捕捞诱导鱼类生物学特征进化研究进展

随着捕捞强度加大,渔业生物为了应对捕捞压力、维持种族繁衍,逐渐产生适应性进化,这一过程称为捕捞诱导进化(Fishing-induced evolution,FIE),通常表现为渔业生物个体变小、性成熟提前,个体对捕捞的敏感性增强,进一步导致渔业种群结构简单、生态系统稳定性下降和渔业经济效益降低。认知捕捞诱导的渔业生物适应性进化,掌握捕捞对渔业种群的作用机制,有利于制定科学合理的渔业资源养护与管理策略。虽然FIE方面已进行了大量研究,但FIE在生理、生态及基因层面上对渔业生物的具体影响过程尚未明确,尤其是在气候变化、多物种相互作用等的动态环境中,FIE的作用方式更为复杂。作者综述了鱼类FIE的主要研究方法,总结了捕捞对大个体的选择性在鱼类生长、性成熟和行为方面的影响,并分析了这种影响对渔业种群恢复与管理产生的效应,以及今后需要解决的关键科学问题,旨在为FIE的进一步深入研究和渔业资源的科学管理提供帮助。     

Effects of cryptic mortality and the hidden costs of using length limits in fishery management

Fishery collapses cause substantial economic and ecological harm, but common management actions often fail to prevent overfishing. Minimum length limits are perhaps the most common fishing regulation used in both commercial and recreational fisheries, but their conservation benefits can be influenced by discard mortality of fish caught and released below the legal length. We constructed a computer model to evaluate how discard mortality could influence the conservation utility of minimum length regulations. We evaluated policy performance across two disparate fish life‐history types: short‐lived high‐productivity (SLHP) and long‐lived low‐productivity (LLLP) species. For the life‐history types, fishing mortality rates and minimum length limits that we examined, length limits alone generally failed to achieve sustainability when discard mortality rate exceeded about 0.2 for SLHP species and 0.05 for LLLP species. At these levels of discard mortality, reductions in overall fishing mortality (e.g. lower fishing effort) were required to prevent recruitment overfishing if fishing mortality was high. Similarly, relatively low discard mortality rates (>0.05) rendered maximum yield unobtainable and caused a substantial shift in the shape of the yield response surfaces. An analysis of fishery efficiency showed that length limits caused the simulated fisheries to be much less efficient, potentially exposing the target species and ecosystem to increased negative effects of the fishing process. Our findings suggest that for overexploited fisheries with moderate‐to‐high discard mortality rates, reductions in fishing mortality will be required to meet management goals. Resource managers should carefully consider impacts of cryptic mortality sources (e.g. discard mortality) on fishery sustainability, especially in recreational fisheries where release rates are high and effort is increasing in many areas of the world.     

Evolutionary impact assessment: accounting for evolutionary consequences of fishing in an ecosystem approach to fisheries management

Managing fisheries resources to maintain healthy ecosystems is one of the main goals of the ecosystem approach to fisheries (EAF). While a number of international treaties call for the implementation of EAF, there are still gaps in the underlying methodology. One aspect that has received substantial scientific attention recently is fisheries‐induced evolution (FIE). Increasing evidence indicates that intensive fishing has the potential to exert strong directional selection on life‐history traits, behaviour, physiology, and morphology of exploited fish. Of particular concern is that reversing evolutionary responses to fishing can be much more difficult than reversing demographic or phenotypically plastic responses. Furthermore, like climate change, multiple agents cause FIE, with effects accumulating over time. Consequently, FIE may alter the utility derived from fish stocks, which in turn can modify the monetary value living aquatic resources provide to society. Quantifying and predicting the evolutionary effects of fishing is therefore important for both ecological and economic reasons. An important reason this is not happening is the lack of an appropriate assessment framework. We therefore describe the evolutionary impact assessment (EvoIA) as a structured approach for assessing the evolutionary consequences of fishing and evaluating the predicted evolutionary outcomes of alternative management options. EvoIA can contribute to EAF by clarifying how evolution may alter stock properties and ecological relations, support the precautionary approach to fisheries management by addressing a previously overlooked source of uncertainty and risk, and thus contribute to sustainable fisheries.     

Genetic connectivity supports recovery of gorgonian populations affected by climate change

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Genetic differentiation among Atlantic island populations of the brown spiny lobster Panulirus echinatus (Decapoda: Palinuridae)

1. Declines in Panulirus echinatus Smith, 1869 populations along their wide distribution in the Atlantic Ocean have spurred efforts to improve their fisheries management and conservation. In this study, the genetic structure of these populations is reported for the first time.
2. In a survey of 18 species‐specific polymorphic microsatellite markers, 152 individuals were genotyped from five Atlantic oceanic islands, covering most of the insular distribution range of the species. The analyses revealed that P. echinatus is genetically partitioned into two stocks in the Atlantic Ocean. A highly significant genetic structure was observed between north‐east and south‐west Atlantic populations based on fixation index, discriminant analysis of principal components, and structure and barrier analysis.
3. We suggest that the Equatorial Circulation System represents a biophysical barrier that effectively limits migration among Atlantic subtropical gyre systems, as has been described for other species. Other physical and ecological barriers, such as the Mid‐Atlantic Ridge itself, the distance between the eastern and western sides of the Atlantic (Mid‐Atlantic Barrier) and water mass differences, as well as other biological aspects, may also influence larval dispersal and modulate the insular distribution of this species.
4. The results show the existence of two distinct genetic stocks of P. echinatus and have implications for fisheries management in the Atlantic Ocean, including their independent management according to their individual status. The Cabo Verde and the Canarian populations (north‐east Atlantic) showed the lowest level of genetic variability in comparison with the south‐western populations. A combination of factors that have occurred or are occurring in the Canary Islands, such as overfishing and volcanic eruptions, is likely to explain the reduced abundance of this lobster species in the area.

     

Assessing the impact of population decline on mating system in the overexploited Mediterranean red coral

1. Understanding the interactions among demographic parameters, mating system and population dynamics is key to predict the response of populations to global change. The Mediterranean red coral is a precious octocoral suffering from population decline due to overfishing and warming‐driven mass mortality events.
2. While the demographic consequences of these two pressures are well characterized, little is known regarding their impact on population dynamics and evolution of red coral populations. The main objective of this study was to fill this gap focusing more particularly on mating pattern and genetic drift.
3. Combining sibship and progeny arrays analyses, a genetic characterization of the red coral mating system was conducted. In addition, a synchronic approach was developed comparing mating patterns in two populations with contrasting demographic patterns: a pristine‐like population and a declining population.
4. The results show that polyandry is likely to be the norm in red coral. The similar patterns of genetic diversity between adults and larvae combined with the lack of differential reproductive success among putative fathers did not support significant sweepstakes effects during larval production. While instantaneous biparental inbreeding was detected, no long‐term inbreeding was observed even in the declining population. Mating patterns and effective population sizes in the two populations were not statistically different. Nevertheless, a trend towards a slightly higher inbreeding and a lower number of breeders was observed in the declining population.
5. Accordingly, we hypothesized that an increase in male gamete dispersal may buffer the increase of genetic drift expected in the declining population. This feedback between demographic decline and reproductive pattern may potentially take part in the long‐term persistence of red coral populations. However, the negative trend reported in the declining population unambiguously supports the need to maintain high densities of reproductive colonies to the functioning of red coral populations.

     

A new role for MSY in single-species and ecosystem approaches to fisheries stock assessment and management

In 1977, Peter Larkin published his now‐famous paper, ‘An epitaph for the concept of maximum sustained yield’. Larkin criticized the concept of single‐species maximum sustained yield (MSY) for many reasons, including the possibility that it may not guard against recruitment failure, and the impossibility of maximising sustainable yields for all species simultaneously. However, in recent years, there has been a fundamental change in the perception of the fishing mortality associated with MSY (F_MSY) as a limit to be avoided rather than a target that can routinely be exceeded. The concept of F_MSY as a limit is embodied in several United Nations Food and Agriculture Organization (FAO) agreements and guidelines, and has now been incorporated into the US Magnuson–Stevens Fishery Conservation and Management Act. As a result, the United States now requires the development of overfishing definitions based on biological reference points that treat the F_MSY as a limit reference point and must also define a lower limit on biomass below which rebuilding plans with strict time horizons must be developed. This represents a major paradigm shift from the previously mandated (but often unachieved) objective to simply maintain fishing mortalities at levels below those associated with recruitment overfishing. In many cases, it requires substantial reductions in current fishing mortality levels. Therefore, the necessity of the new paradigm is continually questioned. This paper draws on examples from several fisheries, but specifically focuses on the recent US experience illustrating the practical difficulties of reducing fishing mortality to levels below those corresponding to MSY. However, several studies suggest that even more substantial reductions in fishing mortality may be necessary if ecosystem considerations, such as multispecies interactions, maintenance of biodiversity and genetic diversity, and reduction of bycatch and waste, are taken into account. The pros and cons of moving beyond single‐species assessment and management are discussed. A US plan for improving stock assessments indicates that even a ‘basic’ objective such as ‘adequate baseline monitoring of all managed species’ may be extremely costly. Thus, the suggestion of Larkin (1983, 1997) that the costs of research and management should not exceed 10–20% of the landed value of the catch may preclude comprehensive ecosystem management. More importantly, neither single‐species nor ecosystem‐based fisheries management is likely to improve appreciably unless levels of fishing capacity are aligned with resource productivity, as is currently being promoted by FAO and several individual nations.     

Dynamic angling effort influences the value of minimum‐length limits to prevent recruitment overfishing

Recruitment overfishing occurs when stocks are fished to a level where recruitment declines proportionally with adult abundance. Although typically considered a commercial fishery problem, recruitment overfishing can also occur in freshwater recreational fisheries. This study developed an age‐structured model to determine if minimum‐length limits can prevent recruitment overfishing in black crappie, Pomoxis nigromaculatus (LeSueur), and walleye, Sander vitreus (Mitchill) fisheries considering angling effort response to changes in fish abundance. Simulations showed that minimum‐length limits prevented recruitment overfishing of black crappie and walleye, but larger minimum‐length limits were required if angler effort showed only weak responses to changes in fish abundance. Low angler‐effort responsiveness caused fishing mortality rates to remain high when stock abundance declined. By contrast, at high effort responsiveness, anglers left the fishery in response to stock declines and allowed stocks to recover. Angler effort for black crappie and walleye fisheries suggested that angler effort could be highly responsive for some fisheries and relatively stable for others, thereby increasing the risk of recruitment overfishing in real fisheries. Recruitment overfishing should be considered seriously in freshwater recreational fisheries, and more studies are needed to evaluate the responsiveness of angler effort to changes in fish abundance.     

Atlantic cod recovery from the Allee effect zone: contrasting ecological and evolutionary rescue

The ability of a population to recover from disturbances is fundamental for its persistence. Impaired population recovery might be associated with a demographic Allee effect. Immigration from adjacent populations could accelerate the recovery not only by promoting population growth beyond the Allee effect threshold but also by bringing in advantageous genotypes. We explore the nature and role of ecological and evolutionary rescue in an Atlantic cod (Gadus morhua Gadidae) population fished below its Allee effect threshold. We utilize an eco‐evolutionary model and simulate scenarios, where the target population evolves in response to selective fishing and sample immigrants from (a) a source population similarly adapted to fishing (post‐fishing genotypes) or (b) an unexploited source population with natural genetic and phenotypic diversity (pre‐fishing genotypes). Immigration of pre‐fishing genotypes enhances the recovery due to the larger body sizes and the flow of associated genes. Post‐fishing immigrants can also promote the population abundance recovery, but they increase uncertainty about recovery times as compared to pre‐fishing immigrants and do not promote evolutionary recovery. Our results stress the importance of maintaining genetic and phenotypic diversity and suggest that marine protected areas can serve as an important source of immigrants to promote both the demographic and evolutionary recovery of exploited populations.     

基于线粒体Cyt b基因序列的绿鳍马面鲀6个野生群体的遗传结构分析

利用线粒体细胞色素b(cytochrome b,Cyt b)基因分析了我国沿海绿鳍马面鲀(Thamnaconus septentrionalis)6个野生群体(大连、秦皇岛、蓬莱、日照、舟山和汕头)的遗传多样性和群体遗传结构。结果显示,在176个个体915 bp的Cyt b部分序列中共检测到32个变异位点和30个单倍型;6个群体的单倍型多样性为0.883~0.953,核苷酸多样性0.0032~0.0039;群体间固定指数较小,呈中低度分化;AMOVA分析表明,遗传变异主要来源于群体内,群体间遗传分化未达到显著水平。群体历史动态分析表明,绿鳍马面鲀在16.9万~42.2万年前经历种群扩张。总体上,各群体间的遗传分化较小,并未形成相应的地理支系,推测黑潮的输送作用以及绿鳍马面鲀的洄游习性维持了各群体间高强度的基因交流;中更新世中晚期的气候波动对绿鳍马面鲀的种群扩张以及地理分布格局可能具有重要影响。本研究结果加深了人们对于绿鳍马面鲀资源情况的认识,为绿鳍马面鲀资源的保护和合理利用提供了理论依据。     

Gillnet fishing drives lake‐migrating brown trout to near extinction in the Lake Päijänne region,Finland

Abstract Wild stocks of brown trout, Salmo trutta L., collapsed in Finnish inland waters during the 20th Century because dams prevented upstream migration, and low water quality and stream dredging weakened reproduction. The demise in migratory stocks was coupled with overfishing, mainly by gillnetting on lakes. Consequently, the migratory spawning stocks have diminished to negligible levels. The remaining stocks exhibit restricted immigration and emigration, are supplemented by continuous stocking, and their natural genetic diversity is affected by human activities. In recent years, various recovery actions have been implemented including stream channel restorations, fish passage facilities constructed and stocking of eggs and smolts. Gillnetting has also been regulated by banning certain mesh sizes, and catch‐and‐release of wild trout is spreading amongst sport fishers. However, these measures seem to be inadequate and almost no recovery of migratory populations has been reported. The problem of by‐catch in intensive gillnetting continues to threaten stocks and creates disputes between stakeholders.     

Multicentury trends and the sustainability of coral reef fisheries in Hawai‘i and Florida

Global overfishing indicates a need to define fisheries sustainability thresholds and identify social factors promoting successful management, but rates of fishing and factors mediating sustainability over long timescales are largely unknown. Here, we reconstruct fisheries yield for the entire period of human habitation (five to seven centuries) for two coral reef ecosystems with substantially different fisheries histories (Florida Keys and the Hawaiian Islands) and evaluate the management strategies associated with periods of sustainable fishing. This involved a mixed methods approach, in which we estimated yield by fishery sector (commercial, subsistence, recreational and aquaculture) and characterized management strategies associated with periods of sustained high yields. We found differences between the two locations, with Hawai‘i sustaining yields of more than 12 mt km^?2 for four centuries prior to the arrival of Europeans. This period was characterized by adaptive management whose design and enforcement exhibited characteristics of common property resource governance systems, and which effectively protected reef habitat, vulnerable life‐history stages for fish, and species with high susceptibilities to overfishing. Reefs in both Florida and Hawai‘i were exploited intensively after European contact, with sequential export‐driven depletion evident in Florida over the past century. Today, both exhibit strikingly similar modern catch levels, with landings exceeding 10 mt km^?2 and evidence of overfishing. Our results demonstrate that management strategies and social institutions that support strict enforcement by a local rule‐making authority have had substantial impacts on fisheries yields in the past and suggest that long‐term sustainability of fisheries is possible, although rare today.     

Metapopulation ecology in the sea: from Levins' model to marine ecology and fisheries science

Marine and fisheries scientists are increasingly using metapopulation concepts to better understand and model their focal systems. Consequently, they are considering what defines a metapopulation. One perspective on this question emphasizes the importance of extinction probability in local populations. This view probably stems from the focus on extinction in Levins' original metapopulation model, but places unnecessary emphasis on extinction–recolonization dynamics. Metapopulation models with more complex structure than Levins' patch‐occupancy model and its variants allow a broader range of population phenomena to be examined, such as changes in population size, age structure and genetic structure. Analyses along these lines are critical in fisheries science, where presence–absence resolution is far too coarse to understand stock dynamics in a meaningful way. These more detailed investigations can, but need not, aim to assess extinction risk or deal with extinction‐prone local populations. Therefore, we emphasize the coupling of spatial scales as the defining feature of metapopulations. It is the degree of demographic connectivity that characterizes metapopulations, with the dynamics of local populations strongly dependent upon local demographic processes, but also influenced by a nontrivial element of external replenishment. Therefore, estimating rates of interpopulation exchange must be a research priority. We contrast metapopulations with other spatially structured populations that differ in the degree of local closure of their component populations. We conclude with consideration of the implications of metapopulation structure for spatially explicit management, particularly the design of marine protected area networks.     

Factors influencing the scope and quality of science and management decisions

The lecture traces the historical path to overfishing of the world's fish and shellfish stocks, and provides an assessment of marine fish resources in the later half of the 1990s. The basis of overfishing as noted by various fishery scientists is reviewed. Four factors, including institutional paralysis, the rapidity of technological developments, uncertainty of science, and the inability to monitor and enforce regulations are identified as the major problems leading to overfishing. The failure of the world community to deal with extensive overfishing, appears to have motivated managers and scientists to promote a new fishery management paradigm that focuses on a broader set of problems resulting from fishing, and establishes a more conservative decision‐making process founded on precautionary principle and uncertainty. The author feels that the evolving paradigm will result in the rebuilding of a number of stocks in the United States, but is less certain of its adoption on a global scale, and whether or not science will play a more useful role in fisheries management. It is noted that the support for fisheries science and the status of fisheries have followed opposite courses. Over the past half century marine science has boomed, diversified and become intellectually and materially enriched, while the number of overfished stocks and ecological disasters has increased. Looking ahead it is expected that fisheries management will move into a more conservative era. The focus of fisheries has moved from full use of ocean resources to establishing yields that take into account the impacts of fisheries on target and non‐target species and the ecosystem in general. Although there has been wide‐spread abuse in the use of the world's fishery resources and condemnation of the fishing industries, the author feels that the government institutions must bear the primary responsibility for the historical course of fishery management and its failure.     

澳洲鲭太平洋群系的资源评估与管理策略

澳洲鲭是西北太平洋重要的经济种类,了解和掌握澳洲鲭太平洋群系资源开发状况对确保其可持续利用具有重要的意义。根据日本中央水产研究所提供的1995—2015年澳洲鲭太平洋群系的生产统计和资源调查资料,利用基于年龄结构的实际种群模型和单位补充量产量模型等进行资源量评估,分析澳洲鲭太平洋群系资源利用情况及其管理策略。结果显示,历年澳洲鲭太平洋群系资源量虽有波动但仍保持在较高水平,2015年资源量最高约为65万t;年平均捕捞死亡系数呈波动下降趋势,2015年捕捞死亡系数只有0.15,近五年平均捕捞死亡系数Fcur=0.33,单位补充量亲体量是未开发时的32.7%,不存在生长型捕捞过度,也不存在补充型捕捞过度,处于可持续开发状态。研究还探讨了水温变化引起自然死亡波动以及不同开捕年龄对澳洲鲭太平洋群系资源状况的影响。研究表明,该渔业目前开发和利用程度合理,建议使用F0.1做为管理参考点进行渔业资源的管理。     

Incorporating accessibility limitation into the surplus production model

Limited access to aquatic populations hinders estimation of their status and establishment of effective management measures. We propose a modified surplus production model to cope with this problem. The model provides population parameters and biological reference points from a time series of annual accessible abundance data. Simulation tests showed that the model provided biological reference point estimates with little bias when sufficiently long time series were available. Even for short time series, we could obtain nearly unbiased estimates by providing information on the exploitation rate at the maximum sustainable yield (F _MSY). As an application, we fit the modified surplus production model to 7-year accessible biomass estimates of a local population of Japanese spiky sea cucumber Apostichopus japonicus using a Bayesian approach. The results indicated that the stock in the area studied was likely to have experienced recent overfishing and had a high probability of being overfished in the future.     

Abalone ranching: a review on genetic considerations

Haliotis L. (abalone) is an economically important genus comprising approximately 14 species with commercial value. Unfortunately, due to habitat destruction, overfishing and illegal harvesting, this wild resource is being depleted to unrecoverable status worldwide. To supply the world demand, abalone are cultured in artificial hatchery environments. In the past few decades, attention has been paid to use these artificial environments to the benefit of the species by implementing abalone ranching or reseeding. This is done in an attempt to rebuild the collapsed populations and to prevent further declines in fished stocks. In various countries in which commercial production is in operation, this has been attempted with various levels of success. For ranching to be successful, various factors, however, need to be considered, one of the most important probably being the correct genetic management of such a ranching programme. Genetic management should include using broodstock that are representatives of natural populations, minimizing the effects of hatchery practices on the genetic make‐up of the animals to be released and investigating the genetic impacts of this practice on wild populations. In this review, the genetic management and the effects if management is not implemented are discussed as well as the potential of molecular markers to be used as genetic tags in identifying hatchery‐reared animals.     

Exploitation of walleye in a Great Plains reservoir: harvest patterns and management scenarios

Abstract This study assessed exploitation and evaluated management options for walleye, Sander vitreus (Mitchill), in Glen Elder Reservoir, Kansas. A total of 2429 walleye varying from 240 to 687 mm was tagged with Carlin dangler tags during 2000–2003. After correcting for tag loss and non‐reporting, exploitation of walleye was estimated as 68.3%. More than 85% of the tagged walleye were harvested during April to June and 75% were harvested within 6 months after tagging. A Beverton‐Holt yield‐per‐recruit model was used to evaluate six minimum length limits varying from 250 to 610 mm. Given current exploitation rates, population demographics and harvest regulations (381‐mm minimum length limit), the walleye population is probably experiencing recruitment overfishing. Model results suggested that a 610‐mm minimum length limit would be required to prevent growth overfishing and a 533‐mm or longer minimum length limit would prevent recruitment overfishing.     

Profile of a fishery collapse: why mariculture failed to save the large yellow croaker

The large yellow croaker (Larimichthys crocea), endemic to East Asia was once one of the three top commercial marine fishes of China PR. Heavily exploited since the 1950s, wild stocks were so severely depleted by the 1980s that most individuals subsequently sold originated from hatcheries. After peaking at about 200 000 tonnes in the mid‐1970s, catches of the croaker in China PR declined by over 90% within just 2 decades; according to most decline criteria this would categorize the croaker as “threatened” and management measures, including restocking, were developed. The extensive government‐sponsored mariculture program introduced to address food supply and overfishing in the 1980s, particularly of the croaker, was one of the earliest for marine finfish, not only in China PR, a nation with a rich and highly successful history in aquaculture, but globally. In this first, in‐depth, profile of a key fishery and early mariculture development, we integrate ecological and biological information with the fishing, management, mariculture and economic history to trace the collapse of wild stocks and assess why management and mariculture did not result in wild stock recovery. Evidence strongly suggests that a combination of heavy exploitation of spawning and over‐wintering aggregations, poor management and overfishing pressure were major factors in stock declines, with contributions from pollution, habitat degradation and marine ecosystem shift. Although the croaker proved a highly successful mariculture candidate, with approximately 70 000 tonnes produced in 2005, the highest of any marine fish cultured in China PR, mariculture and restocking have failed to restore croaker stocks and may have, inadvertently, led to biodiversity losses. The detailed history of the croaker is a sobering reminder that successful mariculture, albeit important for food production and livelihoods, is not necessarily a solution to overfishing, and moreover, may have compromised fishery recovery by competing for funds, attention, space, and maybe genetic resources.     

Spatial abundance patterns and recruitment of a virus‐affected commercial mollusc fishery

Infectious pathogens figure prominently among those factors threatening marine wildlife. Mass mortality events caused by pathogens can fundamentally alter the structure of wild fish stocks and depress recruitment rates and yield. In the most severe instances, this can precipitate stock collapses resulting in dramatic economic losses to once valuable commercial fisheries. An outbreak of a herpes‐like virus among commercially fished abalone populations in the south‐west fishery of Victoria, Australia, during 2006–2007, has been associated with high mortality rates among all cohorts. Long‐term records from fishery‐independent surveys of blacklip abalone Haliotis rubra (Leach) enabled abundance from pre‐ and post‐viral periods to be analysed to estimate stock density and biomass. The spatial distribution of abundance in relation to physical habitat variables derived from high‐resolution bathymetric LiDAR data was investigated. Significant differences were observed in both measures between pre‐ and post‐viral periods. Although there was some limited evidence of gradual stock improvement in recent years, disease‐affected reefs have remained below productivity rates prior to the disease outbreak suggesting a reduction in larval availability or settlement success. This was corroborated by trends in sublegal sized blacklip abalone abundance that has yet to show substantial recovery post‐disease. Abundance data were modelled as a function of habitat variables using a generalised additive model (GAM) and indicated that high abundance was associated with complex reef structures of coastal waters (<15 m). This study highlights the importance of long‐term surveys to understand abalone recovery following mass mortality and the links between stock abundance and seafloor variability.