Invasion by marine nonindigenous species (NIS) is a spread phenomenon. The tunicate Pyura praeputialis shows pronounced disjoint geographical distribution: along thousands of kilometers in wave-swept headlands on the southeastern coast of Australia, from where it appears to have originated, and exclusively along 60 -70 km inside the Bay of Antofagasta, Chile. mtDNA sequences suggested that the species invaded this rocky shore recently. We used field manipulations and juvenile P. praeputialis transplant techniques to test hypotheses regarding the capacity of the tunicate to survive and grow at different sites and tidal heights inside and outside Antofagasta, and its competitive performance for primary space (inside the Bay) against the native mussel Perumytilus purpuratus. We conclude that survival and growth of P. praeputialis showed no significant differences among sites inside and outside the Bay, and suggest that the restrictive distribution of the species in Chile is caused by a specific oceanographic retention mechanism and͞or its brief larval dispersal. We demonstrated that, inside the Bay, P. praeputialis outcompetes Perumytilus from the Mid-Low intertidal, constraining Perumytilus to the Upper MidIntertidal, modifying the local pattern of intertidal zonation. We show that predation on P. praeputialis juveniles by starfish and snails constitutes a regulatory mechanism for the setting of its low intertidal limit. Major ecological impacts caused by NIS invasions to rocky shores by aggressive primary space users may result in negative aspects, but also may contribute to biodiversity enhancement. We call attention to the need for increment manipulations and testing of ecological hypotheses regarding marine NIS.

BackgroundBiological invasions affecting rocky intertidal zonation patterns, yield information on species interactions. In the Bay of Antofagasta, northern Chile, the non-indigenous tunicate Pyura praeputialis, originally from Australia, has invaded (in the past century or so) and monopolized a major portion of the mid-intertidal rocky shore, displacing upshore the native mussel Perumytilus purpuratus. In Antofagasta the tunicate is subjected to intensive exploitation. Monitoring protocols show that in the past 10 years Antofagasta's tunicate population has experienced a drastic decline, affecting the intertidal zonation pattern.Methodology/Principal FindingsA 12.5 km of coastline, on the southern eastern shore of the Bay of Antofagasta, was studied. Eight sites were systematically (1993–1994) or sporadically (2003–2014) monitored for the seaward-shoreward expansion or reduction of the tunicate Pyura praeputialis, and native mussel and barnacle bands. A notable reduction in the mid-intertidal band of P. praeputialis and a seaward expansion of the mussel, Perumytilus purpuratus, and barnacle bands was observed. We suggest that the major cause for the decline in the tunicate is due to its intensive exploitation by rocky shore Pyura-gathers. The rate of extraction of tunicates by professional Pyura-gathers ranged between 256–740 tunicates hour−1. Between 2009–2014 the density of professional Pyura-gather ranged between 0.5–4.5 km−1 per low tide. Hence, 10 professional Pyura-gathers working 1 h for 10 low tides per month, during 6 months, will remove between 307–888 m2 of tunicates. A drastic decline in tunicate recruitment was observed and several P. praeputialis ecosystems services have been lost.Conclusion and SignificanceIn Antofagasta, the continuous and intensive intertidal gathering of the invasive tunicate Pyura praeputialis, has caused a drastic reduction of its population modifying the zonation pattern. Thereby, native mussel Perumytilus purpuratus has regained its ecological center in the intertidal zone. We recorded a Pyura recruitment failure and loss of ecosystem services.

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The continued degradation of marine ecosystems, along with the ecosystem services they provide, suggest that new, innovative approaches are needed to scale up marine biodiversity protection and promote sustainable fishery practices. We synthesize information from Chile on the key processes involved in the development of alternative strategies for scaling up marine biodiversity conservation and discuss the complementarities with marine protected areas. Defined as "ancillary" marine conservation initiatives under the Convention of Biological Diversity, we suggest that these alternative strategies have the potential to capitalize on local stakeholders' participation and contribute to solving livelihood and governance issues while playing a significant role in scaling up marine conservation. We specifically focus on two recent ancillary initiatives being piloted in Chile. The development of business model innovations which could enable biodiversity benefits from territorial user rights fisheries policies and the creation of municipal conservation areas. We identify how these initiatives could eventually help scale up marine conservation, discuss opportunities and challenges from these pilot experiences and conclude with the need for developing policy frameworks and cross-scale governance approaches which formally acknowledge marine ancillary conservation measures as part of an integrated way to manage marine biodiversity. Exploring and supporting alternative complementary marine conservation strategies is particularly relevant in Chile and Latin America, if biodiversity conservation initiatives are to scale in coverage, contribute to livelihood improvement of local communities, replenish fisheries and play key roles in adaptation to climate change.