips indicated different foraging grounds during the two kinds of trips, and also between long trips performed in subtropical and Antarctic waters.     

Trophic ecology of Grey-headed albatrosses from Marion Island, Southern Ocean: insights from stomach contents and diet tracers

During chick-rearing, albatrosses can alternate between long foraging trips that provide the main source of food for the adults and short foraging trips that they use to feed their young. This flexibility in foraging behaviour can lead to differences in diet composition between adults and chicks and implies that they may be vulnerable in different ways to food shortages. The trophic ecology of the Grey-headed albatross Thalassarche chrysostoma was investigated at the sub-Antarctic Prince Edward Islands during the chick-rearing period in April 2006 using a combination of approaches. Diets of adults and chicks were assessed using stable isotope ratios and fatty acid (FA) profiles of blood and/or stomach oils, in addition to stomach contents analysis. Fish from the family Macrouridae and cephalopods (particularly the onychoteuthid Kondakovia longimana) were the primary prey, whereas crustaceans (krill Euphausia superba) represented a smaller proportion of the stomach contents. Stomach oil FA profiles contained more monounsaturated FA than the profiles of plasma, which were richer in saturated FA and arachidonic acid (20:4n-6). There was also a distinct separation of adults from chicks, with higher levels of monounsaturates in chick plasma, and higher saturated FA levels (particularly 16:0) in the adult plasma. Stable carbon isotope ratios of whole blood were similar in adults and chicks, whereas stable nitrogen isotope ratios showed significant enrichment by >1‰ in chicks. The combined FA, stable isotopes and stomach contents analyses suggest clear differences in diet quality between adults and chicks, with chicks feeding at a higher trophic position through feeding more on highly nutritious fish and adults keeping much of the less nutritious zooplankton for themselves.     

Use of pelagic prey subsidies by demersal predators in rocky reefs: insight from movement patterns of lingcod

Allochthonous subsidies of energy and nutrients can affect community structure in patchy marine habitats, including rocky reefs, and their ecological consequences may depend on the mechanism of energy transfer. Lingcod (Ophiodon elongatus) are demersal predators that trophically link nearshore rocky reefs with offshore pelagic habitats through consumption of pelagic fishes. We quantified lingcod habitat use and movement patterns to make inferences about the temporal and spatial conditions under which lingcod may acquire pelagic prey. Lingcod maintained small home ranges (21,272 ± 13,630 m²) within a rocky reef in the San Juan Archipelago, Washington; eight of nine individuals used rocky habitat exclusively. Depths occupied by lingcod (0–50 m) coincided with pelagic fish distribution on the rocky reef; however, diel patterns in lingcod activity varied inversely with occurrence of pelagic fishes on the reef. Our findings suggest that the pelagic subsidy to lingcod is not strongly mediated through directed off-reef foraging by lingcod.     

Functional morphology and zonation of three species of sea anemones from rocky shores in southern Chile

Three species of sea anemones, clearly exhibiting zonation on the shore, are grouped into intertidal [Phymactis clematis (Drayton, 1846), Anthothoe chilensis (Lesson, 1830)] and infralittoral [Antholoba achates (Drayton, 1846)] forms. A series of observations revealed that intertidal forms have the ability to retain water in the coelenteron during exposure to air,/while the infralittoral form cannot do so. This different behaviour is attributed to morphological characteristics, such as the specific body shape and the structure of the sphincter and retractor muscles. These features, in combination with specific abiotic factors of the various habitats under consideration, determine the upper limits of distribution of each species, while zonation below these limits seems to be affected in addition by biological interactions.     

Field evidence of decreased extractability of copper and nickel added to soils in 6-year field experiments

The phytotoxicity of added copper (Cu) and nickel (Ni) is influenced by soil properties and field aging. However, the differences in the chemical behavior between Cu and Ni are still unclear. Therefore, this study was conducted to investigate the extractability of added Cu and Ni in 6-year field experiments, as well as the link with their phytotoxicity. The results showed that the extractability of added Cu decreased by 6.63% (5.10%–7.90%), 22.5% (20.6%–23.9%), and 6.87% (0%–17.9%) on average for acidic, neutral, and alkaline soil from 1 to 6 years, although the phytotoxicity of added Cu and Ni did not change significantly from 1 to 6 years in the long term field experiment. Because of dissolution of Cu, when the pH decreased below 7.0, the extractability of Cu in alkaline soil by EDTA at pH 4.0 could not reflect the effects of aging. For Ni, the extractability decreased by 18.1% (10.1%–33.0%), 63.0% (59.2%–68.8%), and 22.0% (12.4%–31.8%) from 1 to 6 years in acidic, neutral, and alkaline soils, respectively, indicating the effects of aging on Ni were greater than on Cu. The sum of ten sequential extractions of Cu and Ni showed that added Cu was more extractable than Ni in neutral and alkaline soil, but similar in acidic soil.

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Activity patterns and predatory behavior of an intertidal nemertean from rocky shores: Prosorhochmus nelsoni (Hoplonemertea) from the Southeast Pacific

Understanding the impact of environmental stressors on predator activity is a prerequisite to understanding the underlying mechanisms shaping community structure. The nemertean Prosorhochmus nelsoni is a common predator in the mid-intertidal zone on rocky shores along the Chilean coast, where it can reach very high abundances (up to 260 ind m⁻²) in algal turfs, algal crusts, barnacle crusts, and mixed substrata. Tidal and diurnal scans revealed that the activity of P. nelsoni is primarily restricted to night and early-morning low tides and is relatively low when air temperatures are high. On average, larger worms crawled faster than smaller worms, with their maximum velocity being influenced by substratum type. Their estimated rate of predation is 0.092 prey items nemertean⁻¹ day⁻¹, just below the laboratory rate of ~0.2 amphipods nemertean⁻¹ day⁻¹ previously estimated for this species. P. nelsoni consumes a diverse spectrum of prey items (i.e., amphipods, isopods, decapods, barnacles, and dipterans) and is possibly exerting a significant influence on its prey populations. We suggest that the opportunistic predatory behavior of this intertidal predator is caused by the trade-off between immediate persistence (e.g., avoidance of desiccation) and long-term survival through successful foraging.     

Trophic levels and trophic tangles: the prevalence of omnivory in real food webs

The concept of trophic levels is one of the oldest in ecology and informs our understanding of energy flow and top-down control within food webs, but it has been criticized for ignoring omnivory. We tested whether trophic levels were apparent in 58 real food webs in four habitat types by examining patterns of trophic position. A large proportion of taxa (64.4%) occupied integer trophic positions, suggesting that discrete trophic levels do exist. Importantly however, the majority of those trophic positions were aggregated around integer values of 0 and 1, representing plants and herbivores. For the majority of the real food webs considered here, secondary consumers were no more likely to occupy an integer trophic position than in randomized food webs. This means that, above the herbivore trophic level, food webs are better characterized as a tangled web of omnivores. Omnivory was most common in marine systems, rarest in streams, and intermediate in lakes and terrestrial food webs. Trophic-level-based concepts such as trophic cascades may apply to systems with short food chains, but they become less valid as food chains lengthen.     

Investigations on epibenthic meiofauna I. Abundances on and repopulation of the tube-caps of Diopatra cuprea (Polychaeta: Onuphidae) in a subtropical system

Cryptic meiofauna populated the imbricate, shell-sediment matrix of the tube-caps produced by the polychaete Diopatra cuprea (Bose) in Tampa Bay, Florida, USA. Nematodes, copepods (adults and nauplii), polychaetes (adults and juveniles) and amphipods (adults and juveniles) were the most abundant taxa found on tube-caps. Meiofaunal densities on tube-caps were 4 to 19 times higher than equivalent volumes of sediment in cores taken adjacent to D. cuprea tubes. Recruitment onto tube structure occurred within 1 to 2 d after defaunated tube-caps were replanted into sediments in the field (February–November 1980). Repopulation of copepods (adults and nauplii) attained levels equal to or exceeding natural abundances on tube caps within 1 to 5 d; nematode recovery rates were inconsistent. Short-term experiments using a variety of defaunated tube treatments indicated that immigration onto above-sediment tube-caps proceeds via both water column and sediment pathways. Based on data on tube-cap longevity and construction as well as meiofaunal recruitment rates, we conclude that the generation of new tube-cap structure is exploited rapidly by meiofauna.     

Distribution of algae on tropical rocky shores: spatial and temporal patterns of non-coralline encrusting algae in Hong Kong

Encrusting algae have been described as dominant space occupying species on rocky shores around the world. Despite their abundance, however, most studies classify species under generic names (e.g. Ralfsia sp.) or as a functional group (e.g. encrusting algae), thereby underestimating the number of species present and their ecological importance. Studies on six rocky shores of varying exposure, in Hong Kong, recorded eight common species of encrusting algae. The greatest abundance of encrusting algae was recorded on shores of intermediate exposure, where four distinct zonation bands could be identified; a cyanobacterial Kyrtuthrix-Zone in the upper midlittoral, a Bare-Zone below this, a Mixed-Zone in the lower midlittoral and a Coralline-Zone in the infralittoral fringe. Abundance declined on shores of greater and lower exposure to wave action, where bivalves and barnacles were competitively dominant. Certain species were found in greater abundance on exposed shores (e.g. Dermocarpa sp. and Hildenbrandia occidentalis), while others preferred more sheltered shores (e.g. Hildenbrandia prototypus and Kyrtuthrix maculans). With the exception of some cyanobacterial crusts, the abundance of encrusting algae was always greatest towards the low shore, an area of decreased physical stress and increased herbivore density. Zonation patterns showed seasonal variation associated with the monsoonal climate of Hong Kong. Most species increased in abundance during the cool season, while during the summer months the cover and vertical extent of encrusting algae decreased in relation to summer temperatures, although K. maculans increased in abundance during the summer. On Hong Kong shores, encrusting algae have a high species richness and exhibit within-functional group spatial and temporal variation which is mediated by herbivory and seasonal, physical stresses.     

Economic Growth and Marine Biodiversity: Influence of Human Social Structure on Decline of Marine Trophic Levels

Abstract:  We assessed the effects of economic growth, urbanization, and human population size on marine biodiversity. We used the mean trophic level (MTL) of marine catch as an indicator of marine biodiversity and conducted cross-national time-series analyses (1960–2003) of 102 nations to investigate human social influences on fish catch and trends in MTL. We constructed path models to examine direct and indirect effects relating to marine catch and MTL. Nations' MTLs declined with increased economic growth, increased urbanization, and increased population size, in part because of associated increased catch. These findings contradict the environmental Kuznets curve hypothesis, which claims that economic modernization will reduce human impact on the environment. To make informed decisions on issues of marine resource management, policy makers, nonprofit entities, and professional societies must recognize the need to include social analyses in overall conservation-research strategies. The challenge is to utilize the socioeconomic and ecological research in the service of a comprehensive marine-conservation movement.     

Mesocosm and in situ observations of the burrowing shrimp Calocaris templemani (Decapoda: Thalassinidea) and its bioturbation activities in soft sediments of the Laurentian Trough

In vivo observations in laboratory mesocosms and aquaria, accompanied with in situ photographic surveys, have shown that the burrowing shrimp Calocaris templemani has a significant impact on bottom sediment dynamics and geochemistry in the St. Lawrence Estuary. This burrowing shrimp establishes and maintains complex semi-permanent burrows made up of several interconnected, ‘U-shaped’ galleries with generally four or more openings to the sediment surface. In the Estuary, at 345 m depth, Calocaris average density was estimated at 3.4 individuals m^?2. Observed individual burrows reached a maximum volume of 0.54 L. C. templemani displaces this volume of mostly anoxic sediments from the subsurface layers (down to 15 cm) to the sediment surface, thereby obscuring some of the natural stratification patterns. With an estimated turnover rate of about 8 L m^?2 year^?1 of sediment, our calculations suggest that over a period of about 18.75 years, all the sediment to a depth of 15 cm will have been reworked by C. templemani alone.