Collaborative assessment of California spiny lobster population and fishery responses to a marine reserve network

Assessments of the conservation and fisheries effects of marine reserves typically focus on single reserves where sampling occurs over narrow spatiotemporal scales. A strategy for broadening the collection and interpretation of data is collaborative fisheries research (CFR). Here we report results of a CFR program formed in part to test whether reserves at the Santa Barbara Channel Islands, USA, influenced lobster size and trap yield, and whether abundance changes in reserves led to spillover that influenced trap yield and effort distribution near reserve borders. Industry training of scientists allowed us to sample reserves with fishery relevant metrics that we compared with pre-reserve fishing records, a concurrent port sampling program, fishery effort patterns, the local ecological knowledge (LEK) of fishermen, and fishery-independent visual surveys of lobster abundance. After six years of reserve protection, there was a four- to eightfold increase in trap yield, a 5-10% increase in the mean size (carapace length) of legal sized lobsters, and larger size structure of lobsters trapped inside vs. outside of three replicate reserves. Patterns in trap data were corroborated by visual scuba surveys that indicated a four- to sixfold increase in lobster density inside reserves. Population increases within reserves did not lead to increased trap yields or effort concentrations (fishing the line) immediately outside reserve borders. The absence of these catch and effort trends, which are indicative of spillover, may be due to moderate total mortality (Z = 0.59 for legal sized lobsters outside reserves), which was estimated from analysis of growth and length frequency data collected as part of our CFR program. Spillover at the Channel Islands reserves may be occurring but at levels that are insufficient to influence the fishery dynamics that we measured. Future increases in fishing effort (outside reserves) and lobster biomass (inside reserves) are likely and may lead to increased spillover, and CFR provides an ideal platform for continued assessment of fishery-reserve interactions.     

Shelter selection of the spiny lobster <Emphasis Type="Italic">Palinurus elephas</Emphasis> under different levels of <Emphasis Type="Italic">Octopus vulgaris</Emphasis> predation threat

The skill of recognizing and reacting to predators is often based on a learned component. Few studies have examined the role of learning in spiny lobster anti-predator behavior. We investigated whether European spiny lobster (Palinurus elephas) shelter selection is influenced by olfactory stimuli released by one of the most common lobster predators, the common octopus (Octopus vulgaris), and whether the behavioral response to octopus chemical stimuli is innate or influenced by experience. In experimental arenas, we conditioned wild-caught lobsters with three levels of predation threat: no threat, with no predator–prey interaction; medium threat, with odor and visual predator cues only; high threat, active predation risk. We subsequently tested the shelter choice of the conditioned lobster under different experimental conditions: (1) shelter plus seawater; (2) shelter plus seawater plus chemical octopus cue. Our results showed significant differences in mean shelter occupancy with conditioning level. We conclude that P. elephas individuals use chemosensory systems in predator-avoidance mechanisms. Moreover, lobsters subject to a training period of high-level predation threat were able to learn the octopus chemical stimuli and treat its odor as a cue related to predation risk. The findings relative to the spiny lobster learning abilities could be an important tool for future management of lobster populations, e.g., by re-introduction of reared juveniles, which have not yet experienced predation.     

Cryptic species within the commercially most important lobster in the tropical Atlantic, the spiny lobster Panulirus argus

Panulirus argus (Latreille in Ann Mus Hist Nat Paris 3:388–395, 1804) is the lobster of greatest economic importance throughout its distribution. In this study, mitochondrial (Cytochrome Oxidase I and 16S ribosomal genes) and nuclear (Adenine Nucleotide Transporter gene) sequences were used to evaluate the taxonomic status of P. argus sampled from five sites in the Caribbean Sea and nine sites in the Southwest Atlantic. Phylogenetic analyses indicate that lobsters from the two regions form two monophyletic groups with a molecular divergence similar to that observed between distinct congeneric lobster species and much larger than that found between conspecific lobster populations. Therefore, the Caribbean and the Southwest Atlantic lobster populations originally attributed to P. argus belong to different species, with an estimated time of isolation of around 16 Million years. An important consequence of these findings is that the fisheries of spiny lobsters from the Caribbean and the Southwest Atlantic species must be managed separately.     

Recreational diving impacts and the use of pre-dive briefings as a management strategy on Florida coral reefs

Ecotourism often is promoted as an ecologically sustainable activity, but some ecotourism activities negatively impact coastal ecosystems. Impacts of intensive diving tourism on coral reefs remain poorly understood, especially in the Florida Keys. We determined patterns of recreational dive frequency, diver behaviour, and coral damage on reefs near Key Largo, and assessed how pre-dive briefings and other factors affect these damage rates. Recreational divers contacted live stony corals ~ 18 times per scuba dive; most contacts deposited sediment onto corals, but also caused abrasion to coral tissues and fracture of coral skeletons. Divers who received pre-dive ecological briefings caused significantly less coral damage than those who did not, and divers with cameras and/or gloves caused the most damage. The proportion of damaged corals increased significantly with the estimated rate of recreational diving on each reef, and the percent cover of live corals decreased. We conclude that current rates of recreational diving in Key Largo are unsustainable, resulting in damage to >80 % of coral colonies and reduction of live coral cover to <11 % at heavily-dived sites. We recommend that dive tour operators administer pre-dive ecological briefings to all recreational divers, provide extra briefings to camera and glove users, and employ underwater dive guides who intervene when divers inadvertently damage live stony corals. This study provides a scientific basis to support management of intensive ecotourism on Florida coral reefs.     

Defining ecologically relevant scales for spatial protection with long‐term data on an endangered seabird and local prey availability

Human activities are important drivers of marine ecosystem functioning. However, separating the synergistic effects of fishing and environmental variability on the prey base of nontarget predators is difficult, often because prey availability estimates on appropriate scales are lacking. Understanding how prey abundance at different spatial scales links to population change can help integrate the needs of nontarget predators into fisheries management by defining ecologically relevant areas for spatial protection. We investigated the local population response (number of breeders) of the Bank Cormorant (Phalacrocorax neglectus), a range‐restricted endangered seabird, to the availability of its prey, the heavily fished west coast rock lobster (Jasus lalandii). Using Bayesian state‐space modeled cormorant counts at 3 colonies, 22 years of fisheries‐independent data on local lobster abundance, and generalized additive modeling, we determined the spatial scale pertinent to these relationships in areas with different lobster availability. Cormorant numbers responded positively to lobster availability in the regions with intermediate and high abundance but not where regime shifts and fishing pressure had depleted lobster stocks. The relationships were strongest when lobsters 20–30 km offshore of the colony were considered, a distance greater than the Bank Cormorant's foraging range when breeding, and may have been influenced by prey availability for nonbreeding birds, prey switching, or prey ecology. Our results highlight the importance of considering the scale of ecological relationships in marine spatial planning and suggest that designing spatial protection around focal species can benefit marine predators across their full life cycle. We propose the precautionary implementation of small‐scale marine protected areas, followed by robust assessment and adaptive‐management, to confirm population‐level benefits for the cormorants, their prey, and the wider ecosystem, without negative impacts on local fisheries.     

Creation of a Gilded Trap by the High Economic Value of the Maine Lobster Fishery

Abstract: Unsustainable fishing simplifies food chains and, as with aquaculture, can result in reliance on a few economically valuable species. This lack of diversity may increase risks of ecological and economic disruptions. Centuries of intense fishing have extirpated most apex predators in the Gulf of Maine (United States and Canada), effectively creating an American lobster (Homarus americanus) monoculture. Over the past 20 years, the economic diversity of marine resources harvested in Maine has declined by almost 70%. Today, over 80% of the value of Maine's fish and seafood landings is from highly abundant lobsters. Inflation‐corrected income from lobsters in Maine has steadily increased by nearly 400% since 1985. Fisheries managers, policy makers, and fishers view this as a success. However, such lucrative monocultures increase the social and ecological consequences of future declines in lobsters. In southern New England, disease and stresses related to increases in ocean temperature resulted in more than a 70% decline in lobster abundance, prompting managers to propose closing that fishery. A similar collapse in Maine could fundamentally disrupt the social and economic foundation of its coast. We suggest the current success of Maine's lobster fishery is a gilded trap. Gilded traps are a type of social trap in which collective actions resulting from economically attractive opportunities outweigh concerns over associated social and ecological risks or consequences. Large financial gain creates a strong reinforcing feedback that deepens the trap. Avoiding or escaping gilded traps requires managing for increased biological and economic diversity. This is difficult to do prior to a crisis while financial incentives for maintaining the status quo are large. The long‐term challenge is to shift fisheries management away from single species toward integrated social‐ecological approaches that diversify local ecosystems, societies, and economies.     

Temporal patterns and transport processes in recruitment of spiny lobster (Panulirus argus) postlarvae to south Florida

We used time-series analysis to identify weekly and annual patterns in the supply of spiny lobster, Panulirus argus (Latreille, 1804), postlarvae to the Florida Keys, USA, over an 8 yr period. We also investigated the relationship between postlarval influx and wind forcing as a transport mechanism using the complex vector-scalar correlation analysis. Postlarval supply had a lunar phase periodicity at 4.5 wk intervals, with postlarval abundance peaking between the new moon and first-quarter lunar phases. A distinct annual cycle of postlarval supply with two peak periods was also apparent. Cross-correlation analysis between relative postlarval abundance and a 12 mo cycle showed that the annual peak occurs in spring, centered around March. With the 12 mo periodicity removed, a smaller peak at 5 mo intervals was also well defined. Wind-forcing for 7 d prior to the time of postlarval collection was marginally correlated with postlarval abundance through the entire time-series; the association was strongest during the late fall to early spring months. The analysis indicated that postlarval supply was correlated with winds from the northeast (ca. 45°), which are associated with winter atmospheric fronts. In contrast to results reported for other spiny lobster populations, these patterns suggest that recruitment of lobster postlarvae to south Florida is predictable only at a gross level and is presumably affected by the temporally inconsistent structure of regional oceanic gyres and variability in the timing of lobster spawning in the Caribbean. Received: 24 January 1997 / Accepted: 4 March 1997     

Effects of autotomy on long-term survival and growth of painted spiny lobster (Panulirus versicolor) on the Great Barrier Reef, Australia

The effects of autotomy (shedding of appendages) on survival and growth rates of painted spiny lobster were investigated at Northwest Island (23^° 18?? S, 152^° 43?? E) during the period 2003?C2006. Adult lobsters were captured, tagged, and classified as either uninjured (n?=?68), minimally injured (n?=?39) or moderately injured (n?=?19) depending on the number and type of appendages that were autotomized during capture and handling. Six to thirty-six months after release, 86 lobsters were recaptured (mean time at large?=?305?days). Recapture rates of uninjured (64.7%), minimally injured (71.8%), and moderately injured lobsters (73.7%) were not significantly different. Similarly, mean annualized growth rates of uninjured, minimally injured, and moderately injured lobsters were not significantly different. This suggests that the energetic cost of a single episode of autotomy is either negligible or exists as a trade-off with some other life history trait, such as reduced reproductive performance. These results support the use of certain management tools (e.g., size limits) that prescribe release of non-legal lobsters, regardless of their injury status.     

The spiny lobster Palinurus elephas in Scottish waters

Observations have been made on the biology of spiny lobsters, Palinurus elephas Fabr., caught by divers on the west coast of Scotland from 1972 to 1975. P. elephas in this Scottish population show the same length/carapace length relationship as spiny lobsters taken from the west of Ireland. It occurs inshore during the period April to October on shallow rocky reefs in depths of 5 to 20 m. In females, a pre-mating moult occurs in the inshore areas in late summer. Soon after, mating takes place and the spermatophores are deposited on the sternum of the female. The eggs, which are shed within 1 week to 10 days, are then carried by the female throughout the winter, and hatch in the following spring. Offshore migration takes place in the autumn, after mating and egg laying, and inshore migration takes place in the spring and summer.     

Shelter preference in captive juveniles of European spiny lobster <Emphasis Type="Italic">Palinurus elephas</Emphasis> (Fabricius, 1787)

Physical shelter features (e.g. shape, size and substrate slope) were tested in the laboratory to evaluate the preferences of juveniles of European spiny lobster, Palinurus elephas. Tethering experiments to assess whether substrate slope affects the vulnerability of juveniles to predation were also performed. Our results showed that: (1) semi-circular dens were significantly preferred over square and circular shapes; (2) when not disturbed by a predator, lobsters sheltered in holes with a diameter closely related to their own size, while in the presence of a potential predator lobsters showed no significant preference for a particular shelter size; (3) lobsters significantly preferred dens excavated on sub-vertical (35°) substrates over those excavated on vertical ones (90°); (4) individual lobsters tethered on vertical substrates were subject to greater predation activity than those tethered on horizontal structures. In conclusion, the present study contributes to the understanding of how physical properties of shelters affect the choice of P. elephas juveniles, enhancing their protection and survival rate.     

Circadian shelter occupancy patterns and predator–prey interactions of juvenile Caribbean spiny lobsters in a reef lagoon

The spiny lobster, Panulirus argus, is predominantly nocturnal, remaining inside shelters during the day and foraging outside at night, presumably to minimize predation risk. Predation risk generally decreases with increasing lobster size. Therefore, this study examined the hypothesis that size would influence this basic circadian pattern. Video cameras continuously recorded the shelter occupancy of juvenile lobsters (n = 72) having a carapace length (CL) of 30–62 mm that were tethered to shelters in a shallow reef lagoon. The lobsters’ shelter occupancy was 100% during the day, but declined linearly from shortly before sunset to a minimum of 50% shortly after midnight and then increased linearly, reaching 100% by 1 h after sunrise. The percent time the lobsters spent in the shelters followed a similar trend, but there was wide variability at night (0–100%) for individual lobsters. Lobsters left their shelters 2–30 times night⁻¹, with a majority of excursions lasting <10 min. These results suggest that juvenile P. argus minimize predation risk by remaining in their shelters as long as possible but offset the energetic cost of this behavior by foraging close to their shelters for several short periods at night. This emergence pattern contrasts with those of early benthic phase lobsters (<15 mm CL), which seldom leave their shelters, and adults (>80 mm CL), which have a dusk/early evening peak in activity and leave the shelter for extended periods of time during the night. Furthermore, a minimum shelter occupancy in the middle of the night appears especially well adapted to avoid exposure to daytime predators. Videotaped observations also included interactions between lobsters and two dominant lobster predators, the triggerfish, Balistes capriscus, and the octopus Octopus cf. vulgaris. Lobsters responded differently to these predators: remaining in the shelter when attacked by a triggerfish and fleeing the shelter when attacked by an octopus. Triggerfish were nearly twice as likely to attack a lobster that was outside of the shelter than inside. Once under attack, however, a lobster had nearly the same chance of surviving if it was inside or outside. Results suggest that the patterns of shelter use and emergence change as lobsters grow, probably reflecting the interplay between perception of predation risk and the need to forage. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Determining prey availability for rock lobsters, Jasus lalandii : diver-sampling versus monochrome video photography

Accurate determination of diet is important in ecological studies. Many macrobenthic predators fragment their prey so finely that identification of component organisms is difficult. Knowledge of the food available to masticatory predators such as lobsters can help in determining potential prey and enhance the accuracy of dietary assessments. With SCUBA divers limited by depth and submersion time, benthic analyses are becoming increasingly reliant on camera systems. Here, based on paired samples, we assess the relative merits of using removal-sampling by divers versus monochrome video photography to determine the prey spectrum available to macrobenthic predators. We also relate the results to the gut contents of rock lobsters (Jasus lalandii) that were collected simultaneously. Diver samples took on average four times longer to collect and process than video-image samples. No significant differences were evident between the number of species identified in diver samples and video-image samples (n = 21 paired samples, Student's t = 0.233, P > 0.1), although an analysis of similarity (ANOSIM) test revealed that the results obtained from the two sampling methods were significantly different in terms of species composition (global R = 0.203, P < 0.01). Cluster analysis based on Bray–Curtis similarity to compare diver and video-image samples revealed four clusters of samples with <35% similarity, confirming that the two techniques do sample different assemblages of species. Video images detected and allowed identification of all species of prey recorded in the rock lobsters' stomach contents, probably because rock lobsters prey mainly on relatively large prey which are readily detected by video. Diver-sampling underestimated or failed to detect two important prey types, namely small barnacles and encrusting coralline algae. We concluded that the camera system employed was adequate for assessing the prey assemblages available to macrobenthic predators such as lobsters, and that it was considerably more economical than using divers. Received: 10 September 1998 / Accepted: 6 May 1999     

Annual,semilunar, and diel reproductive rhythms in the Hawaiian labrid Thalassoma duperrey

In September 1978, 5 034 juvenile western rock (spiny) lobsters (Panulirus cygnus George) were tagged on a shallow coastal reef near Cliff Head in Western Australia. Of these, 26% were recaptured and removed from the population during the subsequent commercial fishery between 15 November 1978 and 30 June 1979. A resurvey of the tagging site in September 1979 found that size structure of the population was similar to the tagged population in September 1978 but that the 1979 population was composed of almost completely different individuals and only 2% of the rock lobsters tagged in September 1978 were recaught at this time. A wide range of movement was indicated by the recapture positions of 354 rock lobsters caught off the tagging site. Most travelled almost directly offshore but some also travelled offshore in a northwesterly direction. Both trends were shown in the data from the recaptures of 18 rock lobster released farther offshore from Cliff Head in September 1981, and of 19 released at a close inshore site in earlier years. Maximum offshore rates of movement of up to 622 m d^-1 were recorded for distances up to 37 nautical miles (ca. 68 km). Growth rates for the tagged individuals which moved off, and those that were caught on, the tagging site were similar and close to growth rate under optimal conditions for this species. Under the present high levels of exploitation few, if any, of the rock lobsters remain to become permanent residents in the coastal reefs. It is believed that the north-westerly movements of some of the pre-adult rock lobsters, on their way to the breeding grounds on the outer edge of the continental shelf, are due to the upstream movement of these rock lobsters in a current from the north. The north-westerly movements allow the rock lobster population increased utilization of the food supplies of the coastal area.     

Temperate marine reserves enhance targeted but not untargeted fishes in multiple no-take MPAs.

Although many papers report the effects of no-take marine protected areas (MPAs or reserves), scientifically rigorous empirical studies are rare, particularly for temperate reef fishes. We evaluated the responses of fish populations to protection from fishing in reserves by comparing densities and sizes inside and outside of five no-take reserves in southern California, USA. Our results are robust because we compared responses across multiple rocky-reef reserves in two different years and controlled for possible site differences by (a) ensuring that habitat characteristics were the same inside and outside reserves, and (b) sampling species that are not targeted, which would not be expected to have a direct response to fishing. We compared fish density and size and calculated biomass and egg production across all five sites. Fishes targeted by recreational and/or commercial fisheries consistently exhibited increases in mean density (150%), size (30%), biomass (440%), and egg production (730%) inside reserves. Reserve effects were greatest for legal-sized targeted fishes: significantly greater densities were found exclusively inside reserves for targeted species (580%), the largest size classes existed only inside reserves, and mean biomass was 1000% higher. These responses were unlikely to have been caused by habitat differences because there were no significant differences in habitat characteristics between reserve and control locations. Densities of non-targeted species did not differ between reserve and non-reserve locations, further supporting the conclusions that differences in targeted species between reserve and control locations were due to harvesting rather than site-specific effects. Although MPAs cannot replace traditional fisheries management, the concentration of increased biomass and egg production is a unique MPA benefit that serves both reserves and fisheries. Scientifically rigorous studies that include multiple reserves, such as this study, are needed to inform management and policy decisions.     

Disease avoidance influences shelter use and predation in Caribbean spiny lobster

Shelter competition is uncommon among social animals, as is the case among normally gregarious Caribbean spiny lobsters (Panulirus argus). However, healthy lobsters avoid sheltering with conspecifics infected by a lethal pathogenic virus, PaV1. These contradictory behaviors have implications for shelter use and survival, especially in areas where shelter is limited. In laboratory experiments, we tested shelter competition between paired healthy and diseased juvenile lobsters in shelter-limited mesocosms. Neither healthy nor diseased lobsters dominated access to shelters, but lobsters shared shelter less often when diseased lobsters were present relative to controls with two healthy lobsters. We hypothesized that exclusion of juvenile lobsters from shelter results in increased mortality from predation, especially for the more lethargic, infected individuals. Field tethering trials revealed that predation was indeed higher on infected individuals and on all tethered lobsters deprived of shelter. We then tested in mesocosm experiments how the contrasting risks of predation versus infection by a lethal pathogen influence shelter use. Lobsters were offered a choice of an empty shelter or one containing a diseased lobster in the presence of a predator (i.e., caged octopus) whose presence normally elicits shelter-seeking behavior, and these data were compared with a previous study where the predator was absent. Lobsters selected the empty shelter significantly more often despite the threat of predation, foregoing the protection of group defense in favor of reduced infection risk. These results offer striking evidence of how pathogenic diseases shape not only the behavior of social animals but also their use of shelters and risk of predation.     

Bioeconomic models of marine recreational fishing

The theory of recreational fishing is developed and conditions are derived for optimal management policy, with special attention given to functional relationships that must be empirically verified. Determinants of the optimal allocation between commercial and recreational fishing effort are derived. The theory is extended to include selected peculiar features of recreational fishing: Some anglers sell their catch; a small proportion of the fishing population accounts for a large proportion of the catch; and anglers throw back a fraction of what they catch. Optimal policies are derived under these more realistic conditions.     

Predator-prey relationships between the rock lobster Jasus lalandii and the mussel Aulacomya ater at Robben Island on the Cape West Coast of Africa

The natural diet and mode of feeding of the rock lobster Jasus lalandii (H. Milne Edwards) was determined in a rock-lobster sanctuary near Cape Town, South Africa. Field observations were tested and confirmed by means of aquarium studies. Rock lobsters feed mainly upon ribbed mussels Aulacomya ater (Molina), which comprise the largest component of the sessile benthic fauna. Mussel remains were found as the major constituent in 97% of the rock-lobster stomachs examined. The density of rock lobsters averaged 8,100 per hectare (0.81 m^-2), while mussel biomass averaged more than 5 kg (wet whole weight) m^-2 within the same depth range (12 to 30 m). More than 80% of the mussel biomass comprised large individuals between 60 and 90 mm in length. Large rock lobsters (mainly males) were capable of feeding on all sizes of mussels, although many of these were inaccessible to predation. Smaller rock lobsters became progressively more limited in the size range of mussels which they could crack open and consume. Competition between rock lobsters for small mussels appeared to be intense, as mussels of suitable size for feeding were generally in short supply to most of the rock-lobster population. Hence, feeding and growth rates of rock lobsters are likely to be affected by the relative population densities of predator and prey. Growth rates of rock lobsters could be limited by food supplies even in areas where mussel biomass is comparatively large.     

Marine reserves reduce risk of climate-driven phase shift by reinstating size- and habitat-specific trophic interactions

Spatial closures in the marine environment are widely accepted as effective conservation and fisheries management tools. Given increasing human-derived stressors acting on marine ecosystems, the need for such effective action is urgently clear. Here we explore mechanisms underlying the utility of marine reserves to reinstate trophic dynamics and to increase resilience of kelp beds against climate-driven phase shift to sea urchin barrens on the rapidly warming Tasmanian east coast. Tethering and tagging experiments were used to examine size- and shelter-specific survival of the range-extending sea urchin Centrostephanus rodgersii (Diadematidae) translocated to reefs inside and outside no-take Tasmanian marine reserves. Results show that survival rates of C. rodgersii exposed on flat reef substratum by tethering were approximately seven times (small urchins 10.1 times; large urchins 6.1 times) lower on protected reef within marine reserve boundaries (high abundance of large predatory-capable lobsters) compared to fished reef (large predatory lobsters absent). When able to seek crevice shelter, tag-resighting models estimated that mortality rates of C. rodgersii were lower overall but remained 3.3 times (small urchins 2.1 times; large urchins 6.4 times) higher in the presence of large lobsters inside marine reserves, with higher survival of small urchins owing to greater access to crevices relative to large urchins. Indeed, shelter was 6.3 times and 3.1 times more important to survival of small and large urchins, respectively, on reserved relative to fished reef. Experimental results corroborate with surveys throughout the range extension region, showing greater occurrence of overgrazing on high-relief rocky habitats where shelter for C. rodgersii is readily available. This shows that ecosystem impacts mediated by range extension of such habitat-modifying organisms will be heterogeneous in space, and that marine systems with a more natural complement of large and thus functional predators, as achievable within no-take reserves, will minimize local risk of phase shifts by reinstating size and habitat-specific predator-prey dynamics eroded by fishing. Importantly, our findings also highlight the crucial need to account for the influence of size dynamics and habitat complexity on rates of key predator-prey interactions when managing expectations of ecosystem-level responses within marine reserve boundaries.     

Malthusian overfishing and efforts to overcome it on Kenyan coral reefs

This study examined trends along a gradient of fishing intensity in an artisanal coral reef fishery over a 10-year period along 75 km of Kenya's most populated coastline. As predicted by Malthusian scenarios, catch per unit effort (CPUE), mean trophic level, the functional diversity of fished taxa, and the diversity of gear declined, while total annual catch and catch variability increased along the fishing pressure gradient. The fishery was able to sustain high (approximately 16 Mg x km(-2) x yr(-1)) but variable yields at high fishing pressure due to the dominance of a few productive herbivorous fish species in the catch. The effect of two separate management strategies to overcome this Malthusian pattern was investigated: fisheries area closure and elimination of the dominant and most "competitive" gear. We found that sites within 5 km of the enforced closure showed significantly lower total catch and CPUE, but increased yield stability and trophic level of catch than predicted by regression models normalized for fishing effort. Sites that had excluded illegal beach seine use through active gear management exhibited increased total catch and CPUE. There was a strong interaction between closure and gear management, which indicates that, for closures to be effective at increasing catch, there must be simultaneous efforts at gear management around the periphery of the closures. We propose that Malthusian effects are responsible for the variation in gear and catch and that active management through reduced effort and reductions in the most competitive gear have the greatest potential to increase the functional and trophic diversity and per-person productivity.