Human and climatic drivers affect spatial fishing patterns in a multiple-use marine protected area: The Galapagos Marine Reserve

Assessments of the effectiveness of marine protected areas (MPAs) usually assume that fishing patterns change exclusively due to the implementation of an MPA. This assumption increases the risk of erroneous conclusions in assessing marine zoning, and consequently counter-productive management actions. Accordingly, it is important to understand how fishers respond to a combination of the implementation of no-take zones, and various climatic and human drivers of change. Those adaptive responses could influence the interpretation of assessment of no-take zone effectiveness, yet few studies have examined these aspects. Indeed, such analysis is often unfeasible in developing countries, due to the dominance of data-poor fisheries, which precludes full examination of the social-ecological outcomes of MPAs. In the Galapagos Marine Reserve (Ecuador), however, the availability of long-term spatially explicit fishery monitoring data (1997–2011) for the spiny lobster fishery allows such an analysis. Accordingly, we evaluated how the spatiotemporal allocation of fishing effort in this multiple-use MPA was affected by the interaction of diverse climatic and human drivers, before and after implementation of no-take zones. Geographic information system modelling techniques were used in combination with boosted regression models to identify how these drivers influenced fishers’ behavior. Our results show that the boom-and-bust exploitation of the sea cucumber fishery and the global financial crisis 2007–09, rather than no-take zone implementation, were the most important drivers affecting the distribution of fishing effort across the archipelago. Both drivers triggered substantial macro-scale changes in fishing effort dynamics, which in turn altered the micro-scale dynamics of fishing patterns. Fishers’ adaptive responses were identified, and their management implications analyzed. This leads to recommendations for more effective marine and fishery management in the Galapagos, based on improved assessment of the effectiveness of no-take zones.


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Line 245: Table 1 R: Suggestion taken. Line 258: delete 'up' R: Suggestion taken. Line 263: 17,723 R: Suggestion taken. Line 272: 'from 2009' R: Suggestion taken. Line 274: Need to define CPUE at line 216, not here, so just use 'CPUE' R: Text edited. Line 283: delete the comma at the end of the line R: Suggestion taken. Line 290: Too many sub-headings, should delete some R: Suggestion taken. The number of sub-headings was reduced. Line 292: I would say 'The most potentially relevant' because there is no evidence these are relevant.
R: Suggestion taken. Text edited. Line 296: Again 'that potentially affected' R: Suggestion taken. Text edited. Line 307: 'boom-and-bust' R: Suggestion taken. Line 312: delete 'the Galapagos' as this is not necessary R: Suggestion taken. Line 313: delete the comma R: Suggestion taken. Line 317: Again, delete 'in Galapagos' R: Suggestion taken. Line 343: Don't need a part 1 and 2 for the sub-heading R: Suggestion taken. Sub-headings were edited. Line 346: abbreviation GIS has already been established and does not need to be established here again so just use 'GIS' R: Suggestion taken. Line 357: Delete the apostrophe R: Suggestion taken. Lines 370-371: Don't need a new sub-heading R: Suggestion taken. Sub-heading deleted. Line 374: Should be 'is concentrated' R: Suggestion taken. R: Suggestion taken. Line 1293: 'based on the core areas and distribution ranges of fishing fleets' R: Suggestion taken. Line 1303: Delete 'fishing fleets' R: Suggestion taken. Lines 1305-1315: Again, I find this text unnecessary and it could be deleted.
R: Text was deleted. Lines 1319-1323: This is a sentence, not a paragraph R: Text was edited. Line 1321: EBM has already been defined and does not need to be defined again here so just use EBM R: Text was edited. Line 1328: 'should not be taken for granted; MPAs are not a panacea. (delete the rest 'i.e. a one-size…..' R: Suggestion taken. Line 1332: 'with global climate change' R: Suggestion taken. Line 1336: delete 'learned' and write as 'is that assessments of MPA effectiveness' R: Suggestion taken.

Reviewer 1
This study evaluated how the spatiotemporal allocation of fishing effort for lobsters in the Galapagos multiple-use Marine Protected Area was affected by the interaction of diverse climatic and human drivers, before and after implementation of no-take zones. The study used GIS data on fishing effort and BRTs to attempt to identify how these drivers affected spatial fishing patterns. The paper concludes that the boom-and-bust exploitation of the sea cucumber fishery and the global financial crisis (2007)(2008)(2009), rather than no-take zone implementation, were the most important drivers affecting the distribution of fishing effort for lobsters across the archipelago. The study is spatially and temporally extensive (most of the Galapagos Islands, 1997-2011), the data are fairly well-analyzed and interpreted, and the manuscript is well-written. I have no major disagreements with the conclusions. I also have some sympathy with the suggestions that the MPA network placement could be revisited or even revised, and outside the network TURFS encouraged. My comments are mostly to assist the authors with publication.

Major Comments.
1. This is a paper that, in effect, quantifies spatial and temporal trends in fishing effort of a lobster fishery in a developing country. Yet the emphasis chosen is how this data informs effects of MPA implementation. It is highly commendable that the study includes before and after implementation data. In fact, this is such an important aspect of the study, I would recommend that the authors stress this point more in the paper. However, this MPA network is also well-known as a "classic" case where fishers ensured that no-take zones were NOT placed where fishers fished (Edgar et al 2004 Ref. 22 in this manuscript). That is, it is a case where you might NOT expect much change in spatial effort in the lobster fishery pre-and post-implementation of the MPA network (which is what they found). This very important point is not even mentioned until Lines 999-1010 in the Discussion. I recommend that you mention this much earlier in the paper, probably in the Abstract and Introduction. R: Suggestion taken. Both points suggested are mentioned in the Introduction.
2. You place a substantial amount of faith in the "explanatory" powers of your BRTs.
This needs to be tempered a fraction. 3. The spatial scale at which you measure effort (2.25 km 2 ) may be rather coarse to be making confident statements about the lack of evidence for "fishing the line". Many of the studies of spillover (see references cited below at Line 86) often report this effect at much smaller spatial scales than this. You should at least acknowledge this point. R: We agree that a finer scale probably would be needed to evaluate a spiny lobster spillover effect, but this is not the objective of this study. We evaluated finer and coarser spatial scales to conduct the hotspot analysis and the 2.25km 2 scale was the most proper scale to visualize the results and to evaluate the presence of a fishing the line effect around no-take zones. On the other hand, a fishing the line effect around the Galapagos Marine Reserve was detected by Bucaram at al. 2018 using a coarser scale of analysis (see "Assessing fishing effects inside and outside an MPA: The impact of the Galapagos Marine Reserve on the Industrial pelagic tuna fisheries during the first decade of operation"). Therefore, we think that a 2.25km 2 is an appropriate spatial scale of analysis for our case study.  (2015) and, even though they recognize that MPAs can be affected by diverse drivers of change, they do not evaluate their impact in a quantitative way as we did. Therefore, we edited the text in the following way: "…to our knowledge, no study has examined yet, in a quantitative way, how fishers respond to those situations in which they have to cope simultaneously with implementation of an MPA, and with the R: Suggestion taken. Text was edited.

Materials and Methods.
Line 198. "….and spillover to fishing grounds may occur ([26]" Lines 208, 209. Tourist or tourism (not touristic). R: Suggestion taken. Lines 273-274. Why calculate effort by dividing catch by catch-per-unit-effort (CPUE)? Surely you measured catch and effort directly to calculate CPUE? R: We estimated total fishing effort by dividing total catch by average CPUE per fishing season. We conducted this analysis to estimate the amount of sampling effort per fishing season. For all analysis presented in the paper, we used CPUE data estimated directly from catch and effort data per fishing trip. Line 302. ..affected by the potential drivers (add potential).
R: Suggestion taken. Lines 310-312. You make it clear that the re-zoning was confounded by the sea-cucumber over-exploitation phase (see also Table 3). Thus, when you talk of changes to effort associated with the zoning (e.g. Lines: 600-605, 989-990, 1035-1036) you must acknowledge this confounding. At lines 1096-1099 you DO acknowledge the confounding, and should in other places in the manuscript.
R: Suggestion taken.
R: Suggestion taken. Line 601. Acknowledge confounding of zoning and sea-cucumber over-exploitation phase.
R: Suggestions taken. Line 702. These types of fishers. Line 718. Suggest (not suggests). Line 888. Western side of R: Suggestion taken.

Discussion.
Lines 952-954. Note comment re Fletcher et al (2015) and Hughes et al (2016) above. R: Same response as in point 5 above. Lines1174-1176. Good point. The lobster recovery may not be related to the implementation of the MPAs.
R: No response needed. Lines 1188-1190 and 1218-1220. When suggesting a re-evaluation of the MPA zoning, you must be clear about why the MPAs were established: conservation, fisheries management, or both.
R: Suggestions taken. Text was edited as: "…we suggest re-evaluating the distribution of no-take zones across the GMR to promote the sustainability of the spiny lobster fishery and conserve key biodiversity areas". Lines 1194. The TURFS suggestion outside the MPAs is a good one. Lines 1199-1201. Alcala and Russ (2006) could be cited here also. R: Suggestions taken. Cite added. Lines 1224-1227. Why would an MPA network placed in a biased manner help the fishery if it was set up to avoid the fishery? R: We agree. However, there was no scientific evidence about the long-term impact of no-take zones on the fishing effort dynamic for the spiny lobster fishery before this study. Our study highlights the need to redistribute no-take zones to accomplish conservation and fishery management objectives. Line 1238. ..replicates.
R: Suggestion taken. Lines 1249-1262. In addition to the Kay example in the Channel Islands, which is a good one, you could also mention the Goni et al (2010) lobster example from the Mediterranean.
R: During the edition of the Discussion, we decided to eliminate Lines 1249-1262. We took this decision to put more emphasis in the discussion of other results directly associated to the objectives of our study. Line 1264. Ref. 70 in support of the idea of spillover is inadequate. See references to cite on spillover suggested above.
R: The academic editor suggested to eliminate Lines 1263-1270. We accepted this suggestion as we decide to put more emphasis in other results of the paper. Line 1291. To support (not the support).
R: Suggestion taken. Fig. 2. What do the dark grey and light grey shaded areas of time represent? El Nino/La Nina? Specify in caption.
R: Dark grey and light grey areas represents the periods analyzed. Figure 2 has been edited. The name of the periods was added. Caption was edited. Figs. 3 and 4. What are the units here? Effort (diver hours)? Specify in caption.
R: Standard deviation ellipses (SDE) polygons represent graphical summaries of the central tendency, dispersion and directional trends of fishing fleets. Core areas and distribution ranges refer to those areas covering 68% (1 SDE) and 95% (2 SDE) of the full spatial extent of fishing fleet distribution, respectively. Explanation is provided in Lines 324-327 of the revised manuscript.

General comments
In this study, the authors aim to investigate the effects of management, biophysical data and socioeconomic factors on the distribution of fishing effort. They use a variety of analytical tools to detect global and local drivers, from the Global Financial Crisis and climatic drivers to the distribution of MPAs. Given the need to better understand drivers of social and ecological dynamics, it will be good to see this paper published. There are two primary concerns that need to be addressed, however.
The first (and most serious) is that there is no mention of overfishing as a possible driver. This may be hard to measure, but in any boom-and-bust dynamic this must be one of the factors investigated. By reading this manuscript, the reader has no idea what kind of fishing effort the spiny lobster and sea cucumber populations in this area can sustainably endure. Ideally, the authors need to weave this consideration into the whole manuscript, and if there is no way of adding actual data on this, they need to make a substantial effort to include information from other studies. R: As the objective of this study was to predict fishing effort distribution rather than catch or catch-per-unit-effort (CPUE), we focused our analysis on the human element (effort), rather than the interaction between humans and the target species themselves (catch or CPUE). This approach helped us to simplify the interpretation of the results and more accurately predict fishing effort. On the other hand, as overfishing of the spiny lobster fishery is the consequence of the external drivers of change analyzed in this study, particularly of the boom-and-bust exploitation of the sea cucumber fishery, we did not consider overfishing explicitly as a driver of change. Nevertheless, we analyzed the factors influencing fishing effort, including the previous lobster catch and sea cucumber revenues, which were relevant as fishing effort predictors. These two predictors were affected by the overexploitation of the sea cucumber and spiny lobster fisheries. We edited the text to highlight this fact in the manuscript (Lines 1988-1991 of the revised manuscript). In addition, we explained in the Discussion that the spiny lobster fishery was overexploited due to the overcapitalization caused by the expansion of the sea cucumber fishery and explained the consequences of overexploitation on fishing capacity (Lines 1858-1853).