Habitat and zonation patterns of Littoraria species (Gastropoda: Littorinidae) in Indo-Pacific mangrove forests

The zonation of 15 species of Littoraria in the scabra group is described, using a transect method, at 13 mangrove localities in Australia, Thailand, Malaysia and Hawaii. At these localities the littorinids occurred only on trees, and showed both vertical and horizontal patterns of zonation, with broad overlaps between co-occurring species. It is suggested that vertical distribution is determined mainly by behavioural responses, while the landward limits of horizontal distribution may be set by physiological tolerance. Several species occurred predominantly on either leaf or bark substrates, but no consistent associations with tree species were found. On the basis of local distribution patterns, Littoraria species can be classified as continental or oceanic in character. Oceanic species may be less tolerant of sediment in the water or on the substrate.     

Natural selection for apostasy and crypsis acting on the shell colour polymorphism of a mangrove snail, Littoraria filosa (Sowerby) (Gastropoda: Littorinidae)

Littoraria filosa (Sowerby) is a member of the L. scabra group, found amongst the foliage of mangrove trees in northern Australia. The colour of the shell is polymorphic, showing two discrete ground colours, either yellow or orange-pink, with a variable degree of superimposed brown patterning. At a site on Magnetic Island, northern Queensland, colour frequencies of small snails were similar on different backgrounds. Amongst larger shells yellows were more frequent on Avicennia trees with abundant foliage, and browns on relatively bare trees, suggesting that visual selection for crypsis occurred. There was no evidence of substrate selection by the morphs. Yellow shells were cooler than brown shells, but differences in colour frequencies on sunny and shaded trees, and at different seasons, did not suggest climatic selection. By manipulating the colour frequencies of subpopulations of small snails isolated on individual trees, it was shown that the disappearance of yellow and brown shells was frequency-dependent. This result is consistent with hypotheses of mimicry of background elements by the morphs and of apostatic selection by unknown predators. Only the latter can account for the persistence of the highly conspicuous pink morph at a low frequency.     

The effects of two parasitoids on the life history and metapopulation structure of the intertidal snail Littoraria filosa in different-sized patches of mangrove forest

We describe differences in life history of the intertidal arboreal snail, Littoraria filosa, among patches of mangroves ranging in size from isolated trees to large stands several square kilometres in area. Recruitment of L. filosa occurred from mid spring (October) to early winter (June), recruits grew rapidly and copulating adults were found during the following September–April. Populations within large patches of forest were annuals; all or most individuals died between October–January (spring–midsummer). In contrast, those in smaller peripheral patches were more likely to survive over the summer but survival differed among patches and years. These differences in life history were caused by a parasitoid fly (genus Sarcophaga) that attacked L. filosa 10 mm and longer and was present in all large patches, but absent from, or rare, in smaller peripheral patches. Experimental introductions to isolated trees confirmed that the fly could kill L. filosa. Another sarcophagid parasitoid that attacked L. filosa from 4 to less than 10 mm long was also found in every patch. The combined effects of these parasitoids appear to determine the metapopulation structure of L. filosa. Most adults in large patches were killed by the larger fly during early summer. Summer recruits were often killed by the smaller fly within a month of settlement and when this happened effective recruitment of L. filosa was reduced to autumn. The planktotrophic larval stage of L. filosa lasts less than 1 month, so the source of autumn recruits to all patches must have been adults that survived the early summer, most of which were in small patches or on isolated trees. Consequently these ”peripheral sources” are likely to be important for persistence of the metapopulation of L. filosa. The results of this study demonstrate that metapopulation structure may be determined by complex interactions and that common models cannot be assumed to apply in all habitats. Received: 15 September 1999 / Accepted: 31 January 2000     

Temporal patterns in a fish assemblage of a semiarid mangrove zone in Madagascar

Gillnet sampling was conducted for a year in a tropical mangrove creek (SW Madagascar), characterized by a limited freshwater influence, a high turbidity and a tidal range up to 3 m. Sixty species of juvenile fishes were caught, 44 species being of commercial interest. Catches were dominated by Gerreidae (27% of total abundance), Teraponidae (16%), Carangidae (13%) and Sparidae (12%). The temporary resident fishes in the mangrove zone represented 50% of the species and 97% of the total abundance, the other species being rare (less than five individuals). The species richness, abundance and biomass per netting were low in the middle of the cool season (July-August). Monthly changes in the fish assemblage were particularly complex, with three species groups displaying a clear seasonal pattern, some species succeeding one another in a rather unstructured way, and three species abundant throughout the year. There was no clear structuring effect of temperature, salinity and turbidity on the fish assemblage. However, tidal, lunar and diel effects on the composition of the fish assemblage were evident. The species overlap between the Sarodrano mangrove fauna and the adjacent coral reef fauna is particularly weak with six species in common and shows that the mangrove plays only a very limited nursery role for coral reef 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.     

Characterization of phosphorus fractions in the sediments of a tropical intertidal mangrove ecosystem

Solid phases of phosphorus fractions in the surface and core sediments were studied to understand the biogeochemical cycling and bioavailability of phosphorus in the Pichavaram intertidal mangrove sediments of India. Total P in surface and core sediments ranged between 451–552 and 459–736 μg g⁻¹ respectively and Fe bound P was the dominant fraction. Low levels of Fe bound P in the mangrove zone than the two estuarine zones may be because of high salinity inhibition of phosphate adsorption onto the Fe-oxides/hydroxides. Post-depositional reorganization of P was observed in surface sediments, converting organic P and Fe bound P into the authigenic P. High levels of organic P in the mangrove zone is primarily due to intensive cycling and degradation of organic matter and adsorption of phosphate on the organic molecules. The burial rates and regeneration efficiency of P in the intertidal mangrove ecosystem ranged from 5.41 to 7.27 μmol P cm⁻² year⁻¹ and 0.122 to 0.233 μmol P cm⁻² year⁻¹, respectively. High burial efficiency (≈99%) of P proves the earlier observation of limiting nature of P for the biological productivity. Further, bioavailable P (exchangeable P + Fe bound P + organic P) constituted a considerable proportion of sedimentary P pool of which an average accounted for 55 and 50% in surface and core sediments respectively. The results indicate that significant amount of P is locked in sediments in the form of authigenic P and detrital P which makes P as a limiting nutrient for the biological productivity.     

Euspermatozoa,Paraspermatozoa and Spermatozeugmata of Littoraria (Palustorina) articulata (Prosobranchia: Caenogastropoda) with Special Reference to the Pseudotrich

Abstract Paraspermatozoa and euspermatozoa of the littorinid gastropod Littoraria (Palustorina) articulata are examined using transmission electron microscopy and light microscopy. In the seminal vesicle, both sperm types occur, either as free cells or organized into spermatozeugmata. It is shown that the elongate (120–140 μm), flagellum-like component of the paraspermatozoon is in fact a tubular extension of the plasma membrane which encloses granular material but no axonemes or microtubules. This structure, here termed the pseudotrich, shows no evidence of motility and its function remains obscure. The main body region of the paraspermatozoon (length 32–36 μm) contains numerous spherical vesicles, scattered mitochondria, one or two large, rod-shaped bodies (length 20–24 μm) and a fusiform, granular body (containing DNA; probably a modified nucleus). The rod-shaped bodies, granular body and surrounding matrix are contained by a common membrane, and are therefore separated from other contents of the paraspermatozoon. In each spermatozeugma, euspermatozoa are attached via the tips of their acrosomes to the paraspermatozoan body at the opposite end to the pseudotrich. Euspermatozoa exhibit a conical acrosomal complex (with axial rod and basal plate), a tubular nucleus sheathing the axoneme, a midpiece (5–6 helical mitochondrial elements sheathing the axoneme), an annulus (with two rings), a glycogen piece and an end piece (total sperm length 268–272 μm). The euspermatozoa of L. articulata are similar to those of most littorinids and many other caenogastropods. The presence of a pseudotrich in the paraspermatozoon appears to be restricted to the subgenus Palustorina.     

The spatial and vertical distribution of living (stained) benthic foraminifera from a tropical,intertidal environment,north Queensland,Australia

We investigated the distribution of living (stained) benthic foraminifera across a tropical, intertidal shoreline adjacent to Cocoa Creek, Queensland, Australia for the purpose of better understanding the nature of test production and ultimately fossil assemblage development within such environments. Short cores (up to 1 m) were collected during the wet and dry season, along an elevational gradient comprising non-vegetated intertidal mudflat and higher-intertidal mangrove forest environments. The distribution of stained specimens can be broadly delineated into assemblages characterising ‘upper mangrove’ (2.64–2.91 m above Lowest Astronomical Tide (LAT)) and ‘low mangrove-mudflat’ (1.62–2.18 m above LAT) environments. Agglutinated species were generally limited to upper mangrove stations. Calcareous species occurred within all of the intertidal environments examined but differ in their composition between upper and lower intertidal settings. Upper mangrove faunas were characterised by the agglutinated species Arenoparrella mexicana, Haplophragmoides wilberti, Miliammina fusca, Miliammina obliqua and Trochammina inflata and the calcareous species Helenina anderseni. Live (stained) assemblages at lower intertidal elevations were dominated by the calcareous species Ammonia aoteana, as well as Rosalina spp., Elphidium oceanicum, Triloculina oblonga, Ammonia pustulosa and Shackoinella globosa.     

Meiobenthos of hypersaline tropical mangrove sediment in relation to spring tide inundation

Tropical intertidal sediments often contain porewater of relatively high salinity, especially in areas exposed to longer periods without seawater inundation and high evaporation. Such an area exists on the west coast of Zanzibar: a high intertidal mangrove plateau, flooded only during spring high tides, with sediment porewater salinities commonly exceeding 100 ppt. A field survey was conducted in this area to examine variations in population density of major meiofaunal taxa and the assemblage structure of free-living marine nematodes during spring-neap tidal cycles. Samples were taken on seven occasions for two months, starting from the end of the rainy season. Porewater salinity remained high throughout the sampling period, ranging from 89 to 160 ppt. Neither spring tide inundation nor heavy rains lowered the salinity markedly. The meiofauna consisted only of four taxa, present on all sampling occasions: nematodes, harpacticoid copepods, plathyhelminthes and chironomids. Densities in surface sediments (0–5 cm) were low compared to other mangrove areas, ranging from 271 to 656 animals 10 cm^-2 with nematodes dominant on all sampling occasions (58–87%). Density fluctuations could not be explained by the effects of spring tide inundation, but the meiofauna showed significant correlations with grain size and organic material. Despite the wide range of salinity, only the numbers of chironomids were negatively correlated with increased salinity. Nematode species diversity was low in all samples, although altogether 28 species were recorded in the samples. Four species occurred in more than 50% of the samples (Microlaimus sp. (100%), Metalinhomoeus sp. (76%), Daptonema sp.l (56%), Chromadorina sp. (56%)) while 12 species were found only in one or two samples. Multidimensional scaling ordination (MDS) of the nematode species abundance data indicated little effects of spring tide inundation on the assemblage structure, but rather a successive change from wet to dry season with a reduction in species diversity and increased numbers of the dominant nematode species Microlaimus sp.     

Predation induced changes in behavior and growth rate in three populations of the intertidal snail, Littorina sitkana (Philippi)

We investigated the sublethal effects of a predatory crab, Cancer productus (Randall), on the behavior and growth of its snail prey, Littorina sitkana, by setting up controlled rearing and prey-size selection experiments. L. sitkana were collected from three sites on San Juan Island, WA, USA. These sites varied in snail size, abundance, and vertical distribution, and in the abundance of the crab predator C. productus. Snails from all three populations were raised for 34 days under the following treatments: no-crab control, a non-feeding C. productus encased in mesh box, and an encased C. productus feeding on L. sitkana. The non-feeding crab treatment did not affect snail foraging behavior or growth rate in comparison with the no-crab control. In contrast, the presence of a feeding crab elicited escape behavior in the snails, halted grazing, and consequently reduced growth rates. A population difference in escape behavior was observed: upward migration in snails from rocky shores and hiding in crevices in snails from a mud flat. It thus appears that chemicals leaching from crushed conspecific snails, rather than the presence of the crab predator, act as the “alarm substance” to which L. sitkana react. The magnitude of the growth depression in the presence of feeding crabs was 85%, with no difference among the three populations. Once the feeding crab stimulus was removed, snails in all populations resumed normal growth, suggesting that this response to feeding predators is reversible with changing environmental conditions. Laboratory experiments were set up to determine if all size classes of L. sitkana are equally susceptible to C. productus predation. C. productus consistently selected the largest of three size classes of L. sitkana. These results suggest that slow growth rate and small size in L. sitkana may actually be an adaptation for coexisting with high C. productus abundance, rather than simply a cost of escape behavior.     

Reproduction and feeding ethology of a tropical,intertidal sand-dwelling anemone (Actinoporus elongatus,Carlgren, 1900)

Actinoporus elongatus is a widely distributed, tropical, intertidal anemone from the Indo-Pacific and Caribbean regions. Its recorded occurrences are disjunct and its feeding and reproductive ecology have not previously been described in detail. A population of this sand-dwelling species was studied for one year at Shelly Bay, Queensland, Australia. Individuals were found to be randomly distributed within the study site at a density of 42 anemones per hectare. No immigration, juvenile recruitment or mortality was observed. Histological analysis indicated a protracted period of gamete maturity during the summer months, and that the sex ratio was 1:1. A. elongatus is an opportunistic feeder with two distinct feeding methods — actinopharyngeal eversion and a newly described trapdoor response. Ciliary currents on the tentacular crown and oral disk were used mainly for cleaning and sand removal.Department of Zoology, James Cook University of North QueenslandDepartment of Marine Biology, James Cook University of North QueenslandDepartment of Marine Biology, James Cook University of North Queensland     

Ammonium adsorption in sediments of a tropical mangrove forest (Thailand) and a temperate Wadden Sea area (Denmark)

Dimensionless ammonium adsorption coefficients (K) were determined for tropical mangrove forest sediments and temperate Wadden Sea sediments. The K values were obtained from ammonium adsorption isotherms of KCl (2 M) extractable ammonium versus dissolved ammonium in the porewater; relationships that can be described by a linear model. Linearity was evident even at low porewater concentrations for mangrove sediment, according to isotherms based on KCl extractions on untreated sediment profiles. K-values were low in mangrove forest sediments (0.04–0.10), and higher in Wadden Sea sediments (0.17–1.12). The vertical range in K-values was larger at the vegetated sites, with highest values in subsurface sediments, which suggests differences in quantity and quality of the fine organic and inorganic fractions. The low ammonium adsorption in the mangrove sediments can be explained by a higher concentration of competitive cations, especially iron, in this iron-rich environment. The low adsorption of ammonium in mangrove sediments and vegetated surface sediment of the Wadden Sea was negatively related to the organic content of the sediments, which is in contrast to findings for other marine sediments. We suggest that organic material may have a diluting effect on the exchange capacity in fine-grained sediments, and that organic coatings may block ion exchange sites on clay surfaces. Thus, there may be a organic-rich ‘wetland’ versus organic-poor ‘sediment’ effect. This revised version was published online in July 2006 with corrections to the Cover Date.     

The role of tropical dry forest as a long-term barrier to dispersal: a comparative phylogeographical analysis of dry forest tolerant and intolerant frogs

We used a comparative phylogeographical approach to investigate the origins of the disjunct wet forest biota of the Golfo Dulce region along the Pacific slope of Costa Rica. This region is isolated by Pacific dry forests north and south and isolated from Caribbean wet forests by mountains. We studied three sympatric lowland frog species in the Craugastor fitzingeri species group that prefer wet forest but differ in their response to dry habitats. In dry forest, C. fitzingeri can survive along streams while C. crassidigitus and C. talamancae are entirely absent. We collected samples from across the ranges of all three species, and obtained mitochondrial DNA sequence data from the COI and cytochrome b genes. We observed significant phylogeographical structure in C. crassidigitus and C. talamancae, but much less in C. fitzingeri, demonstrating that mountain barriers and dry forest habitat have reduced mitochondrial gene flow in the strictly wet-forest species. Additionally, we discovered that the Golfo Dulce and Central Panama populations of C. crassidigitus appear to have diverged in the Pliocene or earlier, suggesting that the dry forest separating these populations is old. Our phylogenetic analysis of 12 of approximately 16 species of the C. fitzingeri species group suggests that the three lowland species are each other's closest relatives. Because of this shared phylogenetic history, we attribute the striking differences in phylogeographical structure to the different ecologies of the frogs. In summary, we find that what appear to be minor differences in the natural history of these three closely related species may profoundly impact the potential for dispersal, range size, and cladogenesis.     

Polychaete assemblages inhabiting intertidal soft bottoms associated with mangrove systems in Coiba National Park (East Pacific,Panama)

The Coiba National Park is located off the Pacific coast of Panama andincludes several small islands and a larger one which gives its name to the park. Anumber of rivers in Coiba Island have well developed mangrove systems attheir mouths and two of these were selected for the present study: SantaCruz and El Gambute. The first one comprises an assemblage of Rhizopora mangle, Laguncularia racemosa and Pellicierarhizoporae, and shows no evidence of recent human activity. The secondone is a smaller forest, solely composed of short Rhizoporamangle trees, and is recovering from the impact of human activities.During 1997, the tidal flats associated with each system were studied. Toasses the polychaete distribution within each tidal flat, three transects werearranged from the low to the upper intertidal level. Every transect hadthree sampling sites: the first one was adjacent to the low water level, thesecond one at an intermediate point, and the third one close to the highwater level and the mangrove line. At each site, three randomly chosensamples of sediment were taken for faunal study and a further one wascollected for sedimentology study.The polychaetes were identifiedto species or genus level. For each flat, a matrix summarizing the actualnumber of individuals of each species in every sample was created, andpopulation density, species richness and diversity were computed for eachsite. After matrix data had been transformed, a similarity analysis of thesamples was carried out.     

Soil nutrient dynamics in response to irrigation of a Panamanian tropical moist forest

We measured concentrations of soil nutrients (0–15 and 30–35 cm depths) before and after the dry season in control and dry-season irrigated plots of mature tropical moist forest on Barro Colorado Island (BCI) in central Panama to determine how soil moisture affects availability of plant nutrients. Dry-season irrigation (January through April in 1986, 1987, and 1988) enhanced gravimetric soil water contents to wet-season levels (ca. 400 g kg^–1 but did not cause leaching beyond 0.8 m depth in the soil. Irrigation increased concentrations of exchangeable base cations (Ca²⁺, Mg²⁺, K⁺, Na⁺), but it had little effect on concentrations of inorganic N (NH₄^+C, NO₃^– and S (SO₄^2–). These BCI soils had particularly low concentrations of extractable P especially at the end of the dry season in April, and concentrations increased in response to irrigation and the onset of the rainy season. We also measured the response of soil processes (nitrification and S mineralization) to irrigation and found that they responded positively to increased soil moisture in laboratory incubations, but irrigation had little effect on rates in the field. Other processes (plant uptake, soil organic matter dynamics) must compensate in the field and keep soil nutrient concentrations at relatively low levels.     

Behavioral thermoregulation in a tropical gastropod: links to climate change scenarios

Tropical species are vulnerable to global warming because they live at, or near to, their upper thermal threshold limits. Therefore, the predicted increase in the frequency of warming events in the tropics is expected to be critical for the survival of local species. This study explored the major environmental variables which were thought to be correlated with body temperatures (BTs) of the tropical snail Littoraria scabra at the niche level. A correlation between BT and substrate temperature (ST) was detected from field observations which suggests a possible causal relationship between both substrate and BTs. In contrast, there was no correlation between BT and air temperature. Field observations suggest that 33.4 °C may be L. scabra upper limit of substrate surface temperature, although further experiments are needed to assess if the upper limit of physiological tolerance is actually different. As L. scabra individuals were free to choose their substrata, the observed distribution pattern at the niche level is related to L. scabra's behavior. Additionally, substrate surface temperatures were very heterogeneous at centimeter scale (i.e. from 22.5 to 53.1 °C) and L. scabra was shown to select specific STs (i.e. between 22.5 and 33.4 °C) rather than microhabitat type. Therefore, L. scabra did not seem to behaviorally thermoregulate through microhabitat selection nor aggregation. In contrast, behavioral experiments showed that L. scabra has the ability to actively select a thermally favorable site over short temporal scale (i.e. individual average speed of 1.26 cm min^?1) following exposure to high temperatures above 33.4 °C. Hence, this study supports the crucial need to integrate intertidal invertebrate behavioral responses to thermal constraints in climate change studies.     

Contribution to the study of the microfungi in the saprotrophic specialization in tropical forest litter

Abstract

In this study we have reported some results of the fungal colonization in the litter of different plants of the National Park of Tai (Ivory Coast). We have also examined the correlations between the fungal species and the vegetable matrix in order to detect a saprotrophic specialization.     

Mating system of Carapa procera (Meliaceae) in the french guiana tropical forest

The mating system of the tropical rainforest tree Carapa procera was studied for a population located within a sylvicultural trial in French Guiana. We used the mixed-mating model with the genotypes of 47 open-pollinated progenies at ten polymorphic loci, obtained through isozyme analysis. Seeds were collected on both logged and unlogged (control) plots, which were interspersed. A high multilocus outcrossing rate (0.78) was found for the global population. and there was indirect evidence for nonrandom mating for outcrosses. It is argued that this may be due to some kind of positive assortative mating, probably biparental inbreeding. Correlation of selfing between sibs was not significantly different from zero. and the proportion of full-sibs among outcrossed sib-pairs was significant but low (0.16). We found significant differences between outcrossing rates when computing separate estimates for the 14 trees located on logged plots (tm = 0.63) and the 33 trees located on undisturbed plots (tm = 0.85). It is argued that the decrease in density is probably not the only cause of the decrease in outcrossing rate on logged plots. Lastly, several points concerning the reliability of the estimation are outlined. In particular. the downward bias due to ignoring null alleles during genetic interpretations of zymograms is quantified in a few cases by simulating artificial populations.     

The value of a mangrove area in Sarawak

Many arguments have been presented to justify the conservation of tropical forests. In the case of mangrove forests, their preservation can be argued using economic and employment grounds alone. A case study of the Sarawak Mangroves Forest Reserve, Malaysia is presented. Here, the mangroves support marine fisheries worth US$21.1 million p.a. and up to 3000 jobs, timber products worth US$123,217 p.a., and a tourist industry worth US$3.7 million p.a. If the mangroves were to be damaged, all of the fisheries and timber and many of the tourism benefits would be lost. In addition, highly expensive civil engineering works would be incurred to prevent coastal erosion, flooding and other damage. The area is also one of the only remaining refuges for mangrove flora and fauna in Sarawak. If the area were to be converted to aquaculture ponds or oil palm plantations, levels of revenue would be greatly reduced, and the multiple other benefits of mangroves would be lost. Coastal land pressure is not a limiting factor in the State. Considering their economic, employment, coastal protection and species conservation values, mangroves should be conserved and their importance taken into account at all levels in development planning.