An ordination study to view vegetation structure dynamics in disturbed and undisturbed mangrove forests in Kenya and Sri Lanka

The mangrove vegetation of a disturbed and undisturbed site in bothKenya and Sri Lanka was investigated in the field for three vegetation layers:adult trees, young trees, and juvenile trees. A minimum of 25 sample points, inwhich the vegetation was described and environmental factors (salinity, lightintensity, land/water ratio, abundance of herbivorous crabs and snailabundance)were measured or estimated, were taken on each site. Detrended correspondenceanalysis (DCA) and canonical correspondence analysis (CCA) were used tosummarize the data bulk, to investigate the vegetation dynamics(e.g., comparability of species distribution in the threevegetation layers), and/or to link the vegetation data to the environmentalfactors. Results showed that species clusters were relatively easy todelineate,whether mangrove zonation was present or not. Among the environmental factors,the abundance of propagule predators (mostly sesarmid crabs) contributedsignificantly to the variation in vegetation and could be an explanatoryparameter for the observed vegetation data in a majority of sites. In the sitewhere it was not, the most important factor in the ordination was theland/waterratio, which is important at the ecological level as well (link between waterlevel and vegetation dynamics). However, none of the environmental factorscouldsuccessfully explain the total variability in the vegetation data suggestingthat other, more determining factors exist. Our results further provideinformation on the dynamic or non-dynamic nature of a forest and on its abilityto rejuvenate, and may contribute to appropriate forestry management guidelinesin the future.     

Valuing mangrove ecosystem services: linking nutrient retention function of mangrove forests to enhanced agroecosystem production

Mangroves are highly productive wetland ecosystems strategically located at the interface between land and sea. They play an important role in the biogeochemical cycles of the coastal environment, acting as sources of nutrients to adjacent marine and terrestrial ecosystems through active and passive transport. We examined the nutrient contents in mangrove and nonmangrove soils in and around the Bhitarkanika National Park, India and assessed whether the local agricultural producers were aware of and placed a value on this contribution of mangrove forests in enhancing agroecosystem productivity. Soil samples from both mangrove and nonmangrove areas were analysed and quantity of organic carbon, total nitrogen, available phosphorus and potassium were derived. The replacement cost method was used to derive the value of nutrients in mangrove soils. We estimated that each hectare of mangrove contains additional nutrients worth US$232.49 in comparison to nonmangrove areas. The difference in nutrient content in mangrove versus nonmangrove areas gave the value of US$3.37 million for the nutrients in 145 km² of mangrove forests. The agricultural producers were aware that mangrove forests act as a source of nutrients and were willing to pay a higher price for the land adjoining mangrove forests. Around 92% of the producers ranked nutrient retention as a secondary function of mangrove forests. Despite crop depredation from wild ungulates and conflict with salt water crocodiles the agriculturist finds the benefit to cost ratio of mangrove forests high and more than 76% were in favour of mangrove restoration. This study provides an opportunity to highlight the importance of mangrove ecosystems to the livelihoods of the local people and the urgent need to sustain these through proper policy and market interventions.     

Seed predation by insects in tropical mangrove forests: extent and effects on seed viability and the growth of seedlings

Summary Although insects are known to be important seed predators in most terrestrial forests, their role in marine tidal (mangrove) forests has not been examined. Surveys at 12 sites in tropical Australia showed that between 3.1 and 92.7 percent of the seeds or propagules of 12 mangrove tree species had been attacked by insects. Seeds/propagules of six species (Avicennia marina, Bruguiera gymnorrhiza, B. parviflora, Heritiera littoralis, Xylocarpus australasicus and X. granatum) showed consistently high (>40%) levels of insect damage. Greater than 99% of H. littoralis seeds were attacked by insect predators. The survival and subsequent growth in height and biomass of insect-damaged and non-damaged control seeds/propagules of eight mangrove species were compared in shadehouse experiments. Mangrove species fell into 4 groups with regard to the effect of insect predators on their seeds and seedlings. Xylocarpus australasicus and X. granatum had significantly decreased survival (X 48 and 70%) and growth in height (X 61 and 96%) and biomass (X 66 and 85%). Bruguiera parviflora showed decreased survival (X 59%), but there was no effect of insects on the growth of surviving propagules. In contrast, there was no effect of insect damage on the survival of seedlings of Avicennia marina and Bruguiera exaristata, but decreased growth in height (X 22 and 25%) and biomass (X 22 and 26%). Survival and growth of seedlings of Rhizophora stylosa and Bruguiera gymnorrhiza were not affected. The influence of insect seed predators on the survival and growth of seeds of mangrove species in forests will depend on the relative abundance of seed-eating crabs and intertidal position in mangrove forests.This is Contribution No 499 from the Australian Institute of Marine Science     

Ethnobiology, socio-economics and management of mangrove forests: A review

There is growing research interest in the ethnobiology, socio-economics and management of mangrove forests. Coastal residents who use mangroves and their resources may have considerable botanical and ecological knowledgeable about these forests. A wide variety of forest products are harvested in mangroves, especially wood for fuel and construction, tannins and medicines. Although there are exceptions, mangrove forest products are typically harvested in a small-scale and selective manner, with harvesting efforts and impacts concentrated in stands that are closer to settlements and easiest to access (by land or by sea). Mangroves support diverse, local fisheries, and also provide critical nursery habitat and marine productivity which support wider commercial fisheries. These forests also provide valuable ecosystem services that benefit coastal communities, including coastal land stabilization and storm protection. The overlapping of marine and terrestrial resources in mangroves creates tenure ambiguities that complicate management and may induce conflict between competing interests. Mangroves have been cut and cleared extensively to make way for brackish water aquaculture and infrastructure development. More attention is now given to managing remaining forests sustainably and to restoring those degraded from past use. Recent advances in remotely sensed, geo-spatial monitoring provide opportunities for researchers and planners to better understand and improve the management of these unique forested wetlands.     

Seedling recruitment patterns in a Belizean mangrove forest: effects of establishment ability and physico-chemical factors

A field study was conducted to evaluate the relative importance of factors affecting seedling establishment and survival on a mangrove-dominated island in Belize. An examination of spatial patterns of seedling relative densities in relation to reproductive adults and physico-chemical conditions provided correlative information on factors affecting mangrove regeneration patterns. Distance from reproductive adults explained 89–94% of the variation in relative density of Rhizophora mangle seedlings, whereas availability of resources (light and NH₄) explained 73–80% of variation in Avicennia germinans seedling relative density. Just after dispersal (December), 89% of the variation in Laguncularia racemosa seedling relative density was attributable to distance from reproductive adults, but 7 months later (July) 74% of the variation was explained by intensity of flooding- and salinity-related stresses. Survivorship (after 2.5 years) of propagules and seedlings of R. mangle and A. germinans transplanted to zones of contrasting physico-chemical conditions demonstrated that: (1) mortality was highest during the establishment phase and major causes were failure to strand before viability was lost, consumption by predators and desiccation; and (2) after establishment, differences in sensitivity to physicochemical stress factors such as flooding (A. germinans) and initial orientation of the seedling axis (R. mangle) exerted a further influence on seedling survival. The results indicate that seedling recruitment in these neotropical forests is strongly influenced by dispersal patterns, differential establishment abilities and effects of physico-chemical factors that vary with elevation and distance from the shoreline.     

Organic carbon dynamics in mangrove ecosystems: A review

Our current knowledge on production, composition, transport, pathways and transformations of organic carbon in tropical mangrove environments is reviewed and discussed. Organic carbon entering mangrove foodwebs is either produced autochthonously or imported by tides and/or rivers. Mangrove litter and benthic microalgae are usually the most important autochthonous carbon sources. Depending on local conditions, phytoplankton and seagrass detritus imported with tides may represent a significant supplementary carbon input. Litter handling by the fauna not only affects microbial carbon transformations, but also the amount of organic carbon available for export. Most mangrove detritus that enters the sediment is degraded by microorganisms. Aerobic respiration and anaerobic sulfate reduction are usually considered the most important microbial respiration processes, but recent evidence suggests that iron respiration may be important in mangrove sediments as well. Organic carbon that escapes microbial degradation is stored in sediments and in some mangrove ecosystems, organic-rich sediments may extend to several meters depth. Many mangrove forests also lose a significant fraction of their net primary production to coastal waters. Large differences occur between mangrove forests with respect to litter production and export. Mangrove-derived DOC is also released into the water column and can add to the total organic carbon export. Numerous compounds have been characterized from mangrove tissues, including carbohydrates, amino acids, lignin-derived phenols, tannins, fatty acids, triterpenoids and n-alkanes. Many of these may, together with stable isotopes, exhibit a strong source signature and are potentially useful tracers of mangrove-derived organic matter. Our knowledge on mangrove carbon dynamics has improved considerably in recent years, but there are still significant gaps and shortcomings. These are emphasized and relevant research directions are suggested.     

Investigating the role of impoundment and forest structure on the resistance and resilience of mangrove forests to hurricanes

The present study evaluates how a prolonged artificial flooding regime in impounded mangrove forests influences the regeneration after medium-sized forest destruction created by two hurricanes in 2004 in the Indian River Lagoon, Florida, USA. Disturbance patterns, forest structure, and regeneration were investigated. We found disturbed areas, characterized by uprooted and snapped trees, and intact forests in close proximity. The canopy turnover was greater in forests in higher succession stages with median tree diameter at breast height of 7.6 ± 5.7 cm compared to the intact forest with 3.7 ± 1.2 cm. Larger trees with lower densities were more susceptible to hurricane damage. We observed that regeneration of the open patches was dominated by the flood-tolerant species Rhizophora mangle (89.9%) instead of the faster-growing pioneer species Laguncularia racemosa (7.0%). Some of the disturbed areas created by the hurricanes were not recolonized. The regeneration rate of the disturbed areas expressed by vegetation closure >50% differed between almost zero to 100% in three different impoundments. We concluded that the artificial flooding regime imposed by impoundment is the predominant selective force in the successive process according to the species composition. However, we were not able to detect the driving factors that prevented mangrove establishment in disturbed areas.     

Allometry, biomass, and productivity of mangrove forests: A review

We review 72 published articles to elucidate characteristics of biomass allocation and productivity of mangrove forests and also introduce recent progress on the study of mangrove allometry to solve the site- and species-specific problems. This includes the testing of a common allometric equation, which may be applicable to mangroves worldwide. The biomass of mangrove forests varies with age, dominant species, and locality. In primary mangrove forests, the above-ground biomass tends to be relatively low near the sea and increases inland. On a global scale, mangrove forests in the tropics have much higher above-ground biomass than those in temperate areas. Mangroves often accumulate large amounts of biomass in their roots, and the above-ground biomass to below-ground biomass ratio of mangrove forests is significantly low compared to that of upland forests (ANCOVA, P < 0.01). Several studies have reported on the growth increment of biomass and litter production in mangrove forests. We introduce some recent studies using the so-called “summation method” and investigate the trends in net primary production (NPP). For crown heights below 10 m, the above-ground NPP of mangrove forests is significantly higher (ANOVA, P < 0.01) than in those of tropical upland forests. The above-ground litter production is generally high in mangrove forests. Moreover, in many mangrove forests, the rate of soil respiration is low, possibly because of anaerobic soil conditions. These trends in biomass allocation, NPP, and soil respiration will result in high net ecosystem production, making mangrove forests highly efficient carbon sinks in the tropics.     

Dynamics of animal movement in an ecological context: dragonfly wing damage reduces flight performance and predation success

Much of our understanding of the control and dynamics of animal movement derives from controlled laboratory experiments. While many aspects of animal movement can be probed only in these settings, a more complete understanding of animal locomotion may be gained by linking experiments on relatively simple motions in the laboratory to studies of more complex behaviours in natural settings. To demonstrate the utility of this approach, we examined the effects of wing damage on dragonfly flight performance in both a laboratory drop–escape response and the more natural context of aerial predation. The laboratory experiment shows that hindwing area loss reduces vertical acceleration and average flight velocity, and the predation experiment demonstrates that this type of wing damage results in a significant decline in capture success. Taken together, these results suggest that wing damage may take a serious toll on wild dragonflies, potentially reducing both reproductive success and survival.     

Long-term retrospection on mangrove development using transdisciplinary approaches: A review

Large ecosystem processes often take place beyond the observation time of a researcher. Yet, through retrospective research scientists can approach and understand ecosystem changes. This contributes to the fundamental understanding of both human-induced and natural dynamics in ecosystems world-wide. This also holds for fast changing coastal areas with mangrove ecosystems, which are important for biodiversity, for coastal protection, and for the daily livelihood of millions of people in tropical coastal developing countries. In addition, retrospective research generates a basis for predictions that can be used early on to protect an ecosystem. In attempting to protect ecosystems from adverse human-induced change and destruction, and to manage them for sustainability, scientists are only beginning to investigate and understand natural ecosystem dynamics. It is important and advisable to gather, combine and analyse all possible data that allow a researcher to look back in time. This paper reviews the available retrospective methods, and highlights the transdisciplinary way (i.e. combination between basic and applied sciences on one hand, and social and human sciences on the other) in which retrospective research on a scale between months and centuries can be carried out, but it also includes methods on larger scales that may be marginally relevant. The paper particularly emphasizes the lack of transdisciplinary (not interdisciplinary) integration between sciences in retrospective research on mangrove forests in the past.     

Regeneration of<Emphasis Type="Italic"> Rhizophora mangle</Emphasis> in a Caribbean mangrove forest: interacting effects of canopy disturbance and a stem-boring beetle

Current theory predicts that in low-density, seed-limited plant populations, seed predation will be more important than competition in determining the number of individuals that reach maturity. However, when plant density is high, competition for microsites suitable for establishment and growth is expected to have a relatively greater effect. This dichotomous perspective does not account for situations in which the risk of seed predation differs inside versus outside recruitment microsites. We report the results of a field experiment and sampling studies that demonstrate such an interaction between microsite quality and the risk of propagule predation in mangrove forests on the Caribbean coast of Panama, where it appears to play a key role in shaping the demography and dynamics of the mangrove, Rhizophora mangle. Rhizophora's water-borne propagules establish wherever they strand, but long-term sampling revealed that only those that do so in or near lightning-created canopy gaps survive and grow to maturity. These microsites afford better growth conditions than the surrounding understory and, as importantly, provide a refuge from predation by the scolytid beetle, Coccotrypes rhizophorae. This refuge effect was confirmed with a field experiment in which Rhizophora seedlings were planted at different positions relative to gap edges, from 5 m inside to 20 m outside the gap. Mortality due to beetle attack increased linearly from an average of 10% inside a gap to 72% at 20 m into the forest. The interaction between canopy disturbance and propagule predation may be having a large impact on the composition of our study forests. Being shade-tolerant, Rhizophora seedlings that escape or survive beetle attack can persist in the understory for years. However, the high rate of beetle-induced mortality effectively eliminates the contribution of advance regeneration by Rhizophora saplings to gap succession. This may explain why the shade-intolerant mangrove, Laguncularia racemosa, is able to co-dominate the canopy in low intertidal forests at our study sites.     

Molecular ecology and biogeography of mangrove trees towards conceptual insights on gene flow and barriers: A review

In this review the most recent contributions to the field of molecular ecology and biogeography of mangrove trees are considered. Emphasis is on the obtained information of the different molecular marker methods used in mangrove genetics and on the potential to infer biogeographical patterns. Isozymes on average showed low or even no polymorphism in mangrove trees similar as known in seagrasses. The outcrossing Avicennia seems to be the most variable mangrove tree for isozymes. Both low amounts of interpretable allozymes and difficulties in maintaining the enzyme activity have reduced the number of successful studies during the isozyme era. Dominant marker methods (RAPD, AFLP and ISSR) were successful to demonstrate differences in amplified DNA products at large-scale geographical distances within Avicennia species and to estimate species relationships. Hybrid testing seldom revealed hybridization among tree species. The most promising markers (microsatellites or SSR) were only recently developed and will continue to provide evidence in future studies. SSR loci in Avicennia seem to show relatively low levels of polymorphism, though clearly demonstrating that populations located at the edge of the species range can be even more depauperated. Populations located more central in their native range and situated along the same coastline such as reported in Rhizophora, are expected to be only weakly differentiated due to increased levels of gene flow. Haplotypic chloroplast variants (PCR-RFLP) or sequences revealed strong genetic structuring between populations of Avicennia, Kandelia and Ceriops from different biogeographical oceanic regions. Recent views on long-distance dispersal and on gene flow across oceans as well as along the same coastline are discussed. A comparative analysis on genetic variables across species and regions indicated general trends in the partitioning of genetic variation. A conceptual map with a worldwide overview of those regions where high levels of gene flow were reported and of other regions that were considered as effective barriers, is presented. As an aim to increase the number of reliable comparisons of genetic variables across species or regions and to increase the relevance of mangrove genetics for local conservation issues, recommendations on the molecular markers and on the sampling design of individuals and populations are given within a conceptual context of evolutionary significant units.     

Fauna as passengers and drivers in vegetation restoration: A synthesis of processes and evidence

Efforts to restore terrestrial woody ecosystems to former agricultural land are typically based on plant‐focused actions, often neglecting fauna. However, the processes that maintain or restore the health and integrity of these ecosystems involve many animal–plant interactions. Here, I synthesise information about these relationships and the implications for revegetation practice. Fauna have often been viewed as passengers, responding passively to plant‐focused revegetation. This view involves two surrogacy assumptions: first, that vegetation attributes can indicate habitat sufficiency for fauna; second, that animals will be capable of dispersing to the restored habitat and of establishing populations there. Habitat sufficiency depends on how resources such as food and nest sites can be indicated by vegetation attributes and how they interact with an animal's species‐specific requirements. Dispersal and establishment depend on proximity to source populations in habitat elsewhere, the type of intervening habitat and the intrinsic mobility of different species. Evidence about the effects of age, revegetation type and spatial context in relation to animal communities indicates that it can often be invalid to assume vegetation surrogacy. Fauna can also drive the developmental trajectories of floristic diversity and composition during revegetation, because animal–plant interactions frequently mediate life‐history transitions that determine seedling recruitment. Frugivore‐mediated seed dispersal is the best studied, but animals also directly influence early‐stage tree recruitment, especially through their roles in seed predation, seedling herbivory and indirectly through top‐down cascades that include large carnivores. These processes have been insufficiently recognised or studied, although some recent work highlights their significance. Intervening to alter abundances of functionally important animals could be useful in accelerating the redevelopment of woody vegetation. Further research is needed to clarify animals’ roles as both passengers and drivers during revegetation, especially manipulative experiments and innovative restoration trials, in which animals and plants are considered together from the outset.     

The habitat function of mangroves for terrestrial and marine fauna: A review

Mangroves are defined by the presence of trees that mainly occur in the intertidal zone, between land and sea, in the (sub) tropics. The intertidal zone is characterised by highly variable environmental factors, such as temperature, sedimentation and tidal currents. The aerial roots of mangroves partly stabilise this environment and provide a substratum on which many species of plants and animals live. Above the water, the mangrove trees and canopy provide important habitat for a wide range of species. These include birds, insects, mammals and reptiles. Below the water, the mangrove roots are overgrown by epibionts such as tunicates, sponges, algae, and bivalves. The soft substratum in the mangroves forms habitat for various infaunal and epifaunal species, while the space between roots provides shelter and food for motile fauna such as prawns, crabs and fishes. Mangrove litter is transformed into detritus, which partly supports the mangrove food web. Plankton, epiphytic algae and microphytobenthos also form an important basis for the mangrove food web. Due to the high abundance of food and shelter, and low predation pressure, mangroves form an ideal habitat for a variety of animal species, during part or all of their life cycles. As such, mangroves may function as nursery habitats for (commercially important) crab, prawn and fish species, and support offshore fish populations and fisheries. Evidence for linkages between mangroves and offshore habitats by animal migrations is still scarce, but highly needed for management and conservation purposes. Here, we firstly reviewed the habitat function of mangroves by common taxa of terrestrial and marine animals. Secondly, we reviewed the literature with regard to the degree of interlinkage between mangroves and adjacent habitats, a research area which has received increasing attention in the last decade. Finally, we reviewed current insights into the degree to which mangrove litter fuels the mangrove food web, since this has been the subject of long-standing debate.     

A review of mangrove rehabilitation in the Philippines: successes,failures and future prospects

From half a million hectares at the turn of the century, Philippine mangroves have declined to only 120,000 ha while fish/shrimp culture ponds have increased to 232,000 ha. Mangrove replanting programs have thus been popular, from community initiatives (1930s–1950s) to government-sponsored projects (1970s) to large-scale international development assistance programs (1980s to present). Planting costs escalated from less than US$100 to over $500/ha, with half of the latter amount allocated to administration, supervision and project management. Despite heavy funds for massive rehabilitation of mangrove forests over the last two decades, the long-term survival rates of mangroves are generally low at 10–20%. Poor survival can be mainly traced to two factors: inappropriate species and site selection. The favored but unsuitable Rhizophora are planted in sandy substrates of exposed coastlines instead of the natural colonizers Avicennia and Sonneratia. More significantly, planting sites are generally in the lower intertidal to subtidal zones where mangroves do not thrive rather than the optimal middle to upper intertidal levels, for a simple reason. Such ideal sites have long been converted to brackishwater fishponds whereas the former are open access areas with no ownership problems. The issue of pond ownership may be complex and difficult, but such should not outweigh ecological requirements: mangroves should be planted where fishponds are, not on seagrass beds and tidal flats where they never existed. This paper reviews eight mangrove initiatives in the Philippines and evaluates the biophysical and institutional factors behind success or failure. The authors recommend specific protocols (among them pushing for a 4:1 mangrove to pond ratio recommended for a healthy ecosystem) and wider policy directions to make mangrove rehabilitation in the country more effective.     

Aromatic interactions in peptides: impact on structure and function

Aromatic interactions, including pi-pi, cation-pi, aryl-sulfur, and carbohydrate-pi interactions, have been shown to be prevalent in proteins through protein structure analysis, suggesting that they are important contributors to protein structure. However, the magnitude and significance of aromatic interactions is not defined by such studies. Investigation of aromatic interactions in the context of structured peptides has complemented studies of protein structure and has provided a wealth of information regarding the role of aromatic interactions in protein structure and function. Recent advances in this area are reviewed.     

Canopy gaps formed by mangrove trimming: an experimental test of impact on litter fall and standing litter stock in Southwest Florida (USA)

Mangroves in Florida (USA) are subject to horticultural pruning that may increase the size of canopy gaps and alter rates of litter production and accumulation. Mangrove canopy gap formation is a common phenomenon, known to alter abiotic conditions near the forest floor. Using a series of field experiments in Rookery Bay, Florida, the effects of mangrove trimming on canopy density, mangrove litter production, standing litter stocks, and the decomposition rate of Rhizophora mangle leaves on the forest floor were assessed. Litter trap collections over the year following mangrove trimming indicated that pruned mangrove stands (canopy coverage: 42.8±0.9%; mean±S.E.) produced approximately one-half of the litter of mangrove stands with relatively complete canopies (canopy coverage: 72.1±0.5%). However, there was no significant difference between the mass of standing litter on the forest floor beneath reduced canopy and intact canopy mangroves. Also, R. mangle leaves held on the forest floor in fiberglass litter bags at both reduced canopy and control sites did not decompose at different rates over 28 days. These results indicate that while system-wide mangrove litter production should be reduced by the formation of these gaps in mangrove forests, postproduction influences may obscure any site-specific declines in standing litter stocks.     

Environmental drivers in mangrove establishment and early development: A review

Mangroves have a global distribution within coastal tropical and subtropical climates, and have even expanded to some temperate locales. Where they do occur, mangroves provide a plethora of goods and services, ranging from coastal protection from storms and erosion to direct income for human societies. The mangrove literature has become rather voluminous, prompting many subdisciplines within a field that earlier in the 20th century received little focus. Much of this research has become diffuse by sheer numbers, requiring detailed syntheses to make research results widely available to resource managers. In this review, we take an inclusive approach in focusing on eco-physiological and growth constraints to the establishment and early development of mangrove seedlings in the intertidal zone. This is a critical life stage for mangroves, i.e., the period between dispersal and recruitment to the sapling stage. We begin with some of the research that has set the precedent for seedling-level eco-physiological research in mangroves, and then we focus on recent advances (circa. 1995 to present) in our understanding of temperature, carbon dioxide, salinity, light, nutrient, flooding, and specific biotic influences on seedling survival and growth. As such, we take a new approach in describing seedling response to global factors (e.g., temperature) along with site-specific factors (e.g., salinity). All variables will strongly influence the future of seedling dynamics in ways perhaps not yet documented in mature forests. Furthermore, understanding how different mangrove species can respond to global factors and regional influences is useful for diagnosing observed mortality within mangrove wetlands, managed or natural. This review provides an updated eco-physiological knowledge base for future research and reforestation activity, and for understanding important links among climate change, local physico-chemical condition, and establishment and early growth of mangrove seedlings.     

Experimental separation of effects of consumers on sessile prey in the low zone of a rocky shore in the Bay of Panama: direct and indirect consequences of food web complexity

The effects of predation by a diverse assemblage of consumers on community structure of sessile prey was evaluated in the low rocky intertidal zone at Taboguilla Island in the Bay of Panama. Four functional groups of consumers were defined: (1) large fishes, (2) small fishes and crabs, (3) herbivorous molluscs, and (4) predaceous gastropods, (l) and (2) included fast-moving consumers and (3) and (4) included slow-moving consumers. Experimental treatments were: no consumers deleted (all groups present), most combinations of deletions of single groups (i.e., one group absent, three present), pairs of groups deleted (two absent, two present), trios of groups deleted (three absent, one present), and the entire consumer assemblage deleted (all groups absent). Changes in abundance (percent cover) of crustose algae, solitary sessile invertebrates, foliose algae, and colonial sessile invertebrates were quantified periodically in 2–4 plots of each treatment from February 1977 to January 1980 after the initiation of the experiment in January 1977.

Space on this shore is normally dominated by crustose algae; foliose algae, solitary sessile invertebrates, and colonial sessile invertebrates are all rare. After deletion of all consumers, ephemeral green algae increased from 0 to nearly 70% cover. Thereafter, a succession of spatial dominants occurred, with peak abundances as follows: the foliose coralline alga Jania spp. by July 1977, the barnacle Balanus inexpectatus by April 1978, and the rock oyster Chama echinata by January 1980. Although no longer occupying primary rock space, Jania persisted as a dominant or co-dominant turf species (with the brown alga Giffordia mitchelliae and/or the hydrozoan Abietinaria sp.) by colonizing shells of sessile animals as they became abundant instead of the rock surface.

Multivariate analysis variance (MANOVA) indicated that the effect of each group was as follows. Molluscan herbivores grazed foliose algae down to the grazer-resistant, but competitively inferior algal crusts, altered the relative abundances of the crusts, and inhibited recruitment of sessile invertebrates. Predaceous gastropods reduced the abundance of solitary sessile animals. Small fishes and crabs, and large fishes reduced the cover of solitary and colonial sessile animals and foliose algae, although they were incapable of grazing the foliose algae down to the rock surface. Many of the effects of each consumer group on prey groups or species were indirect; some effects were positive and some were negative. The variety of these indirect effects was due to both consumer-prey interactions among the consumers, and competitive or commensalistic interactions among the sessile prey. Comparison of the sum of the effects of each of the single consumer groups (i.e., the sum of the effect observed in treatments with one group absent, three present) with the total effects of all consumers (i.e., the effect observed in the treatment with all groups absent) indicates that a “keystone” consumer was not present in this community. Rather, the impacts of the consumer groups were similar but, due to dietary overlap and compensatory changes among the consumers, not readily detected in deletions of single consumer groups. The normally observed dominance of space by crustose algae is thus maintained by persistent, intense predation by a diverse assemblage of consumers on potentially dominant sessile animals and foliose algae. The large difference in structure between this and temperate intertidal communities seems due to differences in degree, not kind of ecological processes which produce the structure.