Comparing fishers' and scientific estimates of size at maturity and maximum body size as indicators for overfishing.

We tested whether fishers' local ecological knowledge (LEK) of two fish life-history parameters, size at maturity (SAM) at maximum body size (MS), was comparable to scientific estimates (SEK) of the same parameters, and whether LEK influenced fishers' perceptions of sustainability. Local ecological knowledge was documented for 82 fishers from a small-scale fishery in Samaná Bay, Dominican Republic, whereas SEK was compiled from the scientific literature. Size at maturity estimates derived from LEK and SEK overlapped for most of the 15 commonly harvested species (10 of 15). In contrast, fishers' maximum size estimates were usually lower than (eight species), or overlapped with (five species) scientific estimates. Fishers' size-based estimates of catch composition indicate greater potential for overfishing than estimates based on SEK. Fishers' estimates of size at capture relative to size at maturity suggest routine inclusion of juveniles in the catch (9 of 15 species), and fishers' estimates suggest that harvested fish are substantially smaller than maximum body size for most species (11 of 15 species). Scientific estimates also suggest that harvested fish are generally smaller than maximum body size (13 of 15), but suggest that the catch is dominated by adults for most species (9 of 15 species), and that juveniles are present in the catch for fewer species (6 of 15). Most Samaná fishers characterized the current state of their fishery as poor (73%) and as having changed for the worse over the past 20 yr (60%). Fishers stated that concern about overfishing, catching small fish, and catching immature fish contributed to these perceptions, indicating a possible influence of catch-size composition on their perceptions. Future work should test this link more explicitly because we found no evidence that the minority of fishers with more positive perceptions of their fishery reported systematically different estimates of catch-size composition than those with the more negative majority view. Although fishers' and scientific estimates of size at maturity and maximum size parameters sometimes differed, the fact that fishers make routine quantitative assessments of maturity and body size suggests potential for future collaborative monitoring efforts to generate estimates usable by scientists and meaningful to fishers.

Evaluating how variants of floristic quality assessment indicate wetland condition.

Biological indicators are useful tools for the assessment of ecosystem condition. Multi-metric and multi-taxa indicators may respond to a broader range of disturbances than simpler indicators, but their complexity can make them difficult to interpret, which is critical to indicator utility for ecosystem management. Floristic Quality Assessment (FQA) is an example of a biological assessment approach that has been widely tested for indicating freshwater wetland condition, but less attention has been given to clarifying the factors controlling its response. FQA quantifies the aggregate of vascular plant species tolerance to habitat degradation (conservatism), and model variants have incorporated species richness, abundance, and indigenity (native or non-native). To assess bias, we tested FQA variants in open-canopy freshwater wetlands against three independent reference measures, using practical vegetation sampling methods. FQA variants incorporating species richness did not correlate with our reference measures and were influenced by wetland size and hydrogeomorphic class. In contrast, FQA variants lacking measures of species richness responded linearly to reference measures quantifying individual and aggregate stresses, suggesting a broad response to cumulative degradation. FQA variants incorporating non-native species, and a variant additionally incorporating relative species abundance, improved performance over using only native species. We relate our empirical findings to ecological theory to clarify the functional properties and implications of the FQA variants. Our analysis indicates that (1) aggregate conservatism reliably declines with increased disturbance; (2) species richness has varying relationships with disturbance and increases with site area, confounding FQA response; and (3) non-native species signal human disturbance. We propose that incorporating species abundance can improve FQA site-level relevance with little extra sampling effort. Using our practical sampling methods, an FQA variant ignoring species richness and incorporating non-native species and relative species abundance can be logistically efficient, easily understood, and effective for wetland assessment.

High population density enhances recruitment and survival of a harvested coral reef fish.

A negative relationship between population growth and population density (direct density dependence) is necessary for population regulation and is assumed in most models of harvested populations. Experimental tests for density dependence are lacking for large-bodied, harvested fish because of the difficulty of manipulating population density over large areas. We studied a harvested coral reef fish, Lutjanus apodus (schoolmaster snapper), using eight large, isolated natural reefs (0.4-1.6 ha) in the Bahamas as replicates. An initial observational test for density dependence was followed by a manipulation of population density. The manipulation weakened an association between density and shelter-providing habitat features and revealed a positive effect of population density on recruitment and survival (inverse density dependence), but no effect of density on somatic growth. The snappers on an individual reef were organized into a few shoals, and we hypothesize that large shoals on high-density reefs were less vulnerable to large piscivores (groupers and barracudas) than the small shoals on low-density reefs. Reductions in predation risk for individuals in large social groups are well documented, but because snapper shoals occupied reefs the size of small marine reserves, these ecological interactions may influence the outcome of management actions.

Marine reserves as linked social-ecological systems.

Marine reserves are increasingly recognized as having linked social and ecological dynamics. This study investigates how the ecological performance of 56 marine reserves throughout the Philippines, Caribbean, and Western Indian Ocean (WIO) is related to both reserve design features and the socioeconomic characteristics in associated coastal communities. Ecological performance was measured as fish biomass in the reserve relative to nearby areas. Of the socioeconomic variables considered, human population density and compliance with reserve rules had the strongest effects on fish biomass, but the effects of these variables were region specific. Relationships between population density and the reserve effect on fish biomass were negative in the Caribbean, positive in the WIO, and not detectable in the Philippines. Differing associations between population density and reserve effectiveness defy simple explanation but may depend on human migration to effective reserves, depletion of fish stocks outside reserves, or other social factors that change with population density. Higher levels of compliance reported by resource users was related to higher fish biomass in reserves compared with outside, but this relationship was only statistically significant in the Caribbean. A heuristic model based on correlations between social, cultural, political, economic, and other contextual conditions in 127 marine reserves showed that high levels of compliance with reserve rules were related to complex social interactions rather than simply to enforcement of reserve rules. Comparative research of this type is important for uncovering the complexities surrounding human dimensions of marine reserves and improving reserve management.

Host-parasite Interactions between a Copepod (Pharodes tortugensis) and Small Reef-associated Gobies (Coryphopterus) in the British Virgin Islands.

The effects of parasitic copepods on free-living hosts are infrequently documented, and the copepod Pharodes tortugensis has remained virtually unstudied since described. For the first time, we document its host range in the British Virgin Islands (BVI), the prevalence and intensity of infections on wild hosts, and its impacts on host morphology and performance. Infections were observed on four benthic gobies in the BVI (Coryphopterus glaucofraenum, C. venezuelae, C. dicrus and C. eidolon) but not on other host species previously reported from other parts of the western Atlantic. Infected gobies were widespread in the BVI (detected at 33 of 52 sites, prevalence from 1-25%) but extremely rare elsewhere in the Caribbean (detected at 2 of 16 sites, prevalence < 0.006%). As is typical of macroparasite infections, P. tortugensis was over-dispersed in BVI host populations (mean intensity = 4.7, range = 1-17). Infections were most common in juvenile and female hosts, and rarely found in larger male hosts. The copepods attach in the branchial chamber of the goby; female copepods show high attachment fidelity to the ventral surface of the chamber, while male copepods attached most often to the first two gill arches and in the branchial chamber adjacent to the female. Infections caused substantial damage to the host's branchial chamber and gill filaments. Parasitized gobies also had larger livers and smaller gonads than unparasitized individuals of similar length. The changes in organ mass of infected gobies were not sizeable enough to affect total body mass, and host condition (the body-length vs. body-mass relationship) was similar for gobies with and without infections. Parasitized gobies were, however, significantly smaller in body mass at a given age, reflecting slower overall growth. Effects of P. tortugensis on individual hosts were broadly similar to those of other parasitic copepods that infect fish gills and, for unknown reasons, the BVI appears to be a persistent hotspot of infections on these goby hosts.

Using DNA barcoding to identify host-parasite interactions between cryptic species of goby (Coryphopterus: Gobiidae, Perciformes) and parasitic copepods (Pharodes tortugensis: Chondracanthidae, Cyclopoida).

Previous work, using morphological characters, identified a generalist copepod parasite (Pharodes tortugensis) at high prevalence on two common gobies (Coryphopterus glaucofraenum and C. dicrus) in the British Virgin Islands (BVI). DNA barcoding subsequently revealed C. glaucofraenum to be three morphologically similar species (C. glaucofraenum, C. venezuelae and C. tortugae), casting doubt on host identities in the BVI and the classification of the parasite as a single species. Mitochondrial cytochrome c oxidase subunit I (COI) data from 67 gobies in the BVI showed that, in addition to C. dicrus, host gobies were a mix of C. glaucofraenum and C. venezuelae, while C. tortugae was unexpectedly absent from the study area. COI data (n = 70) indicated that the copepod infecting all three hosts was a single species, almost certainly P. tortugensis. The pharodes-coryphopterus interaction has a strong impact on host dynamics in the BVI, and a revised understanding of these dynamics must account for any differences among the three newly confirmed hosts in transmission of, and susceptibility to, the shared parasite. No other infected hosts were discovered at our sites, but P. tortugensis is reportedly widespread and infects 12 additional host species elsewhere. Further DNA barcoding is thus needed to test whether P. tortugensis is truly a widespread generalist, or instead represents a group of more specialized cryptic species.

Inter-cohort competition drives density dependence and selective mortality in a marine fish.

For organisms with complex life cycles, the transition between life stages and between habitats can act as a significant demographic and selective bottleneck. In particular, competition with older and larger conspecifics and heterospecifics may influence the number and characteristics of individuals successfully making the transition. We investigated whether the availability of enemy-free space mediated the interaction between adult goldspot gobies (Gnatholepis thompsoni), a common tropical reef fish, and juvenile conspecifics that had recently settled from the plankton. We added rocks, which provide refuge from predators, to one-half of each of five entire coral reefs in the Bahamas and measured the survival and growth of recent settlers in relation to adult goby densities. We also evaluated whether mortality was selective with respect to three larval traits (age at settlement, size at settlement, and presettlement growth rate) and measured the influence of refuge availability and adult goby density on selection intensity. Selective mortality was measured by comparing larval traits of newly settled gobies (< or = 5 d postsettlement) with those of survivors (2-3 week postsettlement juveniles). We detected a negative relationship between juvenile survival and adult goby density in both low- and high-refuge habitats, though experimental refuge addition reduced the intensity of this density dependence. Juvenile growth also declined with increasing adult goby density, but this effect was similar in both low- and high-refuge habitats. Refuge availability had no consistent effect on selective mortality, but adult goby density was significantly related to the intensity of size-selective mortality: bigger juveniles were favored where adults were abundant, and smaller juveniles were favored where adults were rare. Given the typically large difference in sizes of juveniles and adults, similar stage-structured interactions may be common but underappreciated in many marine species.

Musical chairs mortality functions: density-dependent deaths caused by competition for unguarded refuges.

Structural refuges within which prey can escape from predators can be an important limiting resource for the prey. In a manner that resembles the childhood game of musical chairs, many prey species rapidly retreat to shared, unguarded refuges whenever a predator threatens, and only when refuges are relatively abundant do all prey individuals actually escape. The key feature of this process is that the per capita prey mortality rate depends on the ratio of prey individuals to refuges. We introduce a new class of mortality functions with this feature and then demonstrate statistically that they describe field mortality data from a well-studied coral reef fish species, the Caribbean bridled goby Coryphopterus glaucofraenum, substantially better than do several mortality functions of more conventional form.

Boat anchoring contributes substantially to coral reef degradation in the British Virgin Islands.

Community decline is often linked to anthropogenic activities. Coral reef declines, for example, have been linked to overfishing and climate change, but impacts of coastal development, pollution, and tourism have received increasing attention. Here, we isolated the impact of one little-studied aspect of recreational activity on coral reefs-damage from boat anchoring-by performing a survey of 24 sites in the British Virgin Islands (BVI) subject to varying levels of anchoring activity. The percent cover of hard corals and sea fans was reduced by a factor of ∼1.7 and ∼2.6 respectively at highly anchored sites. Hard coral colonies were  40% smaller in surface area and ∼60% less dense at sites experiencing high anchoring frequency. In addition, highly anchored sites supported only ∼60% of the species richness of little anchored sites. Frequently anchored sites were ∼60% as structurally complex and supported less than half as many fish as those rarely anchored, with 5 of 7 fish functional groups affected. Roughly 24% of BVI coral reef by area appears suitable for anchoring, suggesting that impacts associated with boat anchoring may be both locally severe and more pervasive than previously appreciated.

Behavioural mechanisms underlying parasite-mediated competition for refuges in a coral reef fish.

Parasites have been increasingly recognized as participants in indirect ecological interactions, including those mediated by parasite-induced changes to host behaviour (trait-mediated indirect interactions or TMIIs). In most documented examples, host behaviours altered by parasites increase susceptibility to predation because the predator is also a host (host-manipulation). Here, we test for a TMII in which a parasitic copepod modifies the predator-prey interaction between a small goby host and several larger predatory fish. Gobies compete for crevices in the reef to avoid predation and goby mortality increases more rapidly with increasing refuge shortage for parasitized gobies than for those free of parasites. We found interactive effects of refuge shortage and parasitism on two behaviours we predicted might be associated with parasite-mediated competition for refuges. First, as refuge-shortage increases, the rate of aggression among gobies increases and parasitism intensifies this interaction. Second, goby proximity to refuges increases as refuges become scarce, but parasitism nullifies this increase. In combination, these parasite-induced changes in behaviour may explain why parasitized gobies are poor competitors for refuges. Because the parasite is not trophically transmitted via host manipulation, these altered behaviours in parasitized gobies are likely coincidental to infection.

Parasitism and a shortage of refuges jointly mediate the strength of density dependence in a reef fish.

Various predator-prey, host-pathogen, and competitive interactions can combine to cause density dependence in population growth. Despite this possibility, most empirical tests for density-dependent interactions have focused on single mechanisms. Here we tested the hypothesis that two mechanisms of density dependence, parasitism and a shortage of refuges, jointly influence the strength of density-dependent mortality. We used mark-recapture analysis to estimate mortality of the host species, the bridled goby (Coryphopterus glaucofraenum). Sixty-three marked gobies were infected with a copepod gill parasite (Pharodes tortugensis), and 188 were uninfected. We used the spatial scale at which gobies were clustered naturally (approximately 4 m2) as an ecologically relevant neighborhood and measured goby density and the availability of refuges from predators within each goby's neighborhood. Goby survival generally declined with increasing density, and this decline was steeper for gobies with access to few refuges than for gobies in neighborhoods where refuges were common. The negative effects of high density and refuge shortage were also more severe for parasitized gobies than for gobies free of parasites. This parasite has characteristics typical of emerging diseases and appears to have altered the strength of a preexisting density-dependent interaction.

Episodic Disturbance from Boat Anchoring Is a Major Contributor to, but Does Not Alter the Trajectory of, Long-Term Coral Reef Decline.

Isolating the relative effects of episodic disturbances and chronic stressors on long-term community change is challenging. We assessed the impact of an episodic disturbance associated with human visitation (boat anchoring) relative to other drivers of long-term change on coral reefs. A one-time anchoring event at Crab Cove, British Virgin Islands, in 2004 caused rapid losses of coral and reef structural complexity that were equal to the cumulative decline over 23 years observed at an adjacent site. The abundance of small site-attached reef fishes dropped by approximately one quarter after the anchoring event, but this drop was not immediate and only fully apparent two years after the anchoring event. There was no obvious recovery from the impact, and no evidence that this episodic impact accelerated or retarded subsequent declines from other causes. This apparent lack of synergism between the effect of this episodic human impact and other chronic stressors is consistent with the few other long-term studies of episodic impacts, and suggests that action to mitigate anchor damage should yield predictable benefits.

Strong density-dependent survival and recruitment regulate the abundance of a coral reef fish.

Debate on the control of population dynamics in reef fishes has centred on whether patterns in abundance are determined by the supply of planktonic recruits, or by post-recruitment processes. Recruitment limitation implies little or no regulation of the reef-associated population, and is supported by several experimental studies that failed to detect density dependence. Previous manipulations of population density have, however, focused on juveniles, and there have been no tests for density-dependent interactions among adult reef fishes. I tested for population regulation in Coryphopterus glaucofraenum, a small, short-lived goby that is common in the Caribbean. Adult density was manipulated on artificial reefs and adults were also monitored on reefs where they varied in density naturally. Survival of adult gobies showed a strong inverse relationship with their initial density across a realistic range of densities. Individually marked gobies, however, grew at similar rates across all densities, suggesting that density-dependent survival was not associated with depressed growth, and so may result from predation or parasitism rather than from food shortage. Like adult survival, the accumulation of new recruits on reefs was also much lower at high adult densities than at low densities. Suppression of recruitment by adults may occur because adults cause either reduced larval settlement or reduced early post-settlement survival. In summary, this study has documented a previously unrecorded regulatory mechanism for reef fish populations (density-dependent adult mortality) and provided a particularly strong example of a well-established mechanism (density-dependent recruitment). In combination, these two compensatory mechanisms have the potential to strongly regulate the abundance of this species, and rule out the control of abundance by the supply of recruits.

Effects of trout on the diel periodicity of drifting in baetid mayflies.

Some benthic invertebrates in streams make frequent, short journeys downstream in the water column (=drifting). In most streams there are larger numbers of invertebrates in the drift at night than during the day. We tested the hypothesis that nocturnal drifting is a response to avoid predation from fish that feed in the water column during the day. We surveyed diel patterns of drifting by nymphs of the mayfly Baetis coelestis in several streams containing (n=5) and lacking (n=7) populations of rainbow trout, Oncorhynchus mykiss. Drifting was more nocturnal in the presence of trout (85% of daily drift occurred at night) than in their absence (50% of daily drift occurred at night). This shift in periodicity is due to reduced daytime drifting in streams with trout, because at a given nighttime drift density, the daytime drift density of B. coelestis was lower in streams occupied by trout than in troutless streams. Large size classes of B. coelestis were underrepresented in the daytime drift in trout streams compared to nighttime drift in trout streams, and to both day and night drift in troutless streams. Differences in daytime drift density between streams with and without trout were the result of differences in mayfly drift behaviour among streams because predation rates by trout were too low to significantly reduce densities of drifting B. coelestis. We tested for rapid (over 3 days) phenotypic responses to trout presence by adding trout in cages to three of the troutless streams. Nighttime drifting was unaffected by the addition of trout, but daytime drift densities were reduced by 28% below cages containing trout relative to control cages (lacking trout) placed upstream. Drift responses were measured 15 m downstream of the cages suggesting that mayflies detected trout using chemical cues. Overall, these data support the hypothesis that infrequent daytime drifting is an avoidance response to fish that feed in the water column during the day. Avoidance is more pronounced in large individuals and is, at least partially, a phenotypic response mediated by chemical cues.

Growth of estuarine fish is associated with the combined concentration of sediment contaminants and shows no adaptation or acclimation to past conditions.

We tested whether the growth rates of small benthic fish (Gillichthys mirabilis) in three southern California estuaries corresponded with the local concentrations of contaminants. Fish originating from each estuary were transplanted to cages in each estuary in two reciprocal transplant experiments. The growth rates of caged fish, and the size-distribution of natural populations, showed the same pattern of difference among estuaries. Twelve metals and organic contaminants occurred in bulk sediments at concentrations close to their individual ERL values, and a simple index of their combined concentration (the mean ERL quotient) was inversely correlated to the growth of caged fish. Metals in the water column occurred at lower concentrations, relative to toxicity thresholds, than those in sediments and were unrelated to fish growth. Fish used in the field caging experiments, and other fish held in the laboratory under constant conditions, showed no difference in growth according to their estuary of origin. Fish originating from different estuaries also showed no consistent differences in their tissue burden of organic contaminants. Our results thus suggested no genetic adaptation or physiological acclimation to the past contaminant regime, but revealed a possible association between fish growth rates and the combined concentration of multiple sediment contaminants.

Recent region-wide declines in Caribbean reef fish abundance.

Profound ecological changes are occurring on coral reefs throughout the tropics, with marked coral cover losses and concomitant algal increases, particularly in the Caribbean region. Historical declines in the abundance of large Caribbean reef fishes likely reflect centuries of overexploitation. However, effects of drastic recent degradation of reef habitats on reef fish assemblages have yet to be established. By using meta-analysis, we analyzed time series of reef fish density obtained from 48 studies that include 318 reefs across the Caribbean and span the time period 1955-2007. Our analyses show that overall reef fish density has been declining significantly for more than a decade, at rates that are consistent across all subregions of the Caribbean basin (2.7% to 6.0% loss per year) and in three of six trophic groups. Changes in fish density over the past half-century are modest relative to concurrent changes in benthic cover on Caribbean reefs. However, the recent significant decline in overall fish abundance and its consistency across several trophic groups and among both fished and nonfished species indicate that Caribbean fishes have begun to respond negatively to habitat degradation.

SPATIAL DENSITY DEPENDENCE SCALES UP BUT DOES NOT PRODUCE TEMPORAL DENSITY DEPENDENCE IN A REEF FISH.

Field experiments provide rigorous tests of ecological hypotheses but are typically of short duration and use small spatial replicates. We assessed empirically whether the results of experiments testing for density dependence applied at larger spatial domains and explained temporal population dynamics. We studied a small coral reef fish, the goldspot goby (Gnatholepis thompsoni), in the Bahamas. We assessed the effects of interactions with conspecifics and with an ecologically similar species, the bridled goby (Coryphopterus glaucofraenum). Two density manipulations on small reef patches revealed that goldspot goby mortality over one month increased as conspecifics became crowded. On five large natural reefs, we correlated the initial year-class density of both species (annual larval settlement) with the subsequent decline of goldspot goby year-classes for five years. Mortality was correlated with conspecific density among reefs for all years, but not among years for all reefs. Thus, spatial density dependence in mortality scaled up qualitatively from small patches to entire reefs but was not associated with temporal density dependence. Our results support the conclusion that field experiments may be extrapolated to larger spatial domains with care, but that using small spatial comparisons to predict temporal responses is difficult without knowing the underlying biological mechanisms.

Assessing the magnitude of intra- and interspecific competition in two coral reef fishes.

Many field experiments have tested for effects of competition in nature, but relatively few have used designs allowing simultaneous assessment of the influence of intra- and interspecific competition. Using a response surface design and a press manipulation of densities, we tested effects of competition within and between two species of coral reef fishes (Coryphopterus glaucofraneum and Gnatholepis thompsoni). By tracking individually tagged fishes, we showed that the per-capita effect of intraspecific competitors on individual growth was at least twice as great as the effect of interspecific competitors. Growth rate was better predicted by measures of density that incorporated body size, rather than numerical density, suggesting interference competition. Individuals of both species interacted aggressively with conspecifics at least twice as often as with heterospecifics. Individuals of both species also covered more area while foraging and spent less time in shelter when crowded than when at lower densities. In combination, these behaviours suggest that increased metabolic costs at high density contribute to competitive effects on growth. These competitive interactions occurred among adult fishes, so reduced growth may translate to reduced fecundity as well as reduced survival, and so contribute to population regulation.

Small-scale field experiments accurately scale up to predict density dependence in reef fish populations at large scales.

Field experiments provide rigorous tests of ecological hypotheses but are usually limited to small spatial scales. It is thus unclear whether these findings extrapolate to larger scales relevant to conservation and management. We show that the results of experiments detecting density-dependent mortality of reef fish on small habitat patches scale up to have similar effects on much larger entire reefs that are the size of small marine reserves and approach the scale at which some reef fisheries operate. We suggest that accurate scaling is due to the type of species interaction causing local density dependence and the fact that localized events can be aggregated to describe larger-scale interactions with minimal distortion. Careful extrapolation from small-scale experiments identifying species interactions and their effects should improve our ability to predict the outcomes of alternative management strategies for coral reef fishes and their habitats.