Characterization of the benthos, marine debris and bottom fish at Gray’s Reef National Marine Sanctuary

Executive Summary: Baseline characterization of resources is an essential part of marine protected area (MPA) management and is critical to inform adaptive management. Gray’s Reef National Marine Sanctuary (GRNMS) currently lacks adequate characterization of several key resources as identified in the 2006 Final Management Plan. The objectives of this characterization were to fulfill this need by characterizing the bottom fish, benthic features, marine debris, and the relationships among them for the different bottom types within the sanctuary: ledges, sparse live bottom, rippled sand, and flat sand. Particular attention was given to characterizing the different ledge types, their fish communities, and the marine debris associated with them given the importance of this bottom type to the sanctuary. The characterization has been divided into four sections. Section 1 provides a brief overview of the project, its relevance to sanctuary needs, methods of site selection, and general field procedures. Section 2 provides the survey methods, results, discussion, and recommendations for monitoring specific to the benthic characterization. Section 3 describes the characterization of marine debris. Section 4 is specific to the characterization of bottom fish. Field surveys were conducted during August 2004, May 2005, and August 2005. A total of 179 surveys were completed over ledge bottom (n=92), sparse live bottom (n=51), flat sand (n=20), and rippled sand (n=16). There were three components to each field survey: fish counting, benthic assessment, and quantification of marine debris. All components occurred within a 25 x 4 m belt transect. Two divers performed the transect at each survey site. One diver was responsible for identification of fish species, size, and abundance using a visual survey. The second diver was responsible for characterization of benthic features using five randomly placed 1 m2 quadrats, measuring ledge height and other benthic structures, and quantifying marine debris within the entire transect. GRNMS is composed of four main bottom types: flat sand, rippled sand, sparsely colonized live bottom, and densely colonized live bottom (ledges). Independent evaluation of the thematic accuracy of the GRNMS benthic map produced by Kendall et al. (2005) revealed high overall accuracy (93%). Most discrepancies between map and diver classification occurred during August 2004 and likely can be attributed to several factors, including actual map or diver errors, and changes in the bottom type due to physical forces. The four bottom types have distinct physical and biological characteristics. Flat and rippled sand bottom types were composed primarily of sand substrate and secondarily shell rubble. Flat sand and rippled sand bottom types were characterized by low percent cover (0-2%) of benthic organisms at all sites. Although the sand bottom types were largely devoid of epifauna, numerous burrows indicate the presence of infaunal organisms. Sparse live bottom and ledges were colonized by macroalgae and numerous invertebrates, including coral, gorgonians, sponges, and “other” benthic species (such as tunicates, anemones, and bryozoans). Ledges and sparse live bottom were similar in terms of diversity (H’) given the level of classification used here. However, percent cover of benthic species, with the exception of gorgonians, was significantly greater on ledge than on sparse live bottom. Percent biotic cover at sparse live bottom ranged from 0.7-26.3%, but was greater than 10% at only 7 out of 51 sites. Colonization on sparse live bottom is likely inhibited by shifting sands, as most sites were covered in a layer of sediment up to several centimeters thick. On ledge bottom type, percent cover ranged from 0.42-100%, with the highest percent cover at ledges in the central and south-central region of GRNMS. Biotic cover on ledges is influenced by local ledge characteristics. Cluster analysis of ledge dimensions (total height, undercut height, undercut width) resulted in three main categories of ledges, which were classified as short, medium, and tall. Median total percent cover was 97.6%, 75.1%, and 17.7% on tall, medium, and short ledges, respectively. Total percent cover and cover of macroalgae, sponges, and other organisms was significantly lower on short ledges compared to medium and tall ledges, but did not vary significantly between medium and tall ledges. Like sparse live bottom, short ledges may be susceptible to burial by sand, however the results indicate that ledge height may only be important to a certain threshold. There are likely other factors not considered here that also influence spatial distribution and community structure (e.g., small scale complexity, ocean currents, differential settlement patterns, and biological interactions). GRNMS is a popular site for recreational fishing and boating, and there has been increased concern about the accumulation of debris in the sanctuary and potential effects on sanctuary resources. Understanding the types, abundance, and distribution of debris is essential to improving debris removal and education efforts. Approximately two-thirds of all observed debris items found during the field surveys were fishing gear, and about half of the fishing related debris was monofilament fishing line. Other fishing related debris included leaders and spear gun parts, and non-gear debris included cans, bottles, and rope. The spatial distribution of debris was concentrated in the center of the sanctuary and was most frequently associated with ledges rather than at other bottom types. Several factors may contribute to this observation. Ledges are often targeted by fishermen due to the association of recreationally important fish species with this bottom type. In addition, ledges are structurally complex and are often densely colonized by biota, providing numerous places for debris to become stuck or entangled. Analysis of observed boat locations indicated that higher boat activity, which is an indication of fishing, occurs in the center of the sanctuary. On ledges, the presence and abundance of debris was significantly related to observed boat density and physiographic features including ledge height, ledge area, and percent cover. While it is likely that most fishing related debris originates from boats inside the sanctuary, preliminary investigation of ocean current data indicate that currents may influence the distribution and local retention of more mobile items. Fish communities at GRNMS are closely linked to benthic habitats. A list of species encountered, probability of occurrence, abundance, and biomass by habitat is provided. Species richness, diversity, composition, abundance, and biomass of fish all showed striking differences depending on bottom type with ledges showing the highest values of nearly all metrics. Species membership was distinctly separated by bottom type as well, although very short, sparsely colonized ledges often had a similar community composition to that of sparse live bottom. Analysis of fish communities at ledges alone indicated that species richness and total abundance of fish were positively related to total percent cover of sessile invertebrates and ledge height. Either ledge attribute was sufficient to result in high abundance or species richness of fish. Fish diversity (H`) was negatively correlated with undercut height due to schools of fish species that utilize ledge undercuts such as Pareques species. Concurrent analysis of ledge types and fish communities indicated that there are five distinct combinations of ledge type and species assemblage. These include, 1) short ledges with little or no undercut that lacked many of the undercut associated species except Urophycis earlii ; 2) tall, heavily colonized, deeply undercut ledges typically with Archosargus probatocephalus, Mycteroperca sp., and Pareques sp.; 3) tall, heavily colonized but less undercut with high occurrence of Lagodon rhomboides and Balistes capriscus; 4) short, heavily colonized ledges typically with Centropristis ocyurus, Halichoeres caudalis, and Stenotomus sp.; and 5) tall, heavily colonized, less undercut typically with Archosargus probatocephalus, Caranx crysos and Seriola sp.. Higher levels of boating activity and presumably fishing pressure did not appear to influence species composition or abundance at the community level although individual species appeared affected. These results indicate that merely knowing the basic characteristics of a ledge such as total height, undercut width, and percent cover of sessile invertebrates would allow good prediction of not only species richness and abundance of fish but also which particular fish species assemblages are likely to occur there. Comparisons with prior studies indicate some major changes in the fish community at GRNMS over the last two decades although the causes of the changes are unknown. Species of interest to recreational fishermen including Centropristis striata, Mycteroperca microlepis, and Mycteroperca phenax were examined in relation to bottom features, areas of assumed high versus low fishing pressure, and spatial dispersion. Both Mycteroperca species were found more frequently when undercut height of ledges was taller. They often were found together in small mixed species groups at ledges in the north central and southwest central regions of the sanctuary. Both had lower mode size and proportion of fish above the fishery size limit in heavily fished areas of the sanctuary (i.e. high boat density) despite the presence of better habitat in that region. Black sea bass, C. striata, occurred at 98% of the ledges surveyed and appeared to be evenly distributed throughout the sanctuary. Abundance was best explained by a positive relationship with percent cover of sessile biota but was also negatively related to presence of either Mycteroperca species. This may be due to predation by the Mycteroperca species or avoidance of sites where they are present by C. striata. Suggestions for monitoring bottom features, marine debris, and bottom fish at GRNMS are provided at the end of each chapter. The present assessment has established quantitative baseline characteristics of many of the key resources and use issues at GRNMS. The methods can be used as a model for future assessments to track the trajectory of GRNMS resources. Belt transects are ideally suited to providing efficient and quantitative assessment of bottom features, debris, and fish at GRNMS. The limited visibility, sensitivity of sessile biota, and linear nature of ledge habitats greatly diminish the utility of other sampling techniques. Ledges should receive the bulk of future characterization effort due to their importance to the sanctuary and high variability in physical structure, benthic composition, and fish assemblages. (PDF contains 107 pages.)

Quantifying seascape structure: extending terrestrial spatial pattern metrics to the marine realm

Spatial pattern metrics have routinely been applied to characterize and quantify structural features of terrestrial landscapes and have demonstrated great utility in landscape ecology and conservation planning. The important role of spatial structure in ecology and management is now commonly recognized, and recent advances in marine remote sensing technology have facilitated the application of spatial pattern metrics to the marine environment. However, it is not yet clear whether concepts, metrics, and statistical techniques developed for terrestrial ecosystems are relevant for marine species and seascapes. To address this gap in our knowledge, we reviewed, synthesized, and evaluated the utility and application of spatial pattern metrics in the marine science literature over the past 30 yr (1980 to 2010). In total, 23 studies characterized seascape structure, of which 17 quantified spatial patterns using a 2-dimensional patch-mosaic model and 5 used a continuously varying 3-dimensional surface model. Most seascape studies followed terrestrial-based studies in their search for ecological patterns and applied or modified existing metrics. Only 1 truly unique metric was found (hydrodynamic aperture applied to Pacific atolls). While there are still relatively few studies using spatial pattern metrics in the marine environment, they have suffered from similar misuse as reported for terrestrial studies, such as the lack of a priori considerations or the problem of collinearity between metrics. Spatial pattern metrics offer great potential for ecological research and environmental management in marine systems, and future studies should focus on (1) the dynamic boundary between the land and sea; (2) quantifying 3-dimensional spatial patterns; and (3) assessing and monitoring seascape change.

ROV-based Deep Water Monitoring of the Northern Channel Islands Marine Protected Areas Annual Report - 2007

Research cruises were conducted in August-October 2007 to complete the third annual remotely operated vehicle (ROV)-based assessments of nearshore rocky bottom finfish at ten sites in the northern Channel Islands. Annual surveys at the Channel Islands have been conducted since 2004 at four sites and were expanded to ten sites in 2005 to monitor potential marine protected area (MPA)effects on baseline fish density. Six of the ten sites are in MPAs and four in nearby fished reference areas. In 2007 the amount of soft-only substrate on the 141 track lines surveyed was again estimated in real-time in order to target rocky bottom habitat. These real-time estimates of hard and mixed substrate for all ten sites averaged 57%, 1% more than the post-processed average of 56%. Surveys generated 69.9 km of usable video for use in finfish density calculations, with target rocky bottom habitat accounting for 56% (39.1 km) for all sites combined. The amount of rocky habitat sampled by site averaged 3.8 km and ranged from 3.3 km sampled at South Point, a State Marine Reserve (SMR) off Santa Rosa Island, to 4.7 km at Anacapa Island SMR. A sampling goal of 75 transects at all 10 sites was met using real-time habitat estimates combined with precautionary over-sampling by 10%. A total of seventy kilometers of sampling is projected to produce at least seventy-five 100 m2 transects per site. Thirteen of 26 finfish taxa observed were selected for quantitative evaluation over the time series based on a minimum criterion of abundance (0.05/100 m2). Ten of these 13 finfish appear to be more abundant at the state marine reserves relative to fished areas when densities were averaged across the 2005 to 2007 period. One of the species that appears to be more abundant in fished areas was señorita, a relatively small prey species that is not a commercial or recreational target. (PDF contains 83 pages.)

Fish Habitat Utilization Patterns and Evaluation of the Efficacy of Marine Protected Areas in Hawaii: Integration of NOAA Digital Benthic Habitat Mapping and Coral Reef Ecological Studies

Over the past four decades, the state of Hawaii has developed a system of eleven Marine Life Conservation Districts (MLCDs) to conserve and replenish marine resources around the state. Initially established to provide opportunities for public interaction with the marine environment, these MLCDs vary in size, habitat quality, and management regimes, providing an excellent opportunity to test hypotheses concerning marine protected area (MPA) design and function using multiple discreet sampling units. NOAA/NOS/NCCOS/Center for Coastal Monitoring and Assessment’s Biogeography Team developed digital benthic habitat maps for all MLCD and adjacent habitats. These maps were used to evaluate the efficacy of existing MLCDs for biodiversity conservation and fisheries replenishment, using a spatially explicit stratified random sampling design. Coupling the distribution of habitats and species habitat affinities using GIS technology elucidates species habitat utilization patterns at scales that are commensurate with ecosystem processes and is useful in defining essential fish habitat and biologically relevant boundaries for MPAs. Analysis of benthic cover validated the a priori classification of habitat types and provided justification for using these habitat strata to conduct stratified random sampling and analyses of fish habitat utilization patterns. Results showed that the abundance and distribution of species and assemblages exhibited strong correlations with habitat types. Fish assemblages in the colonized and uncolonized hardbottom habitats were found to be most similar among all of the habitat types. Much of the macroalgae habitat sampled was macroalgae growing on hard substrate, and as a result showed similarities with the other hardbottom assemblages. The fish assemblages in the sand habitats were highly variable but distinct from the other habitat types. Management regime also played an important role in the abundance and distribution of fish assemblages. MLCDs had higher values for most fish assemblage characteristics (e.g. biomass, size, diversity) compared with adjacent fished areas and Fisheries Management Areas (FMAs) across all habitat types. In addition, apex predators and other targeted resources species were more abundant and larger in the MLCDs, illustrating the effectiveness of these closures in conserving fish populations. Habitat complexity, quality, size and level of protection from fishing were important determinates of MLCD effectiveness with respect to their associated fish assemblages. (PDF contains 217 pages)

Benthic habitat mapping in the Olympic Coast National Marine Sanctuary: Classification of side scan sonar data from survey HMPR-108-2002-01: Version I

In September 2002, side scan sonar was used to image a portion of the sea floor in the northern OCNMS and was mosaiced at 1-meter pixel resolution using 100 kHz data collected at 300-meter range scale. Video from a remotely-operated vehicle (ROV), bathymetry data, sedimentary samples, and sonar mapping have been integrated to describe geological and biological aspects of habitat and polygon features have been created and attributed with a hierarchical deep-water marine benthic classification scheme (Greene et al. 1999). The data can be used with geographic information system (GIS) software for display, query, and analysis. Textural analysis of the sonar images provided a relatively automated method for delineating substrate into three broad classes representing soft, mixed sediment, and hard bottom. Microhabitat and presence of certain biologic attributes were also populated into the polygon features, but strictly limited to areas where video groundtruthing occurred. Further groundtruthing work in specific areas would improve confidence in the classified habitat map. (PDF contains 22 pages.)

Distribution and sighting frequency of reef fishes in the Florida Keys National Marine Sanctuary

This study analyzed species richness, distribution, and sighting frequency of selected reef fishes to describe species assemblage composition, abundance, and spatial distribution patterns among sites and regions (Upper Keys, Middle Keys, Lower Keys, and Dry Tortugas) within the Florida Keys National Marine Sanctuary (FKNMS) barrier reef ecosystem. Data were obtained from the Reef Environmental Education Foundation (REEF) Fish Survey Project, a volunteer fish-monitoring program. A total of 4,324 visual fish surveys conducted at 112 sites throughout the FKNMS were used in these analyses. The data set contained sighting information on 341 fish species comprising 68 families. Species richness was generally highest in the Upper Keys sites (maximum was 220 species at Molasses Reef) and lowest in the Dry Tortugas sites. Encounter rates differed among regions, with the Dry Tortugas having the highest rate, potentially a result of differences in the evenness in fishes and the lower diversity of habitat types in the Dry Tortugas region. Geographic coverage maps were developed for 29 frequently observed species. Fourteen of these species showed significant regional variation in mean sighting frequency (%SF). Six species had significantly lower mean %SF and eight species had significantly higher mean %SF in the Dry Tortugas compared with other regions. Hierarchical clustering based on species composition (presence-absence) and species % SF revealed interesting patterns of similarities among sites that varied across spatial scales. Results presented here indicate that phenomena affecting reef fish composition in the FKNMS operate at multiple spatial scales, including a biogeographic scale that defines the character of the region as a whole, a reef scale (~50-100 km) that include meso-scale physical oceanographic processes and regional variation in reef structure and associated reef habitats, and a local scale that includes level of protection, cross-shelf location and a suite of physical characteristics of a given reef. It is likely that at both regional and local scales, species habitat requirements strongly influence the patterns revealed in this study, and are particularly limiting for species that are less frequently observed in the Dry Tortugas. The results of this report serve as a benchmark for the current status of the reef fishes in the FKNMS. In addition, these data provide the basis for analyses on reserve effects and the biogeographic coupling of benthic habitats and fish assemblages that are currently underway. (PDF contains 61 pages.)

The status of shark and ray fishery resources in the Gulf of California: applied research to improve management and conservation

Seasonal surveys were conducted during 1998–1999 in Baja California, Baja California Sur, Sonora, and Sinaloa to determine the extent and activities of artisanal elasmobranch fisheries in the Gulf of California. One hundred and forty–seven fishing sites, or camps, were documented, the majority of which (n = 83) were located in Baja California Sur. Among camps with adequate fisheries information, the great majority (85.7%) targeted elasmobranchs during some part of the year. Most small, demersal sharks and rays were landed in mixed species fisheries that also targeted demersal teleosts, but large sharks were usually targeted in directed drift gillnet or, to a lesser extent, surface longline fisheries. Artisanal fishermen were highly opportunistic, and temporally switched targets depending on the local productivity of teleost, invertebrate, and elasmobranch fishery resources. Major fisheries for small sharks (< 1.5 m, “cazón”) were documented in Baja California during spring, in Sonora during autumn–spring, and in Sinaloa during winter and spring. Triakid sharks (Mustelus spp.) dominated cazón landings in the northern states, whereas juvenile scalloped hammerheads (Sphyrna lewini) primarily supported the fishery in Sinaloa. Large sharks (> 1.5 m, “tiburón”) were minor components of artisanal elasmobranch fisheries in Sonora and Sinaloa, but were commonly targeted during summer and early autumn in Baja California and Baja California Sur. The pelagic thresher shark (Alopias pelagicus) and silky shark (Carcharhinus falciformis) were most commonly landed in Baja California, whereas a diverse assemblage of pelagic and large coastal sharks was noted among Baja California Sur landings. Rays dominated summer landings in Baja California and Sinaloa, when elevated catch rates of the shovelnose guitarfish (Rhinobatos productus, 13.2 individuals/vessel/trip) and golden cownose ray (Rhinoptera steindachneri, 11.1 individuals/vesse/trip) primarily supported the respective fisheries. The Sonoran artisanal elasmobranch fishery was the most expansive recorded during this study, and rays (especially R. productus) dominated spring and summer landings in this state. Seasonal catch rates of small demersal sharks and rays were considerably greater in Sonora than in other surveyed states. Many tiburón populations (e.g., C. leucas, C. limbatus, C. obscurus, Galeocerdo cuvier) have likely been overfished, possibly shifting effort towards coastal populations of cazón and rays. Management recommendations, including conducting demographic analyses using available life history data, determining and protecting nursery areas, and enacting seasonal closures in areas of elasmobranch aggregation (e.g., reproduction, feeding), are proposed. Without effective, enforceable management to sustain or rebuild targeted elasmobranch populations in the Gulf of California, collapse of many fisheries is a likely outcome. (PDF contains 243 pages)

Marine biology of the Sierra Leone River Estuary. 1. The physical environment

The Sierra Leone River Estuary is a relatively young drowned river valley, it is shallow except for a deep channel which passes close to the Freetown shoreline. The upper reaches merge into a network of creeks and channels fringed by large areas of mangrove swamps. It is a tidal estuary of the semi-mixed type with the saline oceanic water entering it on a diurnal cycle. The climate of Sierra Leone is marked by a very distinct change between a very wet rainy season and a dry season. The tidal range of the Estuary (spring 3.03m; neap 2.28m) does not impede normal use of the harbour. The tidal variations can be felt as far as 42 miles inland along the water courses of the Sierra Leone River and its tributaries. The volume of fresh water entering the Estuary is large during the rainy season and greatly reduced during the dry season. Consequently there is a marked fall in salinity during the rainy season and higher salinities due to the marine influence prevailing during the dry season. The nature of the shores and bottom, the hydrography and chemistry of the estuarine system have been outlined in relation to the prevailing climatic conditions.

Hypoxic and low pH water in the nearshore marine environments of Monterey Bay, California: characterizing a decade of oxygen and pH, and drivers of variability.

A decade-long time series recorded in southern Monterey Bay, California demonstrates that the shallow, near-shore environment (17 m depth) is regularly inundated with pulses of cold, hypoxic and low pH water. During these episodes, oxygen can drop to biologically threatening levels, and pH levels were lower than expected. Weekly water chemistry monitoring revealed that the saturation state of aragonite (the more soluble form of calcium carbonate) was often below saturation and had a moderate positive relationship with pH, however, analytical and human error could be high. Pulses of hypoxia and low pH water with the greatest intensity arise at the onset of the spring upwelling season, and fluctuations are strongly semidurnal (tidal) and diurnal. Arrival of cold, hypoxic water on the inner shelf typically occurs 3 days after the arrival of a strong upwelling event and appears to be driven by upwelling modulated by internal tidal fluctuations. I found no relationship between the timing of low-oxygen events and the diel solar cycle nor with terrestrial nutrient input. These observations are consistent with advection of hypoxic water from the deep, offshore environment where water masses experience a general decline of temperature, oxygen and pH with depth, and inconsistent with biochemical forcing. Comparisons with concurrent temperature and oxygen time series taken ~20 km away at the head of the Monterey Canyon show similar patterns but even more intense hypoxic events due to stronger semidiurnal forcing there. Analysis of the durations of exposure to low oxygen levels establishes a framework for assessing the ecological relevance of these events. Increasing oceanic hypoxia and acidification of both surface and deep waters may increase the number, intensity, duration and spatial extent of future intrusions along the Pacific coast. Evaluation of the resiliency of nearshore ecosystems such as kelp forests, rocky reefs and sandy habitats, will require consideration of these events.

Season- and depth-dependent variability of a demersal fish assemblage in a large fjord estuary (Puget Sound, Washington)

Fjord estuaries are common along the northeast Pacific coastline, but little information is available on fish assemblage structure and its spatiotemporal variability. Here, we examined changes in diversity metrics, species biomasses, and biomass spectra (the distribution of biomass across body size classes) over three seasons (fall, winter, summer) and at multiple depths (20 to 160 m) in Puget Sound, Washington, a deep and highly urbanized fjord estuary on the U.S. west coast. Our results indicate that this fish assemblage is dominated by cartilaginous species (spotted ratfish [Hydrolagus colliei] and spiny dogfish [Squalus acanthias]) and therefore differs fundamentally from fish assemblages found in shallower estuaries in the northeast Pacific. Diversity was greatest in shallow waters (<40 m), where the assemblage was composed primarily of flatfishes and sculpins, and lowest in deep waters (>80 m) that are more common in Puget Sound and that are dominated by spotted ratf ish and seasonally (fall and summer) by spiny dogfish. Strong depth-dependent variation in the demersal fish assemblage may be a general feature of deep fjord estuaries and indicates pronounced spatial variability in the food web. Future comparisons with less impacted fjords may offer insight into whether cartilaginous species naturally dominate these systems or only do so under conditions related to human-caused ecosystem degradation. Information on species distributions is critical for marine spatial planning and for modeling energy flows in coastal food webs. The data presented here will aid these endeavors and highlight areas for future research in this important yet understudied system.

Anadromous fish as marine nutrient vectors

The tidal freshwater of Virginia supports anadromous herring (Alosa spp.) spawning runs in the spring; however, their importance as nutrient delivery vectors to the freshwater fish food web remains unknown. The stable isotope signatures of fishes from 21 species and four different guilds (predators, carnivores, generalists, and planktivores) were examined in this study to test the hypothesis that marine derived nutrients (MDNs) brought by anadromous fish would be traced into the guilds that incorporated them. Spawning anadromous fish were 13C and 34S-enriched (δ13C and δ34S of approximately 18‰ and 17.7‰, respectively) relative to resident freshwater fish. Of the guilds examined, only predators showed 13C and 34S-enrichment similar to the anadromous fish; however, some generalist catfish also showed enriched signatures. Specific fatty acid δ13C signatures for gizzard shad (Dorosoma cepedianum), blue catfish (Ictalurus furcatus), and alewife (Alosa pseudoharengus), show a 10‰ range among fishes, clearly reflecting isotopically distinct dietary sources. The δ13C and δ34S distribution and range among the freshwater fishes suggest that both autochthonous and allochthonous (terrestrial C3 photosynthetic production and MDN) nutrient sources are important to the tidal freshwater fish community.

Diet composition and food habits of demersal and pelagic marine fishes from Terengganu waters, east coast of Peninsular Malaysia

Fish stomachs from 18 demersal and pelagic fishes from the coast of Terengganu in Malaysia were examined. The components of the fishes’ diets varied in number, weight, and their frequency of occurrence. The major food items in the stomachs of each species were determined using an Index of Relative Importance. A conceptual food web structure indicates that fish species in the study area can be classified into three predatory groups: (1) predators on largely planktivorous or pelagic species; (2) predators on largely benthophagous or demersal species; and (3) mixed feeders that consume both pelagic and demersal species.