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Figure 4. Factor Analysis of Mixed Data. The eight most important attributes (six significant and two tentative
as detected by random forests and Boruta) detected for overall management success (OMS) were used as active
variables. The OMS was added as a supplementary variable. Black filled squares indicate centroids of each factor
levels. Coloured points indicate individual MPAs, colours indicating MPA success score. White-filled squares
with coloured outlines indicate centroids of MPAs sharing the same OMS and ellipses indicate centroids 95%
confidence intervals.

no-take zone could provide both ecological and socio-economic benefits. Larger no-take zones protect a larger
portion of fish stocks and promote density dependent spill-over processes thus enhancing fisheries catches
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in the buffer zones 17,24,54 . On the other hand restrictions granting fishing rights exclusively to local fishermen
could provide additional benefits and increase OMS. From this perspective combining Territorial Use Rights for
Fisheries (TURFs) with MPAs could further increase the perceived legitimacy of management decisions by pro-
viding fishermen with “ownership” of local fisheries resources. Examples of benefits related to TURFs are available
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globally , however this management tool remains largely under-utilised in the Mediterranean.
Interestingly, leadership among fishermen was not a determinant to successful management of SSF. This is in
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contrast with results from fisheries of other regions and also seems counterintuitive given the need for fisher-
men engagement. The presence of a leader among fishermen can potentially facilitate dialogue between fisher-
men and MPA management bodies, therefore enhancing fishermen engagement into SSF management. However,
inconsistency in our results could be specific to MPAs where management bodies can catalyse the action of the
fishermen community toward support in management, mimicking the role usually acted by fishermen leaders.
Highly successful MPAs (i.e. OMS = 3) show high similarity in the five most important attributes (as denoted
by their clear grouping in Factor Analysis of Mixed Data; Fig. 4) with moderate correlations among four attrib-
utes (i.e. fishermen engagement, presence of fishermen in management board, presence of a management plan and
presence of activity promoting sustainable fisheries) (Supplementary Figure S1). Conversely, MPAs with lower OMS
have large variability in conditions determining their lack of success. This suggests that a wide range of conditions
can result in unsuccessful/moderately successful SSF management in MPAs, while a very specific combination of
circumstances can determine successful cases. Because success in our study cases was determined mainly by five
shared attributes, and few studies had subsets of these five attributes, we are unable to determine if subsets would
result in successful SSF-MPA management. However, the presence of these five attributes may be inherent of
MPA management (e.g. promotion activities of sustainable fisheries is likely to be supported when a management
plan is in place and fishermen are engaged) where good practices can facilitate the onset of other good practices.
To quantify tangible benefits delivered by successful MPAs, we collected extensive field data at one successful
case study (Torre Guaceto MPA, Italy) by using underwater visual census and monitoring of SSF catch landings.
At this MPA, four of the five key attributes have been implemented except for presence of a fishermen represent-
ative on the MPA management board. At Torre Guaceto the implementation of these four key attributes led to:
a (i) 428% increase in total fish biomass in NTZ compared to external, unprotected areas, (ii) 128% increase in
fishermen revenues when they operate within the MPA buffer zone compared to fishing outside the MPA, and
(iii) dramatic increase in the commitment of local fishermen to environmental issues (i.e. the number of fines
for illegal fishing after MPA establishment dropped to nearly zero after the implementation of the key features.
Furthermore, the fishermen now participate in research and environmental programs).
Single-species management strategies generally employed for large-scale fisheries are not suited for the
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dynamic context of SSFs. However, alternative management strategies for SSF have been missing because success-
ful examples were yet to be characterised. Here, we identify five key attributes that are suitable to manage SSFs.
Significant economic and social commitments are required to implement the key attributes we have highlighted.

Scientific RepoRts | 6:38135 | DOI: 10.1038/srep38135 6