Roles of larval supply and behavior in determining settlement of barnacles in a temperate mangrove forest

Recruitment is often a major influence on the spatial distribution of populations of benthic marine invertebrates, but the contributions of different components of recruitment are not well known, with the added complication that the relative importance of various life-history processes may be scale-dependent. Previously, we have shown that over a large scale across a mangrove (Avicennia marina) forest in southeastern Australia, settlement of the barnacle Elminius covertus explained its patterns of recruitment, which in turn explained the distribution of adults on mangrove pneumatophores. Post-settlement mortality had little influence on this pattern. In contrast, small-scale vertical distributions of adult barnacles along individual pneumatophores were determined by the pattern of recruitment, which differed from the pattern of settlement, so post-settlement mortality determined the vertical patterns of adults.

In this study, we tested whether larval supply and/or settlement behavior influence the observed settlement patterns of E. covertus across a forest (from seaward to landward zones). We also tested whether larval supply could explain the vertical settlement patterns along the pneumatophores. A pumping system was used to collect cypris larvae from seaward, mid and landward zones of a mangrove forest and an adjacent, unvegetated shore and from three heights above the sediment surface. We also used transplantation of wooden stakes bearing microbial films and barnacle recruits between horizontal zones of the forest to determine whether settlement was influenced by these films or recruits.

Both cyprid supply and cyprid behavior were important factors in determining the patterns of settlement of E. covertus across the forest. Cyprid supply was a result of three-fold differences in immersion times of different (landward, mid and seaward) zones across the forest and a decrease in density of cyprids in the water column from the seaward zone of the forest to the landward sections. In the absence of mangroves immediately adjacent to the forest, there was no temporally consistent difference in cyprid density across the shore and even the differences in immersion time did not produce consistent differences in cyprid supply across the shore. Wooden substrata that had been immersed at seaward sections of the forest attracted consistently more settlers than substrata immersed initially at other sections of the forest and settlement could be induced beyond the normal distribution of adults of E. covertus by stakes transplanted from the seaward zone.

The vertical settlement pattern could not be explained by the supply of cyprids, suggesting that larval behavior must determine the vertical settlement pattern.     

Juvenile morphology and occurrence patterns of three Butis species (Gobioidei: Eleotridae) in a mangrove estuary, southern Thailand

Juveniles of three eleotrid Butis species (B. butis, B. humeralis, and B. koilomatodon) are described; their occurrence patterns were examined in Sikao Creek, a mangrove estuary located in southern Thailand. Juveniles of each species were distinguished by the following characters: B. butis with no bands on body and pale pelvic fins; B. humeralis with no bands on body and densely pigmented pelvic fins; and B. koilomatodon with 5–6 regular bands on body and a fleshy process (preorbital knob) on the snout. Although B. butis shared the aforementioned characters with B. amboinensis found in the same estuary, the former was distinguished from the latter by having a greater number of pectoral fin rays (18–21 vs. 17) and a deeper caudal peduncle. Distribution patterns of the three Butis species in Sikao Creek were distinguishable from each other. Smaller B. butis [mean ± SD = 22.7 ± 16.9 mm in standard length (SL), n = 32] occurred in the upper reach of the estuary, while larger specimens (52.4 ± 26.2 mm SL, n = 18 and 51.5 ± 29.7 mm SL, n = 10, respectively) were found in the middle and lower reaches and none in the marine area. In B. humeralis and B. koilomatodon, only juveniles were caught except for one adult specimen each. Juveniles (8.9–16.5 mm SL, n = 79) of B. humeralis occurred in the upper and middle reaches and the marine area. B. koilomatodon juveniles (9.9–13.7 mm SL, n = 30) were distributed in all areas from the lower to upper reaches.     

The importance of crevices to the intertidal snail Littoraria articulata (Philippi) in a tropical mangrove forest

The intertidal gastropod, Littoraria articulata (Philippi), is common on the trunks of mangroves at Coorooman Creek, Central Queensland. Individuals of shell length less than 3 mm were only found in empty tests of the barnacle Hexaminius popeiana (Foster). Larger snails were exposed on the trunk or occasionally in crevices such as knot holes. The relationship between shell length and shell strength differed between barnacle dwelling and larger L. articulata: in both cases it was linear, but the rate of increase of strength with increasing length was significantly greater for barnacle dwelling snails than for exposed ones. In contrast, there was an exponential relationship between shell strength and length for the weaker shelled Littoraria filosa (Sowerby), which occurs higher on the tree and does not inhabit barnacles. The survival of L. articulata was affected by the presence of crevices: significantly fewer disappeared from posts with artificial crivices than from those which lacked them, and some snails testhered to posts appeared to have been eaten by fish. The importance of crevices is discussed in relation to predation pressure, growth and habitat selection by larval L. articulata. This revised version was published online in July 2006 with corrections to the Cover Date.     

Sporulation ofEnteromorpha spp. (Chlorophyta) and overwintering of spores in sediments of the Wadden Sea,Island Sylt,North Sea

For the last two decades dense mats of species of the filamentous green algaeEnteromorpha spp. have regulary occurred on tidal flats of Köningshafen Bay (island Sylt, North Sea, FRG). In calm areas overwintering of adult plants or plant fragments is a common process to guarantee the mass development during the next season. In contrast, the distribution ofEnteromorpha on exposed sandy tidal flats depends on recruitment by juvenile stages. In 1993Enteromorpha spore settlement was recorded regularly in the field. Polyethylene dishes were placed in the field and left for a period of seven days and lateron cultivated in the laboratory to checkEnteromorpha germling development. During summer 1993 — at a minimum distance of 200 m to the nearest adultEnteromorpha populations — a total of at least 82×10⁶ spores m^–2 settled. During winter the number of spores attached to the collecting dishes was close to zero and the adjacent sand flats were free of any visibleEnteromorpha plants. In further experiments it was shown that the development ofEnteromorpha juveniles in the next spring depended on the overwintering capacity of spores. More than 2×10⁶ spores m^–2 attached to large sand grains and other substrata (e.g. Hydrobia ulvae) survived the winter. In a laboratory experiment several species ofEnteromorpha were able to survive in total darkness for at least 10 months.     

Zonation of ground beetles (Coleoptera: Carabidae) and spiders (Araneida) in salt marshes at the North and the Baltic Sea and the impact of the predicted sea level increase

The ground beetles and spiders of two salt marshes at the German Northand Baltic Sea coast were investigated by pitfall traps in 1997 and 1998. Whilethe sites at the North Sea coast are tidal salt marshes, the salt marshes at theBaltic Sea are not influenced by tides. Pitfall traps were installed in agradient from 20 to 150 cm above MHT (mean high tide: 157cm + NN, NN: 500 cm above 0 at Amsterdam gauge) atthe North Sea coast or NN at the Baltic Sea coast at six or sevensampling elevations, each with five replicates. Conductivity, water content, organic substance,frequency or duration of floodings, sand content and pH of the soil weredetermined. The flooding regime is the major factor controlling the zonation ofinvertebrates. Two and three invertebrate assemblages at the North and BalticSea, respectively, were distinguished. These corresponded well with thevegetational zones. The border between the two zones was at 60–80cm above MHT at the North Sea. The three zones at the Baltic Seaextended between 20 and 30 cm, 40 to 80 cm and 100 to150 cm above NN. The elevation of the mean abundance of speciesabove MHT or NN was calculated. A tide simulation experiment resulted in ashifting population and in an increasing activity under a tidal regime aspredicted for the global climate change conditions in 2050. From the actualelevation of the mean abundance, the habitat size of salt marsh species wascalculated for a moderate and worse scenario of global climate change. Habitatreduction becomes highest for species of the lower salt marsh zone. Under worseconditions the gradiental length of habitat will only amount to a maximum of 20m at the slopes of the dikes.     

Activity and growth efficiency of heterotrophic bacteria in a salt marsh (Ria de Aveiro, Portugal)

Bacterial utilization of monomers is recognized as an important step in the biogeochemical cycling of organic matter. In this study we have compared the heterotrophic activity of bacterial communities from different micro-habitats within a salt marsh environment (Ria de Aveiro, Portugal) in order to establish spatial patterns of bacterial abundance, monomer turnover rates (Tr) and bacterial growth efficiency (BGE). Differences in bacterial abundance and activity could be found between distinct plant rhizospheres. BGE tended to be lower at Halimione portulacoides banks, when compared to Sarcocornia perennis subsp. perennis banks which, on the contrary, showed the highest bacterial densities. Experiments of amendment of natural samples with organic and inorganic supplements indicated that salt marsh bacteria are not strongly regulated by salinity but the increased availability of labile organic matter causes a significant metabolic shift towards mineralization.     

Activity patterns, feeding and burrowing behaviour of the crab Ucides cordatus (Ucididae) in a high intertidal mangrove forest in North Brazil

Activity patterns, feeding and burrowing behaviour of the economically important semi-terrestrial mangrove crab Ucides cordatus (Ucididae, L. 1763) was studied in a high intertidal Rhizophora mangle forest stand in Bragança, North Brazil. Video observations in the rainy and dry season were conducted over 24 h cycles at different lunar phases to investigate the behaviour of these litter-feeding crabs outside their burrows. During the rainy season, crabs stayed inside their burrows for 79% and 92% of the time during day and night, respectively. Time spent for feeding, burrowing and other activities outside their burrows was significantly longer during the day with 9.9% (night: 1.7%) and at waning and waxing moon with 9% (full and new moon: 0.9%). At neap tides (no tidal inundation) foraging and feeding activities outside burrows were clearly light-dependent, increasing at dawn and decreasing at dusk. Highest activities during daytime relate to the visual localisation of food. During the dry season, crabs spent less time inside burrows at neap tides than during the rainy season (80% and 91%, respectively). However, time spent for feeding activities was similar during both seasons. During almost all observation periods crabs collected leaf litter, but rarely fed on it outside burrows. At neap tides nearly all available litter was collected, suggesting that the U. cordatus population is litter-limited during these times. At spring tides (regular tidal inundation) the surface activity of U. cordatus was tide-dependent. Crabs closed their burrow entrances 2-3 h before flooding and re-emerged as soon as the tide retreated. During the day, burrow maintenance was the second most frequent behaviour after feeding. Agonistic interactions were regularly observed and were mainly related to burrow defence. The mean foraging radius of the crabs was only 19 cm (max: 1 m) underneath high Rhizophora mangle trees where crab densities were high. The results point to a high competition for burrows and show that U. cordatus is territorial. It is concluded that several exogenous factors, in particular light, leaf litter availability, flooding of burrows and the presence of conspecifics are important in controlling the crabs' activity patterns.     

Characterisation and analysis of microsatellite loci in a mangrove species, Avicennia marina (Forsk.) Vierh. (Avicenniaceae)

An enriched microsatellite library of the mangrove species Avicennia marina was constructed, in which 85.8% of the clones contained microsatellite sequences. Of the microsatellite repeat sequences isolated, 55.0% were di-nucleotides, 34.2% were tri-nucleotides, 50.0% were perfect, 24.2% were imperfect, and 15.0% were compound. Four different di-nucleotide repeats were isolated with repeat lengths ranging from 5 to 33; ten different tri-nucleotide repeats were isolated with repeat lengths ranging from 3 to 25. The most common di-nucleotide was the AC/TG repeat; the most common tri-nucleotide was the CCG/GGC repeat. Sixteen microsatellite sequences were selected for primer design, and 6 primers were selected to investigate the polymorphism detected among 15 individuals of A. marina from three natural populations in Australia. A total of 40 alleles were detected at 6 microsatellite loci. The number of alleles per microsatellite locus ranged from 5 to 13. On average, 7 alleles were detected per locus. All microsatellite loci showed high levels of gene diversity (heterozygosity), with values ranging from 0.53 to 0.88; the mean value of gene diversity was 0.70. Microsatellite loci were also tested for conservation across Avicennia species. There was a decline in amplification success with increasing divergence between Avicennia species. The results indicate that microsatellites are abundant in the Avicennia genome and can be valuable genetic markers for assessing the effects of deforestation and forest fragmentation in mangrove communities, which is an important issue for mangrove conservation and afforestation schemes. Received: 8 June 1999 / Accepted: 21 September 1999     

Impact of sampling methods on sulfate reduction rates and dissolved organic carbon (DOC) concentrations in vegetated salt marsh sediments

The impact of sediment coring on measured rates of sulfate reduction(SRR) by the whole core ³⁵S-injection technique was assessed inmarshsediment vegetated by Spartina anglica. Simultaneously,therole of extraction method (centrifugation vs. sippers) for determination ofporewater DOC in vegetated sediment was evaluated. SRR was measuredinsitu with radiotracer injected directly into the sediment and in atime series from 1 to 24 h after coring. SRR incubations carriedout within 6 h (June) or 12 h (August) of coringyielded up to an order of magnitude higher rates than measured insitu. The enhancement of SRR was instantaneous but temporary andcorrelated with measured porewater DOC concentrations. Cores sampled fromrootedsediments should therefore not be used for sulfate reduction incubations withinthe first 12 h due to the effect of DOC leaching from roots cutduring the coring procedure. The labile fraction of leached DOC appears to beexhausted after a pre-incubation period of at least 12 h.Measurement of porewater DOC is also problematic in vegetated sediment.Porewater extraction by centrifugation of sediment may result in up to oneorderof magnitude higher DOC concentrations than in porewater obtained by anondestructive sipper technique. DOC is probably forced out of roots duringcentrifugation resulting in erroneously high porewater DOC concentrations.     

The diet of the mud clam Geloina coaxans (Mollusca, Bivalvia) as indicated by fatty acid markers in a subtropical mangrove forest of Okinawa, Japan

Fatty acid compositions in the tissues of the clam Geloina coaxans collected from Oura mangal, Okinawa, Japan, during the cold and warm seasons (January and July 2001, respectively) were compared with those in suspended materials (SM) in order to assess the clams' diet. In both seasons, the suspended mangrove detritus at the sediment-water interface was high as indicated by the mean percentage of even-numbered long-chain fatty acids in SM (12.8-18.4%). The contribution of this marker in the clam tissues, especially during the cold season (3.9%), indicates the consumption of mangrove detritus in considerable amounts by the clams. The occurrence of the fatty acids 16:1ω7, 18:1ω9, 18:2ω6 and 18:3ω3 in SM was most likely due to the mangrove detritus sources, whereas in the SM they together constituted 12.9% and 23.9% of total fatty acid contents during the cold and warm seasons, respectively. As a result, their contribution in the clam tissues was high in the cold (15.4%) and warm seasons (19.0%). These results indicate that mangrove detritus play a significant role in the clams' diet. The mean percentages of bacterial markers (odd-numbered branched fatty acids and vaccenic acid, 18:1ω7) in the SM and tissues during both seasons ranged from 8.1% to 9.5%. This indicates that the clam diet is also dependent on the attached bacteria on the partially decomposed leaf detritus suspended at the sediment-water interface. The relative contribution by microalgae markers (18:4ω3, 20:5ω3 and 22:6ω3) in clam tissues ranged from 4.3% to 7.6%, suggesting considerable microalgae sources in the diets.     

Long-term changes of intertidal and subtidal sediment compositions in a tidal basin in the northern Wadden Sea (SE North Sea)

Wadden Sea tidal flats are highly dynamic regarding the spatial distribution and the grain size composition of their sediments. From 2003 to 2006 surface sediments have been surveyed in an intertidal and a subtidal area within the tidal inlet Königshafen (south-eastern North Sea, northern Wadden Sea, island of Sylt) with the goal to gain information on short-term development trends in the grain size composition. The average grain size (Mean) becomes finer in the sheltered part of the intertidal survey area whereas a coarsening tendency can be observed in the more exposed part of the intertidal and especially in the subtidal survey area. The trend of the most frequent grain size (first Mode) shows the same spatial distribution pattern but is far less distinct. Thus, the changing Mean must be related to an increase in the deposition of fines in the sheltered part of the intertidal Königshafen as well as a general removal of fine-grained material in the exposed intertidal and subtidal Königshafen. In order to see long-term trends the surveys of 2003–2006 were compared to earlier studies conducted in 1932/1933, 1981 and 1989. A significant depletion of mud can be observed in the entire survey area. It is concluded that primarily changed hydrodynamics that may accompany ongoing climate change are responsible for this. However, the loss of fine-grained sediments is additionally amplified by a reduced vegetation cover and coastal protection measures.     

Recognition of facial expressions in a Japanese monkey (Macaca fuscata) and humans (Homo sapiens)

Recognition of facial expressions by a Japanese monkey and two humans was studied. The monkey subject matched 20 photographs of monkey facial expressions and 20 photographs of human facial expressions. Humans sorted the same pictures. Matching accuracy by the monkey was about 80% correct for both human and monkey facial expressions. The confusion matrices of those facial expressions were analyzed by a multi-dimensional scaling procedure (MDSCAL). The resulting MDS plots suggested that the important cues in recognizing facial expressions of monkeys were “thrusting the mouth” and ‘raising the eyebrows.” Comparison of the MDS plots by the monkey subject with those by human subjects suggested that the monkey categorized the human “happiness” faces. This may suggest that the monkey has an ability to recognize human smile face even though it is learned. However, the monkey did not differentiate the human “anger/disgust” faces from the human “sad” faces, while human subjects clearly did. This may correlate with the lack of eyebrow movement in monkeys.     

Response of a raptor community to shrinking area and degradation of tropical rain forest in the south western Ghâts (India)

The influence of irregular or incomplete fragmentation and increasing degradation of natural rain forest on diurnal raptor community was studied in the northern part of the western Ghâts in southwestern India A census of mainly territorial breeding pairs on 400 ha sample quadrats was associated with a measure of the percent cover of the main habitat types and a degree of forest fragmentation including irregular patch shape and habitat heterogeneity
Four groups of 3-4 species were defined according to their decreasing tolerance to forest fragmentation and disturbance, from mostly open grassland species to interior forest specialists Habitat selection, density and sensitivity to landscape structure were investigated The community composition and dynamic of each habitat were the sum of these specific reactions The distribution of species along the succession of increasing fragmentation and forest degradation was thus found to be non random, but did not follow a nested subset pattern The density of forest species declined with forest patch size possibly because of the irregular patch shape, the increased edge effects and the consequent increase of linear distance for a bird to cover within its territory However, sensitivity to habitat structure and disturbance was found to be even higher than sensitivity to area per se The need to conserve the largest patches of little disturbed forest is emphasized, as well as the conservation value of woodlots-open habitat mosaics that are suitable for a different set of species     

Spatial and temporal distribution of Myrophis punctatus (Ophichthidae) and associated fish fauna in a northern Brazilian intertidal mangrove forest

The mangrove forest along the northern Brazilian coast is not inundated during low tide. However, many fish species stay in the mangrove forest during this time. Tidal behaviour strategies are described for fish species that linger in the mangrove forest during low tide. The samples were taken at the end of the dry season (December 1996) and at the end of the rainy season (July 1997). Fish were captured using an ichthyotoxic plant extract (Ichthyotere cunabi). Spatial and temporal fish density and biomass were analyzed statistically. Thirty-six samples were taken with a total density of 2.8 ind m^-2 and a total biomass of 17.4 g m^-2 distributed among seven families and 14 species. Myrophis punctatus was the most important species in number (1.66 ind m^-2) and weight (12.68 g m^-2) of all catches. The total fish densities were not significantly different among areas and between months, although, total biomass differed significantly in time and space. The densities and biomass for the three most dominant species (M. punctatus, Poecilia spp. and Gobionellus smaragdus) differed significantly among species. Only the biomass of these species showed significant monthly differences. The only significant main effect on variance in the densities and biomass of M. punctatus were encountered between months. In addition, the factor area was significantly different for the variable number of species.     

Benthic oxygen respiration, ammonium and phosphorus regeneration in surficial sediments of the Sacca di Goro (Northern Italy) and two French coastal lagoons: a comparative study

During 1994 net sediment-water fluxes of oxygen, ammonium and inorganic phosphorus as well as sediment profiles of organic matter, nitrogen, phosphorus and iron were determined in three shallow eutrophic environments. Investigations were conducted monthly from March to December at five stations in the Sacca di Goro (Po River Delta, Italy). In the late summer, samples were collected from a single site in the Prévost lagoon (French Mediterranean coast) and three stations in the Bassin d'Arcachon (French Atlantic coast). In the Sacca di Goro, water-sediment exchanges of O₂, NH ₄ ⁺ and PO ₄ ^3– were estimated by means of core incubation in the dark. Benthic fluxes for the French lagoons were in part determined experimentally using benthic chambers and in part from the literature.In general in the Sacca di Goro the highest oxygen uptake and nutrient release rates were found at the central sites, affected by macroalgal growth. At the sampling site adjacent to the freshwater inlet, sediment-water exchanges were principally influenced by tidal activity. In terms of organic matter and nutrient levels, sediments from the Sacca di Goro and from the Prévost lagoon, both colonised by the floating macroalga Ulva rigida C. Agardh, were similar. Sediments from the inner sheltered site in the Bassin d'Arcachon, invaded by the rooted macrophyte Ruppia cirrhosa (Pet.) Grande, showed the highest total N and P content (363 ± 157 µmol N cm^–3 and 15 ± 2 µmol P cm^–3 as average values in the top 10 cm of sediment), but were low in pore water ammonium and orthophosphate probably due to the high sequestering capacity of the system and/or efficient coupling between bacterial nutrient regeneration and assimilation by the plant roots. In addition the outer tidal stations in the Bassin d'Arcachon, invaded by rooted macrophytes, were low in pore water nutrients. A different trend was evident in the Prévost lagoon where the concentrations of exchangeable inorganic phosphorus and ammonium were appreciable (0.28 ± 0.07 µmol P cm^–3 and 2.4 ± 1.4 µmol N cm^–3 as average values in the top 10 cm of sediment). High amounts of dissolved organic nitrogen were found in the pore water at all the sites investigated showing the key role of the organic nitrogen in the recycling of nitrogen in these systems.The hypothesis that iron is a key factor in controlling phosphorus release is discussed since the Sacca di Goro, which is subject to dystrophic crises, is richer in iron than the Bassin d'Arcachon, which is a more buffered system.     

Towards a physical description of habitat: quantifying environmental adversity (abiotic stress) in temperate forest and woodland ecosystems

1. Our aim was to develop a quantitative proxy for environmental adversity (abiotic stress) in temperate Eucalyptus and Nothofagus forest and woodland ecosystems.
2. Samples and measurements were collected at 42 sites across a rainfall gradient in southern Australia, an elevation gradient in south-eastern Australia, and a longitudinal transect (temperature gradient) in Patagonia, Argentina.
3. We compared the ability of (a) abiotic variables (14 soil and 21 climatic variables) and (b) the stable carbon isotope (δ¹³C) values of soil organic matter (SOM), to predict variation in leaf area index (LAI; a forest productivity variable).
4. The δ¹³C of SOM (soil aggregates) explained more variation (57%) in LAI than multivariate statistical models that integrated information on many abiotic variables. W* (a climatic water balance model) was also a powerful predictor variable, explaining 37% of the variability in LAI.
5 Synthesis . The stable carbon isotopic signature of soil aggregates is a powerful explanatory variable that may help us to quantify environmental adversity (abiotic stress) in temperate forest and woodland ecosystems.     

Natural and man-induced changes in a tidal channel mangrove system under tropical semiarid climate at the entrance of the Maracaibo lake (Western Venezuela)

Caño Paijana is a tidal channel that connected, until recently, theUruba bay at the mouth of the Limón river, to the Gulf of Venezuelanorth of Maracaibo, Venezuela. It separated the Island of San Carlos fromthe mainland. In the last 50 years the channel has been drying out rapidly,to the extent that presently there is no water flow into or from the seathrough its mouth opening on the Gulf of Venezuela. This mouth wascovered by healthy mangroves (mainly Avicennia germinans) until atleast 1952. The two extremes of the Paijana water channel differ radicallytoday. At Uruba bay end the channel is fringed by dense mangrovevegetation dominated by Rhizophora mangle, while at the Gulf ofVenezuela mouth the vegetation cover is mostly constituted by scrubby,scattered trees of Avicennia germinans, xerophytic shrubs acting asdune fixers, and halophytic strand vegetation. The process is the result ofcomplex interaction between: a) high frequency of low-rainfall years; b)high dune activity during dry years c) dam construction on tributaries ofthe Limón river that reduced discharge of fresh water into Uruba bay.