Marine Policy 51 (2015) 13–20 Contents lists available at ScienceDirect Marine Policy journal homepage: www.elsevier.com/locate/marpol Mismatch between biological, exploitation, and governance scales and ineffective management of sea urchin (Paracentrotus lividus) fisheries in Galicia Rosana Ouréns a,n, Inés Naya a, Juan Freire b a b Recursos Marinos y Pesquerías, Facultad de Ciencias, Universidad de A Coruña, Rúa da Fraga 10, 15008 A Coruña, Spain Teamlabs, Impact Hub Madrid, C. Gobernador 26, 28014 Madrid, Spain ar t ic l e i nf o a b s t r a c t Article history: Received 12 April 2014 Received in revised form 15 July 2014 Accepted 16 July 2014 The spatial structure of fishery resources influences stock dynamics and finally the fishery. Therefore, this aspect should be included as a key topic in the assessment and management of fisheries. The fishery of the sea urchin Paracentrotus lividus in Galicia has been used as case study to demonstrate how the mismatch between biological, fishery and management scales causes failures in the governance, giving rise to over-exploitation. P. lividus is spatially distributed in nested biological units: patches, microstocks, local populations and metapopulations. Fishing operations are local exploiting micro-stocks; however management units in Galician comprise usually more than one local population. This pattern allows the depletion of several micro-stocks without any short-term signals in the exploitation rates over the complete managed territory. Management units should be redefined according to the boundaries of the local populations. In addition, the implementation of reserve networks or a rotation system could allow to effectively managing the resource at a fine-scale. Any of both regulations could also compensate the inverse density dependence that regulates recruitment and fecundity in this species. & 2014 Elsevier Ltd. All rights reserved. Keywords: Metapopulation Density dependence Management Sea urchin Spatial structure 1. Introduction Nowadays it is widely recognised that the fisheries are complex systems integrated by human (i.e. social, economic, and political components) and ecological subsystems in a two-way feedback relationship [1–3]. Social and ecological processes interact at different spatial and temporal scales, and it is essential to match both subsystems to improve stewardship of natural resources and ecosystem services for human well-being and sustainability [4,5]. This perspective is often referred to as the problem of fit [6–9]. A major challenge concerning the problem of fit lies in addressing the governance dimension of ecosystem management [6,9]. Management units are often defined according to historical and political boundaries (for example: community, municipal, regional or national levels), which make little ecological sense [4]. For this reason spatial mismatches between scales of governance and ecosystems are common. Thus, ICES [10] summarised that population and management spatial structures were uncoupled for approx. 33% of the 150 stocks reviewed by the Advisory Committee on n Corresponding author. Present address: Hopkins Marine Station, Stanford University, 120 Ocean View Blvd., Pacific Grove, CA 93950, USA. E-mail address: rosanaoc@gmail.com (R. Ouréns). http://dx.doi.org/10.1016/j.marpol.2014.07.015 0308-597X/& 2014 Elsevier Ltd. All rights reserved. Fishery Management in 1999. For example, the Total Allowed Catch (TAC) for many species is assigned for broad areas that include several local populations [11]. In these cases, the TAC could be appropriate for the overall area but not for each of the subunits, allowing a progressive decline of the reproductive stock [12,13]. The opposite situation occurs in some highly migratory species that range over large ocean areas and whose management problems cannot be solved at the regional or national levels. Here international actions are often required to manage properly these resources [14]. This misfit between scales threatens to undermine the sustainability of the social–ecological system because it can give rise to the reduction of incentives for sustainable management, or to a loss of species, functions, and other system components that are determining in the social–ecological resilience [15,16]. Mismatch between subsystems is evident in echinoid fisheries [17–19]. Echinoids show a strong and persistent spatial structure that has a key role in the population dynamics as well as in the spatial dynamics of the fishing activity. In consequence, the management of the spatial distribution of fishing effort in echinoids is of similar relevance or more important than to manage when and how much to fish [20,21]. To incorporate the “where” in the management, it is needed spatial information to identify possible connections between populations, preferred habitats, and 14 R. Ouréns et al. / Marine Policy 51 (2015) 13–20 areas that require protection, such as nursery and reproductive areas [20,22–24]. However, the conventional assessment models ignore the spatial component and therefore the dimension of the ecological units is often unknown, promoting the problem of fit. Echinoids and other species with a heterogeneous spatial distribution require to be managed at multiple nested scales, including generally a fine-scale [18,19,25,26]. In this regard, areabased management is often suggested as a good management tool for these resources because it allows to manage the fishery at a small scale according to the ecological scales [27,28]. However the outcomes are not conclusive, and whereas in Nova Scotia (Canada), Galicia (Spain) and Baja California (Mexico) this system failed to achieve sustainable resource use [17,29], Chile appears to have had more success [30]. The fishery of the echinoid Paracentrotus lividus in Galicia (NW Spain) is presented here. The aim of this work is to demonstrate that the uncoupling between its spatial population structure and the fishing operational and management units causes failures in the governance, and therefore the social–ecological system does not work properly. The article begins with some needed background information about the echinoid fisheries and management systems worldwide as well as about biological process affecting the spatial distribution of sea urchins. Then, the mismatch between ecological and social subsystems in Galicia is explained, and alternative management models for sustainability are proposed. 2. Sea urchin fisheries worldwide and their management systems Japan is by far the main consumer country of sea urchins [29]. The decline in its production resulted to the development of new fisheries worldwide in the 1980s, seeking to supply the high demand from the Japanese market [31]. Thus, it begins a dramatic rise of the global catches, reaching a historic high in 1995 with 108,969 t. Since then production falls progressively, and in 2012 the world catches did not exceed 63,359 t [32]. This decline is caused by the collapse of some fisheries (e.g. USA, Japan, France, Ireland), but also by the establishment of a management system in other ones, which tries to adjust fishing effort and catches to longterm sustainable levels [33]. Andrew et al. [29] and Williams [33] wrote an extensive review of status and management of sea urchin fisheries worldwide. Some of the most popular regulatory measures are: limited access to the fishery, the use of TACs, the use of minimum legal sizes, closed seasons, gear restrictions and closed areas. Within area-management measures, territorial use rights (TURFs) have been used worldwide in the management of fisheries for echinoid and other benthic species [17,30,34–36]. The success of this system is related to the fact that allows managing resources at the proper spatial scale with specific regulations in each territory. Moreover, TURFs are associated in many cases to a comanagement system where fisher organisations and communities have responsibilities in the management of resources. In this way co-management is expected to produce a higher degree of compliance with management measures, the inclusion of social objectives to regulations, a more active participation of fishers and a greater social cohesion and community development [37,38]. In spite of these potential advantages, TURFs for the sea urchin fishery in Galicia did not prevented temporal closures that affected several local fisheries as a consequence of overfishing [39]. These facts could be a consequence of the use of an inadequate spatial scale for management. The reason for this situation is that the spatial components of the population structure and of the fishing dynamics are not taken into account. 3. Biological characteristics affecting the spatial distribution of echinoids The spatial structure of echinoid populations in general, and of P. lividus specifically, is determined by the oceanographic processes affecting larval dispersal, the habitat selection for settlement, the early mortality of recruits and by the movement and migratory patterns of benthic post-metamorphic phases. Whereas larval dispersal for echinoids operates in scales of 100 s or 1000 s km [40], other processes, such as settlement habitat selection or movement patterns of post-metamorphic phases, might occur in scales of 10 s m. The interaction among processes occurring in this huge range of scales produces a complex spatial structure known as metapopulation [41,42]. From the different definitions proposed for metapopulation [43,44], this term is used here to identify a system of local populations with their own internal dynamics but connected between them by the larval flux. This flux is not so low as to consider insignificant the demographic connectivity among populations; but it is not so high to dilute the internal dynamics of local populations [44–46]. This double scale (local and regional) that characterises ecological and biological processes is a key factor in the genetic structure and evolution of populations [44,46]. Density-dependent mechanisms, recurrent in many biological processes affecting echinoids, influence the spatial distribution and size of populations as well. Thus, the aggregative behaviour of these organisms increases survival rates because represents a defence mechanism against predators and waves [47–49]. Similarly, several studies have documented that fertility rates decrease in areas of low population density [50–52], and, in some echinoid species, the same pattern is evident also for recruitment rates [53– 55]. This latter depensatory or Allee effect [56] determines to a large extent the distribution of recruits because they are concentrated in the patches where adults aggregate. In the case of P. lividus [55] and S. franciscanus [57] these patches with high concentration of recruits are located in the very shallow areas. According to these biological characteristics, echinoid metapopulations comprise spatial units at different scales. Morgan and Shepherd [58] described previously these units, but they will be detailed here again because it is essential to understand their management implications. Echinoids constitute small-scale patches or aggregations where individuals are very close and the physical contact among them is common [48]. The extent of the patches is of about 10 s m2, at least in the case of P. lividus [59,60], and they are separated by bottom areas showing similar habitats and where sea urchins are present isolated and at low densities. The size and location of patches changes in a dynamical way [61] because sea urchins move daily (P. lividus is able to move up to 2 m in 24 h according to Hereu et al. [62]). Patch distribution is not homogeneous in space because they concentrate in zones of high environmental quality. This heterogeneous distribution gives rise to a spatial structure more static at scales of 1000 s m2 that will be named here as micro-stock (Fig. 1), because these units are the smallest sea urchin concentrations that are targeted by the fishing force [63]. A local population (1–10 s km2) comprises several nearby micro-stocks. The limits of the local populations are to some extent arbitrary but they are defined by habitat continuity, being isolated from other local populations by zones without an adequate habitat for sea urchin colonisation. Individuals inside a local population interact and reproduce among them due to individual movements. However these interactions do not occur among individuals pertaining to nearby populations, and local populations are connected among them only by larval dispersal [58]. In the case of P. lividus, Calderón et al. [64] showed genetic R. Ouréns et al. / Marine Policy 51 (2015) 13–20 15 Fig. 1. Diagram of the spatial units involved in the fishery of sea urchin P. lividus. Biological units (ellipses) in increasing size are: patches, micro-stocks, local populations and the metapopulation. Fishing activity units (boxes with dashed line) in increasing size are: micro-socks, fishing grounds and fishing area. Management scales (brackets) in increasing size are: territorial and regional. In this case is depicted as a local population could be divided between two adjacent territories fished and managed independently. differences between Atlantic and Mediterranean populations, but not among populations located in the same basin. This fact indicates the existence of at least two metapopulations, each one expanding 1000 s km2. The spatial scales involved in the fishing activities and management are detailed below. 4. Sea urchin fishery in Galicia Fishing strategies are adapted to the spatial structure of P. lividus, and consequently they show different operative scales. Fishers operate daily at very local scales, concentrating the fishing effort in micro-stocks where stock density is high [63]. The series of micro-stocks that are in short distance allowing a boat to exploit several of them in the same day is denominated fishing ground (Fig. 1). In this way the definition of ground is made by the fishers taking into account the distance among microstocks and the oceanographic conditions that allow the access to the zone. Following this criterion, a local population could include more than one ground. Indeed, this could be the case of a Galician locality, Lira, where fishers differentiate two grounds (north and south of Punta Remedios, Fig. 3) because the differences in the wind regimes allow exploiting one of these in alternative days. However habitat is similar in both grounds and micro-stocks (and patches) are dispersed along the whole area. These characteristics indicate that both grounds may be part of the same local population. Finally, the different grounds exploited by a fleet along the fishing season constitute a fishing area, and its expanse depends of the maximum distance that a boat can move from the home port. According to information provided by fishers, the fishing areas for sea urchins in Galicia are restricted by the TURFs defined by the FA (10 s km2), because the distance that boats used to move before the start of this regulation was larger than the distance allowed today. Nowadays Galicia is the main fishery for P. lividus, landing annually about 700 t [31]. Harvesting occurs mainly in subtidal areas by scuba diving, although the exploitation has been expanded to the intertidal in some Galician locations. The fleet is composed of 174 small boats ( o5 m long) that operate near the coastline and whose crew includes one skipper and 1 or 2 divers. Commercial fishing for sea urchin in Galicia started in the 1960s, but the regulation started only in 1986 [39]. The regulatory system has since become more sophisticated along the time and now it includes a closed season (from May to September), a daily quota per boat and fisher (100 kg per fisher until a maximum of 300 kg per boat), a daily timetable (from 9 to 15 h), a minimum commercial size (55 mm test diameter) and a depth limit for harvesting (o12 m). However, the main change in the fishing management was introduced in 1992 when the Fishing Authority of the regional government of Galicia (now denominated FA) assigned territorial rights to local fisher organisations (“Cofradías”) supervised by the regional government. In addition, the access to the fishery was restricted to a given number of boats using a licence system [65]. Nowadays there are 15 subtidal and 5 intertidal territories for the exploitation of sea urchin. Whereas the subtidal territories accomplish the whole coast, the intertidal ones are located only in the north and south extremes of the Galician coast (Fig. 2). Assuming a maximum harvesting depth of 20 m (according to their own commentaries and observations, divers frequently operate deeper than the 12-m legal limit), most of subtidal territories occupy a surface of approx. 30 km2, with a range from 11 to 130 km2. 4.1. Spatial structure of harvesting activities of P. lividus 4.2. Management scales for the P. lividus fishery in Galicia There are two management scales for the sea urchin fishery in Galicia. The regional one accomplishes the complete Galicia; the 16 R. Ouréns et al. / Marine Policy 51 (2015) 13–20 Fig. 2. Territories for the sea urchin fishery in Galicia in 2012. Grey rectangles represent territories defined in subtidal plans and coloured coastlines represent the 5 intertidal territories. Note that the intertidal territory 1 in the North of Galicia is discontinuous. Fig. 3. Spatial structure of the fishery of sea urchin, P. lividus, in Lira (Galicia). Dots represent the micro-stocks identified by Fernández-Boán et al. [63] and shadow areas are 5 fishing grounds as delimited by the fishers. territorial one is defined by the fishing zones delimited by the FA for this fishery. Regulations applied at the regional scale were presented at the beginning of this section, and they are devoted to control effort or catch: restrictions in the daily timetable and in the harvesting depth, daily quotas and minimum landing size. At territorial scale, there is a co-management between the FA and the fishers' associations. The fishers can propose additional regulations for their territories through the development of annual exploitation plans, which must include an assessment of the previous fishing season, a harvesting and marketing plan for the next season and a financial plan (see Molares and Freire [66] and Macho et al. [65] for details). Finally the FA has to assess the plan proposal and it has the authority to include modifications. Because there are not scientific assessments of the local stocks, exploitation plans use both basic landing data and the fisher knowledge obtained through their personal experience and their involvement in the fishery. In this way, plans use a trial-and-error approach and are adapted yearly using information about the results obtained in previous seasons and the successes and failures obtained by other Cofradías. This procedure could be defined as an informal and imperfect adaptive management system where the socioeconomic objectives play a key role with the same or higher priority respect to biological objectives (that actually could not be quantified due to the lack of assessments). Although TURFs constitute an opportunity to manage the resource at the local scale, Cofradías often do not propose additional regulations for their territories. This lack of proposals might be related with the fact that most of the territories are exploited jointly by various Cofradías (an average of 3.1 organisations per territory). In these cases the diversity in the interests of the different collectives causes differences and conflicts in their vision about how to manage the fishery and the specific regulations that should be implemented locally [39]. Consequently, complimentary regulations are proposed only in extreme situations when the productivity of the stocks have decreased dramatically, and in these cases fishers tend to propose regulations more restrictive over the catches or effort than those ones imposed by the FA. 5. Uncoupling of scales The comparison of the biological, fishery operations and management scales shows a clear uncoupling with relevant consequences for the fishery health (Fig. 4). Management scales are too wide precluding the adoption of regulations that allow managing effectively micro-stocks, which are the units that determine the fishery dynamics. In this way, over-exploitation of several micro-stocks would be possible before any signal of resource depletion was detected at the territorial scale. R. Ouréns et al. / Marine Policy 51 (2015) 13–20 17 Fig. 4. Diagram showing the mismatch between the units that compose the spatial structure of sea urchin, P. lividus, populations, fishing operations and management in Galicia. On the other hand, fishing territories have been defined by the FA using several criteria, such as the historical spatial pattern of fleet activity or the different eco-geographic regions characterised by specific oceanographic conditions. Nevertheless the spatial structure of sea urchin populations has not been taken into account, because most probably the importance of this fact in fishery management has been overlooked. This mismatch opened the possibility for local populations being divided between two fishing territories and consequently being exploited according to the different management plans running in each area. For instance, management plans for intertidal areas are in most cases independent from those for subtidal areas, although both stocks might be part of the same local population. Fig. 2 shows clearly another example of uncoupling of scales in sea urchin intertidal populations in northern Galicia. In this area the coastal sector 1, exploited jointly by 4 associations (San Cibrao, Burela, Ribadeo and San Cosme de Barreiros), is interrupted by the sector 2, exploited and managed in an independent way by another Cofradía (Celeiro). In all these cases in which different territories share the same local population, regulations introduced in a given territory are not going to produce the expected results because of the interference of fishing in adjacent territories. 6. How to adapt sea urchin management in Galicia to the relevant scales? All the facts discussed above should be a reason to review the spatial structure of the regulations of the sea urchin fishery in Galicia, in order to match management and biological scales. Firstly, fishing territories should be redefined to include complete local populations. In addition, the demarcation of individual territories for each Cofradía, or the generation of coordination processes between Cofradías sharing the same territory, could favour the introduction of effective regulations at territorial level. Nowadays management strategy is based in controlling how much and when to fish. Complimentary regulations at the territory scale should manage the distribution of effort in space and in this way assure the sustainability of the harvest of the smaller scale biological units. In this regard, micro-stocks should be a management scale because are both the minimal scale targeted by the fishing force and the smallest biological scale that is stable (patches are dynamical and therefore they are not practical for monitoring). Complimentarily, regulations should take into account Allee effects experienced by echinoid populations (Section 3), because they promote recruitment overfishing at low densities [67]. There are two types of spatial regulations that could be useful to manage the P. lividus fishery in Galicia: rotations and marine reserves. 6.1. Rotations Rotations are based in the delimitation of fishing subareas where harvest alternates. In this way each subarea shows consecutive periods of high and low stock density [68]. The basis of this strategy is that during temporal closures juveniles will attain the minimum commercial size entering the harvestable biomass and recovering the stock for the next fishing season. Due to the depensatory mechanisms that operate in P. lividus, population density in each micro-stock after a fishing season should remain at a level allowing effective reproduction and recruitment [68]. A challenge for future research is the assessment of this threshold density that could be a key indicator of the maximum level of harvesting that could be applied to a microstock of P. lividus. In this regard, Botsford et al. [68] estimated that densities higher than 0.7 ind m 2 allowed fertilisation success for S. franciscanus, using information from the experiments carried out by Levitan et al. [51]. Other relevant design aspect for rotations is the definition of the sizes of the subareas. If the spatial scale is too large fishers could overexploit several micro-stocks and continue to harvest others without any short-term signal of the decrease of the profitability of the subarea (Fig. 5). This is the reason because micro-stocks should be the units for rotations to allow controlling effectively density after a harvest pulse. Several studies of echinoids have simulated fishery yield using different rotation calendars [68–70]. Many of these studies took into account the possibility of density-dependent recruitment, but in any case the rotation scale was analysed despite its potential influence in the efficiency of the rotational system. For instance, 5 fishing areas to be exploited for 6 months every 3 years were established in the fishery of S. franciscanus in Washington. The size of each area was in the order of magnitude of 100 s km2 (see Fig. 1 of Lai and Bradbury [70]), and probably they comprised several micro-stocks and in some cases various local populations. Minimum and maximum landing sizes and a catch quota were imposed and the number of boats participating in the fishery was restricted. This management strategy was active from 1977 until 1995 but 18 R. Ouréns et al. / Marine Policy 51 (2015) 13–20 Fig. 5. Scenarios for the temporal dynamics of a local sea urchin population composed of 6 micro-stocks (circles) and managed using two alternative rotation systems. Boxes 1A and 1B represent the virgin population (black circles), whereas boxes 2 and 3 represent the population status after the first and second fishing seasons respectively. Relative stock density is represented by the colour of circles (virgin: black; fished: grey; over-exploited: white). Arrows indicate the spatial units exploited in each season. In the scenario A the rotation scale corresponds with the fishing ground, and consequently the harvesting rate of micro-stocks is not homogeneous: two of them reduced only slightly their size whereas the third is overexploited and it is unable to recover due to the inverse denso-dependency. In scenario B the rotation scale corresponds with the micro-stocks. The same stocks are exploited in 2B and 2A, but in B the exploitation is made in an alternative and regular way (i.e. 2 months every micro-stock). In this case all the micro-stocks decrease in size but they are able to recover for the second fishing season. catch decreased dramatically during this period [70], perhaps as a consequence of inadequate rotations. 6.2. Marine reserves The use of marine protected areas as tools for fisheries management has gained popularity in the last decades [71–74]. Besides protecting habitats and biodiversity, reserves could increase fishery yield and improve the stock sustainability in the long term due to two different mechanisms: (1) biomass spillover to adjacent, no protected, areas due to migrations or individual movements, and (2) increase the adult density and the production of eggs and larvae, which are transported by currents to the fishing areas [74–76]. The latter mechanism is of special relevance in echinoids and other species showing inverse denso-dependency in reproduction, due to the high densities that adults could attain in protected areas [77]. Moreover, reserves are a tool commonly suggested and used in species with metapopulation structure [46,78–80] because the protection of only one or a few local populations could increase the larval production for the complete metapopulation [46,81]. An alternative design for reserves devoted to protect species with metapopulation structure would consist of the protection of a small area inside each local population (Fig. 6). This option not only would increase fertilisation rates inside the reserve, but also would favour the biomass spillover to the fishing areas. This process would be not possible protecting a complete local population because adjacent populations are only connected by larval dispersal. In this sense, Quinn et al. [78] demonstrated using Fig. 6. Two potential designs for marine reserves devoted to the management of a resource with metapopulation structure. It is assumed that previously to the establishment of the network all local populations contribute similarly to the larval pool. Black ellipses represent micro-stocks in each local population and the shadowed areas are the protected zones. In the scenario A one whole local population is protected. The arrow width is proportional to the larval production, indicating that the protected population becomes key for the contribution of larvae to the metapopulation. In the scenario B a network of reserves is established and all local populations continue to provide similar larval productions. Discontinuous arrows represent migration of sea urchins towards fishing areas. simulation modelling that a reserve network could allow to attain sustainable harvesting rates preventing the collapse of S. franciscanus stocks. These authors suggested the establishment of several small-sized reserves separated by a distance shorter than the larval dispersal range. Because the recruitment of P. lividus occurs in shallow waters ( 5 m) and shows inverse density dependence [55], establishing multiple reserves in shallow habitats (intertidal and subtidal o5 m deep) could be an adequate regulation to promote larval production and recruitment survival for this species. Moreover, the migratory pattern of P. lividus heading to deeper areas [55,82] would secure the spillover of biomass towards the fishery areas. Marine reserves could be established protecting all microstocks located at o5 m, or protecting only a part of this shallow habitat in each local population. To assess both scenarios it is needed to know the proportion of habitat that should be protected to sustain the stocks, and that depends on the life history of the species (reproductive and growth rates, and larval dispersal distances and patterns) and on the harvesting rates. The study of Morgan and Botsford [83], which suggested a reserve system for S. franciscanus occupying approx. 35% of the area of the metapopulation, could be used as a preliminary estimation. This estimate was obtained using simulation models assuming uncertainty in fishery mortality rates and in the spatial patterns of larval dispersal. 7. Conclusion The present work shows that the conventional assessment and management methods, which ignore the spatial issues, are R. Ouréns et al. / Marine Policy 51 (2015) 13–20 not suitable for benthic resources with a complex spatial structure, because they do not allow to understand the spatial distribution of resources and to manage where to fish. Indeed, a common cause for failure in fisheries is the mismatch between the spatial scale of exploited populations and the scale of their assessment and management. Sea urchin fishery in Galicia is an example of this situation. Here, the operational and management units are uncoupled with the spatial population structure, promoting an ineffective governance system (based on TURFs) and the need for temporal closures of some local fisheries. The reason for this mismatch is that the biological structure of the resource has not been taken into account in the delimitation of the fishing territories, making possible the division of a population between two adjacent territories fished and managed independently. Because the spatial structure of populations have been overlooked in fishery management, this mismatch is also likely to occur in other benthic resources managed through TURFs in Galicia, such as goose barnacles, or razor clams. Future research should test this hypothesis and the dialogue between scientists, government and fishers should be promoted to solve the problem of fit. In addition, the metapopulation structure of sea urchins requires to be managed at multiple nested scales. The management in Galicia includes regional and territorial scales (being present at these scales social institutions responsible for management: Galician government and fishers' local associations), but a finer scale is also needed. Spatial regulatory measures should be established in each fishing territory, so that the fishing effort could be controlled at a micro-stock level (minimum biological level that determines the fishery dynamics), and the depensatory mechanisms affecting the recruitment and reproduction of P. lividus could be offset. 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