Effects of Artisanal Fishing on Marine Communities in the Galápagos Islands

Abstract: The Galápagos Islands harbor some of the least impacted marine ecosystems in the tropics, but there are indications that local artisanal fishing is affecting exploited marine communities. To quantify these effects, I sampled communities of fishes and sea urchins at a number of heavily fished and lightly fished sites throughout the central islands of the archipelago. Sites were selected based on information collected as part of a local fisheries monitoring study and standardized across a number of abiotic factors. Abundance and biomass of the primary target species were significantly lower in the heavily fished sites than in the lightly fished sites. Community structure also differed between heavily and lightly fished sites. Cluster analyses of the full community of fishes and a subset of nontarget fishes revealed that sites within a treatment were more similar to one another than sites between treatments. Herbivorous fishes tended to be lower in abundance and sea urchins tended to be higher in abundance in heavily fished sites, but these differences were not significant. My results are encouraging in that the direct effects of artisanal fishing are limited to the primary target species, which probably results from a high specificity of fishing gear. The differences in community structure, however, suggest that artisanal fishing also has cascading effects on noncommercial species throughout the community. An improved understanding of important ecological interactions, increased ecological and fishery monitoring, and effective precautionary management are needed to ensure that human effects in these waters remain minimal.     

Effect of predation threat on repeatability of individual crab behavior revealed by mark-recapture

The persistence of behavioral types in situ and the drivers of persistence are central to predicting the ecological effects of intraspecific behavioral variation. We surveyed individual refuge use of mud crabs (Panopeus herbstii), a behavior related to the strength of a trait-mediated trophic cascade in oyster reefs, in the absence and presence of toadfish (Opsanus tau) predation threat. We then released these crabs into the field and using mark-recapture, measured the repeatability of this behavior in the absence and presence of threat, and how behavioral change was affected by time in the field (a month on average, up to 81 days), crab size, and sex. Because crabs exhibited some evidence of a circatidal rhythm in refuge use, we also tested how tidal height during observation influenced behavioral change. Predation threat increased refuge use, and small crabs used the refuge more than large crabs, particularly under threat. In recaptured crabs, refuge use was more repeatable under threat. Neither time in the field, crab size, crab sex, nor tidal height had any effect on behavioral change. Our results support the non-mutually exclusive hypotheses that (1) prey organisms in the presence, rather than absence, of predation threat should exhibit less behavioral variability because the fear of dying (a severe fitness consequence) should take precedence over less immediately important influences on behavior (e.g., hunger) and that (2) individual behaviors tied to fixed traits (e.g., the body size dependence of refuge use under threat in this study), rather than variable traits, should be more repeatable over time.     

A blended hazard identification methodology to support process diagnosis

A novel hazard identification methodology applied to process systems is presented in this paper. This blended hazard identification (BLHAZID) methodology blends two different types of HAZID methods: the function-driven and component-driven approach. The BLHAZID method is based on a conceptual framework called the Functional Systems Framework, which describes structure–function–goal relationships in process systems.The goals of the BLHAZID methodology are to generate outcomes that contain a high coverage of hazards, describe detailed failure causality in process systems and express this knowledge in a structured form for effective reused in subsequent applications, such as fault diagnosis, operator training, design reviews, fault and event tree construction and hazard updates to satisfy major hazard facility requirements.Both the BLHAZID methodology and the Functional Systems Framework were developed with involvement and advice from two major industrial partners. An industrial case study of a benzene saturation unit is presented to illustrate how the BLHAZID methodology operates in practice.     

Re-creating missing population baselines for Pacific reef sharks

Sharks and other large predators are scarce on most coral reefs, but studies of their historical ecology provide qualitative evidence that predators were once numerous in these ecosystems. Quantifying density of sharks in the absence of humans (baseline) is, however, hindered by a paucity of pertinent time-series data. Recently researchers have used underwater visual surveys, primarily of limited spatial extent or nonstandard design, to infer negative associations between reef shark abundance and human populations. We analyzed data from 1607 towed-diver surveys (>1 ha transects surveyed by observers towed behind a boat) conducted at 46 reefs in the central-western Pacific Ocean, reefs that included some of the world's most pristine coral reefs. Estimates of shark density from towed-diver surveys were substantially lower (<10%) than published estimates from surveys along small transects (<0.02 ha), which is not consistent with inverted biomass pyramids (predator biomass greater than prey biomass) reported by other researchers for pristine reefs. We examined the relation between the density of reef sharks observed in towed-diver surveys and human population in models that accounted for the influence of oceanic primary productivity, sea surface temperature, reef area, and reef physical complexity. We used these models to estimate the density of sharks in the absence of humans. Densities of gray reef sharks (Carcharhinus amblyrhynchos), whitetip reef sharks (Triaenodon obesus), and the group "all reef sharks" increased substantially as human population decreased and as primary productivity and minimum sea surface temperature (or reef area, which was highly correlated with temperature) increased. Simulated baseline densities of reef sharks under the absence of humans were 1.1-2.4/ha for the main Hawaiian Islands, 1.2-2.4/ha for inhabited islands of American Samoa, and 0.9-2.1/ha for inhabited islands in the Mariana Archipelago, which suggests that density of reef sharks has declined to 3-10% of baseline levels in these areas.     

Density-dependent facilitation cascades determine epifaunal community structure in temperate Australian mangroves

Co-occurring foundation species can determine biological community structure via facilitation cascades. We examined the density dependencies of facilitation cascades, including how the density of a basal foundation species influences the density of secondary foundation species, and how the density of secondary foundation species influences community structure. The system in which we assessed density dependencies was a temperate mangrove forest in which pneumatophores trap the fucoid alga Hormosira banksii and provide substrate for the oyster, Saccostrea glomerata. The alga and oyster in turn determine benthic community structure. In the field, algal biomass was positively correlated with pneumatophore density. Oysters, by contrast, were highly over-dispersed and correlated with the presence/absence of pneumatophores. Epifaunal abundance and species richness were positively correlated with algal and oyster abundance, but their effects were independent. The positive effect of pneumatophore density on epifauna was primarily an indirect effect of trapping more algae. Pneumatophores did not directly influence invertebrate communities. Experiments revealed that, at very low pneumatophore densities, algal retention was insufficient to facilitate epifauna above that found on pneumatophores alone. At higher densities, however, increasing the density of pneumatophores increased algal retention, and the density and diversity of associated invertebrates. Shading by the mangrove canopy reduced algal biomass but did not modify the density-dependent nature of the cascade. Our results extend facilitation theory by showing that the density of both basal and secondary foundation species can be critical in triggering facilitation cascades. Our study also reveals that, where foundation species co-occur, multiple, independent cascades may arise from a single basal facilitator. These findings enhance our understanding of the role of density-dependent facilitation cascades in community assembly.     

Dissolved histamine: a potential habitat marker promoting settlement and metamorphosis in sea urchin larvae

Many species of marine invertebrate larvae settle and metamorphose in response to chemicals produced by organisms associated with the adult habitat, and histamine is a cue for larvae of the sea urchin Holopneustes purpurascens. This study investigated the effect of histamine on larval metamorphosis of six sea urchin species. Histamine induced metamorphosis in larvae of three lecithotrophic species (H. purpurascens, Holopneustes inflatus and Heliocidaris erythrogramma) and in one planktotrophic species (Centrostephanus rodgersii). Direct comparisons of metamorphic rates of lecithotrophic and planktotrophic larvae in assays cannot be made due to different proportions of larvae being competent. Histamine (10 μM) induced metamorphosis in 95% of larvae of H. purpurascens and H. inflatus after 1 h, while the coralline alga Amphiroa anceps induced metamorphosis in 40–50% of these larvae. Histamine (10 μM) and A. anceps induced 40 and 80% metamorphosis, respectively, in the larvae of H. erythrogramma after 24 h. Histamine (10 μM) and the coralline alga Corallina sp. induced 30 and 70% metamorphosis, respectively, in the larvae of C. rodgersii after 24 h. No metamorphosis of any larval species occurred in seawater controls. Larvae of two planktotrophic species (Tripneustes gratilla and Heliocidaris tuberculata) did not metamorphose in response to histamine. Seagrasses, the host plants of H. inflatus, induced rapid metamorphosis in larvae of the two Holopneustes species, and several algae induced metamorphosis in C. rodgersii larvae. Histamine leaching from algae and seagrasses may act as a habitat marker and metamorphic cue for larvae of several ecologically important sea urchin species.     

Urban marine ecology in southern California: the ability of riprap structures to serve as rocky intertidal habitat

Increasing human populations along marine coastlines has lead to increasing urbanization of the marine environment. Despite decades of investigations on terrestrial ecosystems, the effect of urbanization on marine life is not well understood. Riprap is the rocky rubble used to build jetties, breakwaters, and armored shorelines. Roughly 30% of the southern California shoreline supports some form of riprap, while 29% of the shoreline is natural rocky substrate. Astonishingly few studies have investigated this anthropogenic rocky habitat even though it rivals a natural habitat in area on a regional scale along a coastline that has been extensively studied. In this study, I compared the diversity and community structure of exposed rocky intertidal communities on four riprap and four natural sites in southern California. I ask the following questions: (1) does diversity or community composition differ between intertidal communities on riprap and natural rocky habitats in southern California, (2) if so, which organisms contribute to those differences, (3) which physical factors are contributing to these differences, and (4) do riprap habitats support higher abundances of invasive species than natural habitats? On average, riprap and natural rocky habitats in wave exposed environments in southern California did not differ from each other in diversity or community composition when considering the entire assemblage. However, when only mobile species were considered, they occurred in greater diversity on natural shores. These differences appear to be driven by wave exposure. The presence of invasive species was negligible in both natural and riprap habitats.     

Worker reproductive parasitism and drift in the western honeybee Apis mellifera

When a honeybee (Apis spp.) colony loses its queen and is unable to rear a new one, some of the workers activate their ovaries and produce eggs. When a colony has a queen (i.e., it is queenright) almost all worker-laid eggs are eaten, but when hopelessly queenless, the workers become more tolerant of worker-laid eggs and rear some of them to adult drones. This increased tolerance renders a queenless colony vulnerable to worker reproductive parasitism, wherein unrelated workers enter the colony and lay eggs. Here, we show that the proportion of unrelated (non-natal) workers significantly decreases after an Apis mellifera colony becomes queenless. The remaining non-natal workers are as likely to have activated ovaries as natal workers, yet they produce more eggs than natal workers, resulting in significantly higher reproductive success for non-natal workers. In a second experiment, we provided queenless and queenright workers with a choice to remain in their own colony or to join a queenless or queenright colony nearby. The experiment was set up such that worker movement was unlikely to be due to simple orientation errors. Very few workers joined another colony, and there was no preference for workers to drift into or out of queenless or queenright colonies, in accordance with the proportion of non-natal workers declining significantly after becoming queenless in the first experiment.     

The ecological significance of lipid/fatty acid synthesis in developing eggs and newly hatched larvae of Pacific cod (Gadus macrocephalus)

The lipid/fatty acid composition of marine fish eggs and larvae is linked with buoyancy regulation, but our understanding of such processes is largely restricted to species with pelagic eggs. In this study, we examined developmental changes in the lipid/fatty acids of eggs and embryos of Pacific cod (Gadus macrocephalus), a species that spawns demersal eggs along coastal shelf edges, but as larvae must make a rapid transition to the upper reaches of the water column. Adult Pacific cod were collected in the Gulf of Alaska during the spawning season and eggs of two females were artificially fertilized with sperm from three males for each female. The eggs were subsequently reared in the laboratory to determine (1) how lipids/fatty acids were catabolized during egg and larval development, and (2) whether lipid/fatty acid catabolism had measurable effects on egg/embryo density. Eggs incubated at 4°C began hatching after 3-weeks and continued to hatch over a 10-day period, during which there was a distinct shift in lipid classes (phospholipids (PL), triacyglycerols (TAG), and sterols (ST)) and essential fatty acids (EFAs: 22:6n-3 (DHA), 20:5n-3 (EPA), and 20:4n-6 (AA)). In the egg stage, total lipid content steadily decreased during the first 60% of development, but just prior to hatch we observed an unexpected 2–3-fold lipid increase (~6–9 μg individual⁻¹) and a significant drop in egg density. The increase in lipids was largely driven by PL, with evidence of long-chained fatty acid synthesis. Late-hatching larvae had progressively decreasing lipid and fatty acid reserves, suggesting a shift from lipogenesis to lipid catabolism with continued larval development. Egg density measures suggest that lipid/fatty acid composition is linked to buoyancy regulation as larvae shift from a demersal to a pelagic existence following hatch. The biochemical pathway by which Pacific cod are apparently able to synthesize EFAs is unknown, therefore representing a remarkable finding meriting further investigation.     

Causes of habitat divergence in two species of agamid lizards in arid central Australia

The deserts of central Australia contain richer communities of lizards than any other arid regions, with the highest diversity occurring in sand dune habitats dominated by hummock-forming spinifex grasses. To investigate the mechanisms that permit coexistence, we studied two species of coexisting agamid lizards that exhibit striking divergence in their use of habitat in the Simpson Desert of central Australia. Here, the military dragon Ctenophorus isolepis is restricted primarily to sites providing > 30% cover of hard spinifex Triodia basedowii, whereas the central netted dragon C. nuchalis occurs in areas with much sparser (< 10%) cover. We constructed four mechanistic models to explain this pattern and then derived hypotheses to test them. One hypothesis, that competition restricts each species to its preferred habitat, was rejected after dyad encounters in field enclosures failed to elicit any habitat shift or any overt interactions between the species. Our next hypotheses were that each species exhibits preferences for different thermal environments or different prey types and that each selects the habitats that maximize access to them. Both were supported. C. isolepis preferred lower temperatures when active and specialized in eating ants < 5 mm long and selected spinifex-dominated areas where these requirements were met. In contrast, C. nuchalis preferred higher temperatures and a diversity of prey, both of which were available mostly in open areas. Finally, we used plasticine models to test the hypothesis that each species faced lower risk of predation in its selected habitat. This was partly supported, as models of both species were attacked more often in the open than under spinifex cover. The results show that habitat divergence occurs along several, probably covarying, niche axes. We suggest that different levels of spinifex cover provide the template for a broad range of ecological interactions, allowing lizard species to partition biotic and abiotic resources and achieve the extraordinarily high levels of local diversity that are observed.