Harnessing island-ocean connections to maximize marine benefits of island conservation.

Islands support unique plants, animals, and human societies found nowhere else on the Earth. Local and global stressors threaten the persistence of island ecosystems, with invasive species being among the most damaging, yet solvable, stressors. While the threat of invasive terrestrial mammals on island flora and fauna is well recognized, recent studies have begun to illustrate their extended and destructive impacts on adjacent marine environments. Eradication of invasive mammals and restoration of native biota are promising tools to address both island and ocean management goals. The magnitude of the marine benefits of island restoration, however, is unlikely to be consistent across the globe. We propose a list of six environmental characteristics most likely to affect the strength of land-sea linkages: precipitation, elevation, vegetation cover, soil hydrology, oceanographic productivity, and wave energy. Global databases allow for the calculation of comparable metrics describing each environmental character across islands. Such metrics can be used today to evaluate relative potential for coupled land-sea conservation efforts and, with sustained investment in monitoring on land and sea, can be used in the future to refine science-based planning tools for integrated land-sea management. As conservation practitioners work to address the effects of climate change, ocean stressors, and biodiversity crises, it is essential that we maximize returns from our management investments. Linking efforts on land, including eradication of island invasive mammals, with marine restoration and protection should offer multiplied benefits to achieve concurrent global conservation goals.

Evidence of Biological Self-Organization in Spatial Patterns of a Common Tropical Alga.

AbstractTropical reef communities contain spatial patterns at multiple scales, observable from microscope and satellite alike. Many of the smaller-scale patterns are generated physiologically (e.g., skeletal structures of corals at <1-m scale), while some of the larger patterns have been attributed to scale-dependent feedbacks (e.g., spur and groove reefs at 10-100-m scales). In describing the spatial patterning of reef benthic communities at landscape levels, we uncovered unique spatial patterning among living marine algae. Populations of the calcifying green alga Halimeda were observed to form a consistent polygonal pattern at a characteristic scale of 3-4 m. The pattern showed no clear evidence of having been formed through biologically created shifts in hydrodynamical conditions or related mechanisms. In considering the specifics of Halimeda growth patterns, a model of self-organization involving separation and patterned extension is proposed, a mechanism revealed in some geological pattern formation. This observation reinforces the diversity of pathways by which striking spatial patterns can occur in ecosystems.

Life history mediates the association between parasite abundance and geographic features.

Although parasites are ubiquitous in marine ecosystems, predicting the abundance of parasites present within marine ecosystems has proven challenging due to the unknown effects of multiple interacting environmental gradients and stressors. Furthermore, parasites often are considered as a uniform group within ecosystems despite their significant diversity. We aim to determine the potential importance of multiple predictors of parasite abundance in coral reef ecosystems, including reef area, island area, human population density, chlorophyll-a, host diversity, coral cover, host abundance and island isolation. Using a model selection approach within a database of more than 1,200 individual fish hosts and their parasites from 11 islands within the Pacific Line Islands archipelago, we reveal that geographic gradients, including island area and island isolation, emerged as the best predictors of parasite abundance. Life history moderated the relationship; parasites with complex life cycles increased in abundance with increasing island isolation, while parasites with direct life cycles decreased with increasing isolation. Direct life cycle parasites increased in abundance with increasing island area, although complex life cycle parasite abundance was not associated with island area. This novel analysis of a unique dataset indicates that parasite abundance in marine systems cannot be predicted precisely without accounting for the independent and interactive effects of each parasite's life history and environmental conditions.

Emerging insights on effects of sharks and other top predators on coral reefs.

Predation is ubiquitous on coral reefs. Among the most charismatic group of reef predators are the top predatory fishes, including sharks and large-bodied bony fishes. Despite the threat presented by top predators, data describing their realized effects on reef community structure and functioning are challenging to produce. Many innovative studies have capitalized on natural experimental conditions to explore predator effects on reefs. Gradients in predator density have been created by spatial patterning of fisheries management. Evidence of prey release has been observed across some reefs, namely that potential prey increase in density when predator density is reduced. While such studies search for evidence of prey release among broad groups or guilds of potential prey, a subset of studies have sought evidence of release at finer population levels. We find that some groups of fishes are particularly vulnerable to the effects of predators and more able to capitalize demographically when predator density is reduced. For example, territorial damselfish appear to realize reliable population expansion with the reduction in predator density, likely because their aggressive, defensive behavior makes them distinctly vulnerable to predation. Relatedly, individual fishes that suffer from debilitating conditions, such as heavy parasite loads, appear to realize relatively stronger levels of prey release with reduced predator density. Studying the effects of predators on coral reefs remains a timely pursuit, and we argue that efforts to focus on the specifics of vulnerability to predation among potential prey and other context-specific dimensions of mortality hold promise to expand our knowledge.

Considering the rates of growth in two taxa of coral across Pacific islands.

Reef-building coral taxa demonstrate considerable flexibility and diversity in reproduction and growth mechanisms. Corals take advantage of this flexibility to increase or decrease size through clonal expansion and loss of live tissue area (i.e. via reproduction and mortality of constituent polyps). The biological lability of reef-building corals may be expected to map onto varying patterns of demography across environmental contexts which can contribute to geographic variation in population dynamics. Here we explore the patterns of growth of two common coral taxa, corymbose Pocillopora and massive Porites, across seven islands in the central and south Pacific. The islands span a natural gradient of environmental conditions, including a range of pelagic primary production, a metric linked to the relative availability of inorganic nutrients and heterotrophic resources for mixotrophic corals, and sea surface temperature and thermal histories. Over a multi-year sampling interval, most coral colonies experienced positive growth (greater planar area of live tissue in second relative to first time point), though the distributions of growth varied across islands. Island-level median growth did not relate simply to estimated pelagic primary productivity or temperature. However, at locations that experienced an extreme warm-water event during the sampling interval, most Porites colonies experienced net losses of live tissue and nearly all Pocillopora colonies experienced complete mortality. While descriptive statistics of demographics offer valuable insights into trends and variability in colony change through time, simplified models predicting growth patterns based on summarized oceanographic metrics appear inadequate for robust demographic prediction. We propose that the complexity of life history strategies among colonial reef-building corals introduces unique demographic flexibility for colonies to respond to a wide breadth of environmental conditions.

Decoding diversity in a coral reef fish species complex with restricted range using metagenomic sequencing of gut contents.

AIM: Identification of the processes that generate and maintain species diversity within the same region can provide insight into biogeographic patterns at broader spatiotemporal scales. Hawkfishes in the genus Paracirrhites are a unique taxon to explore with respect to niche differentiation, exhibiting diagnostic differences in coloration, and an apparent center of distribution outside of the Indo-Malay-Philippine (IMP) biodiversity hotspot for coral reef fishes. Our aim is to use next-generation sequencing methods to leverage samples of a taxon at their center of maximum diversity to explore phylogenetic relationships and a possible mechanism of coexistence. LOCATION: Flint Island, Southern Line Islands, Republic of Kiribati. METHODS: A comprehensive review of museum records, the primary literature, and unpublished field survey records was undertaken to determine ranges for four "arc-eye" hawkfish species in the Paracirrhites species complex and a potential hybrid. Fish from four Paracirrhites species were collected from Flint Island in the Southern Line Islands, Republic of Kiribati. Hindgut contents were sequenced, and subsequent metagenomic analyses were used to assess the phylogenetic relatedness of the host fish, the microbiome community structure, and prey remains for each species. RESULTS: Phylogenetic analyses conducted with recovered mitochondrial genomes revealed clustering of P. bicolor with P. arcatus and P. xanthus with P. nisus, which were unexpected on the basis of previous morphological work in this species complex. Differences in taxonomic composition of gut microbial communities and presumed prey remains indicate likely separation of foraging niches. MAIN CONCLUSIONS: Our findings point toward previously unidentified relationships in this cryptic species complex at its proposed center of distribution. The three species endemic to the Polynesian province (P. nisus, P. xanthus, and P. bicolor) cluster separately from the more broadly distributed P. arcatus on the basis of relative abundance of metazoan sequences in the gut (presumed prey remains). Discordance between gut microbial communities and phylogeny of the host fish further reinforce the hypothesis of niche separation.

Ecophysiology of coral reef primary producers across an upwelling gradient in the tropical central Pacific.

Upwelling is an important source of inorganic nutrients in marine systems, yet little is known about how gradients in upwelling affect primary producers on coral reefs. The Southern Line Islands span a natural gradient of inorganic nutrient concentrations across the equatorial upwelling region in the central Pacific. We used this gradient to test the hypothesis that benthic autotroph ecophysiology is enhanced on nutrient-enriched reefs. We measured metabolism and photophysiology of common benthic taxa, including the algae Porolithon, Avrainvillea, and Halimeda, and the corals Pocillopora and Montipora. We found that temperature (27.2-28.7°C) was inversely related to dissolved inorganic nitrogen (0.46-4.63 µM) and surface chlorophyll a concentrations (0.108-0.147 mg m-3), which increased near the equator. Contrary to our prediction, ecophysiology did not consistently track these patterns in all taxa. Though metabolic rates were generally variable, Porolithon and Avrainvillea photosynthesis was highest at the most productive and equatorial island (northernmost). Porolithon photosynthetic rates also generally increased with proximity to the equator. Photophysiology (maximum quantum yield) increased near the equator and was highest at northern islands in all taxa. Photosynthetic pigments also were variable, but chlorophyll a and carotenoids in Avrainvillea and Montipora were highest at the northern islands. Phycobilin pigments of Porolithon responded most consistently across the upwelling gradient, with higher phycoerythrin concentrations closer to the equator. Our findings demonstrate that the effects of in situ nutrient enrichment on benthic autotrophs may be more complex than laboratory experiments indicate. While upwelling is an important feature in some reef ecosystems, ancillary factors may regulate the associated consequences of nutrient enrichment on benthic reef organisms.

Foraging consistency of coral reef fishes across environmental gradients in the central Pacific.

We take advantage of a natural gradient of human exploitation and oceanic primary production across five central Pacific coral reefs to examine foraging patterns in common coral reef fishes. Using stomach content and stable isotope (δ15N and δ13C) analyses, we examined consistency across islands in estimated foraging patterns. Surprisingly, species within the piscivore-invertivore group exhibited the clearest pattern of foraging consistency across all five islands despite there being a considerable difference in mean body mass (14 g-1.4 kg) and prey size (0.03-3.8 g). In contrast, the diets and isotopic values of the grazer-detritivores varied considerably and exhibited no consistent patterns across islands. When examining foraging patterns across environmental contexts, we found that δ15N values of species of piscivore-invertivore and planktivore closely tracked gradients in oceanic primary production; again, no comparable patterns existed for the grazer-detritivores. The inter-island consistency in foraging patterns within the species of piscivore-invertivore and planktivore and the lack of consistency among species of grazer-detritivores suggests a linkage to different sources of primary production among reef fish functional groups. Our findings suggest that piscivore-invertivores and planktivores are likely linked to well-mixed and isotopically constrained allochthonous oceanic primary production, while grazer-detritivores are likely linked to sources of benthic primary production and autochthonous recycling. Further, our findings suggest that species of piscivore-invertivore, independent of body size, converge toward consuming low trophic level prey, with a hypothesized result of reducing the number of steps between trophic levels and increasing the trophic efficiency at a community level.

Gradients in Primary Production Predict Trophic Strategies of Mixotrophic Corals across Spatial Scales.

Mixotrophy is among the most successful nutritional strategies in terrestrial and marine ecosystems. The ability of organisms to supplement primary nutritional modes along continua of autotrophy and heterotrophy fosters trophic flexibility that can sustain metabolic demands under variable or stressful conditions. Symbiotic, reef-building corals are among the most broadly distributed and ecologically important mixotrophs, yet we lack a basic understanding of how they modify their use of autotrophy and heterotrophy across gradients of food availability. Here, we evaluate how one coral species, Pocillopora meandrina, supplements autotrophic nutrition through heterotrophy within an archipelago and test whether this pattern holds across species globally. Using stable isotope analysis (δ13C) and satellite-derived estimates of nearshore primary production (chlorophyll-a, as a proxy for food availability), we show that P. meandrina incorporates a greater proportion of carbon via heterotrophy when more food is available across five central Pacific islands. We then show that this pattern is consistent globally using data from 15 coral species across 16 locations spanning the Caribbean Sea and the Indian and Pacific Oceans. Globally, surface chlorophyll-a explains 77% of the variation in coral heterotrophic nutrition, 86% for one genus across 10 islands, and 94% when controlling for coral taxonomy within archipelagos. These results demonstrate, for the first time, that satellite-derived estimates of nearshore primary production provide a globally relevant proxy for resource availability that can explain variation in coral trophic ecology. Thus, our model provides a pivotal step toward resolving the biophysical couplings between mixotrophic organisms and spatial patterns of resource availability in the coastal oceans.

Human impacts decouple a fundamental ecological relationship-The positive association between host diversity and parasite diversity.

Human impacts on ecosystems can decouple the fundamental ecological relationships that create patterns of diversity in free-living species. Despite the abundance, ubiquity, and ecological importance of parasites, it is unknown whether the same decoupling effects occur for parasitic species. We investigated the influence of fishing on the relationship between host diversity and parasite diversity for parasites of coral reef fishes on three fished and three unfished islands in the central equatorial Pacific. Fishing was associated with a shallowing of the positive host-diversity-parasite-diversity relationship. This occurred primarily through negative impacts of fishing on the presence of complex life-cycle parasites, which created a biologically impoverished parasite fauna of directly transmitted parasites resilient to changes in host biodiversity. Parasite diversity appears to be decoupled from host diversity by fishing impacts in this coral reef ecosystem, which suggests that such decoupling might also occur for parasites in other ecosystems affected by environmental change.

Reef Fish Survey Techniques: Assessing the Potential for Standardizing Methodologies.

Dramatic changes in populations of fishes living on coral reefs have been documented globally and, in response, the research community has initiated efforts to assess and monitor reef fish assemblages. A variety of visual census techniques are employed, however results are often incomparable due to differential methodological performance. Although comparability of data may promote improved assessment of fish populations, and thus management of often critically important nearshore fisheries, to date no standardized and agreed-upon survey method has emerged. This study describes the use of methods across the research community and identifies potential drivers of method selection. An online survey was distributed to researchers from academic, governmental, and non-governmental organizations internationally. Although many methods were identified, 89% of survey-based projects employed one of three methods-belt transect, stationary point count, and some variation of the timed swim method. The selection of survey method was independent of the research design (i.e., assessment goal) and region of study, but was related to the researcher's home institution. While some researchers expressed willingness to modify their current survey protocols to more standardized protocols (76%), their willingness decreased when methodologies were tied to long-term datasets spanning five or more years. Willingness to modify current methodologies was also less common among academic researchers than resource managers. By understanding both the current application of methods and the reported motivations for method selection, we hope to focus discussions towards increasing the comparability of quantitative reef fish survey data.

Productivity and fishing pressure drive variability in fish parasite assemblages of the Line Islands, equatorial Pacific.

Variability in primary productivity and fishing pressure can shape the abundance, species composition, and diversity of marine life. Though parasites comprise nearly half of marine species, their responses to these important forces remain little explored. We quantified parasite assemblages at two spatial scales, across a gradient in productivity and fishing pressure that spans six coral islands of the Line Islands archipelago and within the largest Line Island, Kiritimati, which experiences a west-to-east gradient in fishing pressure and upwelling-driven productivity. In the across-islands data set, we found that increasing productivity was correlated with increased parasite abundance overall, but that the effects of productivity differed among parasite groups. Trophically transmitted parasites increased in abundance with increasing productivity, but directly transmitted parasites did not exhibit significant changes. This probably arises because productivity has stronger effects on the abundance of the planktonic crustaceans and herbivorous snails that serve as the intermediate hosts of trophically transmitted parasites than on the higher-trophic level fishes that are the sole hosts of directly transmitted parasites. We also found that specialist parasites increased in response to increasing productivity, while generalists did not, possibly because specialist parasites tend to be more strongly limited by host availability than are generalist parasites. After the effect of productivity was controlled for, fishing was correlated with decreases in the abundance of trophically transmitted parasites, while directly transmitted parasites appeared to track host density; we observed increases in the abundance of parasites using hosts that experienced fishing-driven compensatory increases in abundance. The within-island data set confirmed these patterns for the combined effects of productivity and fishing on parasite abundance, suggesting that our conclusions are robust across a span of spatial scales. Overall, these results indicate that there are strong and variable effects of anthropogenic and natural drivers on parasite abundance and taxonomic richness. These effects are likely to be mediated by parasite traits, particularly by parasite transmission strategies.

Spawning aggregation behavior and reproductive ecology of the giant bumphead parrotfish, Bolbometopon muricatum, in a remote marine reserve.

The giant bumphead parrotfish (Bolbometopon muricatum) has experienced precipitous population declines throughout its range due to its importance as a highly-prized fishery target and cultural resource. Because of its diet, Bolbometopon may serve as a keystone species on Indo-Pacific coral reefs, yet comprehensive descriptions of its reproductive ecology do not exist. We used a variety of underwater visual census (UVC) methods to study an intact population of Bolbometopon at Wake Atoll, a remote and protected coral atoll in the west Pacific. Key observations include spawning activities in the morning around the full and last quarter moon, with possible spawning extending to the new moon. We observed peaks in aggregation size just prior to and following the full and last quarter moon, respectively, and observed a distinct break in spawning at the site that persisted for four days; individuals returned to the aggregation site one day prior to the last quarter moon and resumed spawning the following day. The mating system was lek-based, characterized by early male arrival at the spawning site followed by vigorous defense (including head-butting between large males) of small territories. These territories were apparently used to attract females that arrived later in large schools, causing substantial changes in the sex ratio on the aggregation site at any given time during the morning spawning period. Aggression between males and courtship of females led to pair spawning within the upper water column. Mating interference was not witnessed but we noted instances suggesting that sperm competition might occur. Densities of Bolbometopon on the aggregation site averaged 10.07(±3.24 SE) fish per hectare (ha) with maximum densities of 51.5 fish per ha. By comparing our observations to the results of biennial surveys conducted by the National Oceanic and Atmospheric Administration (NOAA) Coral Reef Ecosystem Division (CRED), we confirmed spatial consistency of the aggregation across years as well as a temporal break in spawning activity and aggregation that occurred during the lunar phase. We estimated the area encompassed by the spawning aggregation to be 0.72 ha, suggesting that spawning site closures and temporal closures centered around the full to the new moon might form one component of a management and conservation plan for this species. Our study of the mating system and spawning aggregation behavior of Bolbometopon from the protected, relatively pristine population at Wake Atoll provides crucial baselines of population density, sex ratio composition, and productivity of a spawning aggregation site from an oceanic atoll. Such information is key for conservation efforts and provides a basic platform for the design of marine protected areas for this threatened iconic coral reef fish, as well as for species with similar ecological and life history characteristics.

The shallow-water fish assemblage of Isla del Coco National Park, Costa Rica: structure and patterns in an isolated, predator-dominated ecosystem / El conjunto de peces de aguas poco profundas del Parque Nacional Isla del Coco, Costa Rica: estructura y patrones en un ecosistema aislado dominado por depredadores

Fishes at Isla del Coco National Park, Costa Rica, were surveyed as part of a larger scientific expedition to the area in September 2009. The average total biomass of nearshore fishes was 7.8 tonnes per ha, among the largest observed in the tropics, with apex predators such as sharks, jacks, and groupers accounting for nearly 40% of the total biomass. The abundance of reef and pelagic sharks, particularly large aggregations of threatened species such as the scalloped hammerhead shark (up to 42 hammerheads ha-1) and large schools of jacks and snappers show the capacity for high biomass in unfished ecosystems in the Eastern Tropical Pacific. However, the abundance of hammerhead and reef whitetip sharks appears to have been declining since the late 1990s, and likely causes may include increasing fishing pressure on sharks in the region and illegal fishing inside the Park. One Galapagos shark tagged on September 20, 2009 in the Isla del Coco National Park moved 255km southeast towards Malpelo Island in Colombia, when it stopped transmitting. These results contribute to the evidence that sharks conduct large-scale movements between marine protected areas (Isla del Coco, Malpelo, Galápagos) in the Eastern tropical Pacific and emphasize the need for regional-scale management. More than half of the species and 90% of the individuals observed were endemic to the tropical eastern Pacific. These high biomass and endemicity values highlight the uniqueness of the fish assemblage at Isla del Coco and its importance as a global biodiversity hotspot.

La biomasa promedio de peces costeros en el Parque Nacional Isla del Coco en septiembre de 2010 fue de 7,8 toneladas por hectárea, entre las más elevadas halladas jamás en zonas tropicales. Los grandes depredadores representaron el 40% de la biomasa total. La abundancia de tiburones costeros y pelágicos, particularmente las enormes agregaciones de tiburón martillo (hasta 42 individuos por hectárea) y los extensos bancos de carángidos y lutjánidos, muestran la capacidad que tienen los ecosistemas marinos no pescados para albergar elevadas biomasas de peces, y hacen de la Isla del Coco un lugar único en el mundo. No obstante, la abundancia de tiburones parece estar decreciendo desde 1999, probablemente a causa de la creciente presión pesquera en la región y la pesca ilegal en el interior del Parque. Un tiburón de Galápagos marcado se dirigió 255km en dirección a la Isla de Malpelo, Colombia. Estos resultados sugieren que los tiburones realizan importantes movimientos entre áreas marinas protegidas (Isla del Coco, Malpelo, Galápagos) en el Pacífico Tropical Oriental y remarcan la necesidad de una gestión regional de estos animales. Más del 50% de las especies y el 90% de individuos observados en los contajes eran endémicos del Pacífico Tropical Oriental.

Assessing coral reefs on a Pacific-wide scale using the microbialization score.

The majority of the world's coral reefs are in various stages of decline. While a suite of disturbances (overfishing, eutrophication, and global climate change) have been identified, the mechanism(s) of reef system decline remain elusive. Increased microbial and viral loading with higher percentages of opportunistic and specific microbial pathogens have been identified as potentially unifying features of coral reefs in decline. Due to their relative size and high per cell activity, a small change in microbial biomass may signal a large reallocation of available energy in an ecosystem; that is the microbialization of the coral reef. Our hypothesis was that human activities alter the energy budget of the reef system, specifically by altering the allocation of metabolic energy between microbes and macrobes. To determine if this is occurring on a regional scale, we calculated the basal metabolic rates for the fish and microbial communities at 99 sites on twenty-nine coral islands throughout the Pacific Ocean using previously established scaling relationships. From these metabolic rate predictions, we derived a new metric for assessing and comparing reef health called the microbialization score. The microbialization score represents the percentage of the combined fish and microbial predicted metabolic rate that is microbial. Our results demonstrate a strong positive correlation between reef microbialization scores and human impact. In contrast, microbialization scores did not significantly correlate with ocean net primary production, local chla concentrations, or the combined metabolic rate of the fish and microbial communities. These findings support the hypothesis that human activities are shifting energy to the microbes, at the expense of the macrobes. Regardless of oceanographic context, the microbialization score is a powerful metric for assessing the level of human impact a reef system is experiencing.

Extraordinary aggressive behavior from the giant coral reef fish, Bolbometopon muricatum, in a remote marine reserve.

Human impacts to terrestrial and marine communities are widespread and typically begin with the local extirpation of large-bodied animals. In the marine environment, few pristine areas relatively free of human impact remain to provide baselines of ecosystem function and goals for restoration efforts. Recent comparisons of remote and/or protected coral reefs versus impacted sites suggest remote systems are dominated by apex predators, yet in these systems the ecological role of non-predatory, large-bodied, highly vulnerable species such as the giant bumphead parrotfish (Bolbometopon muricatum) has received less attention. Overfishing of Bolbometopon has lead to precipitous declines in population density and avoidance of humans throughout its range, contributing to its status as a candidate species under the U. S. Endangered Species Act and limiting opportunities to study unexploited populations. Here we show that extraordinary ecological processes, such as violent headbutting contests by the world's largest parrotfish, can be revealed by studying unexploited ecosystems, such as the coral reefs of Wake Atoll where we studied an abundant population of Bolbometopon. Bolbometopon is among the largest of coral reef fishes and is a well known, charismatic species, yet to our knowledge, no scientific documentation of ritualized headbutting exists for marine fishes. Our observations of aggressive headbutting by Bolbometopon underscore that remote locations and marine reserves, by inhibiting negative responses to human observers and by allowing the persistence of historical conditions, can provide valuable opportunities to study ecosystems in their natural state, thereby facilitating the discovery, conservation, and interpretation of a range of sometimes remarkable behavioral and ecological processes.

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.