Global conservation outcomes depend on marine protected areas with five key features.

In line with global targets agreed under the Convention on Biological Diversity, the number of marine protected areas (MPAs) is increasing rapidly, yet socio-economic benefits generated by MPAs remain difficult to predict and under debate. MPAs often fail to reach their full potential as a consequence of factors such as illegal harvesting, regulations that legally allow detrimental harvesting, or emigration of animals outside boundaries because of continuous habitat or inadequate size of reserve. Here we show that the conservation benefits of 87 MPAs investigated worldwide increase exponentially with the accumulation of five key features: no take, well enforced, old (>10 years), large (>100 km(2)), and isolated by deep water or sand. Using effective MPAs with four or five key features as an unfished standard, comparisons of underwater survey data from effective MPAs with predictions based on survey data from fished coasts indicate that total fish biomass has declined about two-thirds from historical baselines as a result of fishing. Effective MPAs also had twice as many large (>250 mm total length) fish species per transect, five times more large fish biomass, and fourteen times more shark biomass than fished areas. Most (59%) of the MPAs studied had only one or two key features and were not ecologically distinguishable from fished sites. Our results show that global conservation targets based on area alone will not optimize protection of marine biodiversity. More emphasis is needed on better MPA design, durable management and compliance to ensure that MPAs achieve their desired conservation value.

Global human footprint on the linkage between biodiversity and ecosystem functioning in reef fishes.

Difficulties in scaling up theoretical and experimental results have raised controversy over the consequences of biodiversity loss for the functioning of natural ecosystems. Using a global survey of reef fish assemblages, we show that in contrast to previous theoretical and experimental studies, ecosystem functioning (as measured by standing biomass) scales in a non-saturating manner with biodiversity (as measured by species and functional richness) in this ecosystem. Our field study also shows a significant and negative interaction between human population density and biodiversity on ecosystem functioning (i.e., for the same human density there were larger reductions in standing biomass at more diverse reefs). Human effects were found to be related to fishing, coastal development, and land use stressors, and currently affect over 75% of the world's coral reefs. Our results indicate that the consequences of biodiversity loss in coral reefs have been considerably underestimated based on existing knowledge and that reef fish assemblages, particularly the most diverse, are greatly vulnerable to the expansion and intensity of anthropogenic stressors in coastal areas.

Caulerpa chemnitzia in Darwin threatening Galapagos coral reefs.

Coral reefs are rare in the Galapagos and there is concern that, like in many areas around the world, they may be degrading due to increasing anthropogenic pressure, which can cause changes and reorganizations of structure and function with associated phase shifts. Algae of the genus Caulerpa J.V. Lamouroux, 1809 are known as widespread and persistent marine invaders. They grow rapidly, particularly in disturbed areas where they can opportunistically monopolize substratum and compete with native species, thus reducing biodiversity. Caulerpa chemnitzia increased in abundance and overgrew corals on the reef since 2012, ultimately raising fears that a phase-shift from coral to algae might be imminent. However, from 2019 onwards algae populations strongly contracted and while not having returned to baseline level, there is currently low risk of corals being displaced. Visual censuses were conducted on a yearly basis since 2004 using sample quadrats (0.5 x 0.5m) every 5 m along a 50-m-long transects at a depth of 6-15 m at 5 permanent subtidal ecological monitoring sites around Darwin. In addition, 10 m photo-transects were taken using a graduated meter-long measuring stick in the centre of the frame in 2012, 2014, 2016, 2017, 2018 and 2021 at a depth of 15m at Wellington reef. The authors hypothesize that this species could have expanded its distribution over Wellington Reef because of its known morphological plasticity due to a response to change in the environment, in this case high temperature and low nutrients. As ENSO events are predicted to increase in intensity and frequency due to the impact of climate change it is important to develop and implement a functional alert system. Early Detection Rapid Response (EDRR) protocols are recommended to avoid climate driven Non-Indigenous Species (NIS) entering the GMR or for native species becoming invasive due to warming-related phase shifts.

Some environmental and biological determinants of coral richness, resilience and reef building in Galápagos (Ecuador).

Throughout the Galápagos, differences in coral reef development and coral population dynamics were evaluated by monitoring populations from 2000-2019, and environmental parameters (sea temperatures, pH, NO3-, PO43-) from 2015-19. The chief goal was to explain apparent coral community differences between the northern (Darwin and Wolf) and southern (Sta. Cruz, Fernandina, San Cristóbal, Española, Isabela) islands. Site coral species richness was highest at Darwin and Wolf. In the three most common coral taxa, a declining North (N)-South (S) trend in colony sizes existed for Porites lobata and Pocillopora spp., but not for Pavona  spp. Frequent coral recruitment was observed in all areas. Algal competition was highest at Darwin, but competition by bioeroding sea urchins and burrowing fauna (polychaete worms, bivalve mollusks) increased from N to S with declining coral skeletal density. A biophysical model suggested strong connectivity among southern islands with weaker connectivity to Wolf and even less to Darwin. Also, strong connectivity was observed between Darwin and Wolf, but from there only intermittently to the south. From prevailing ocean current trajectories, coral larvae from Darwin and Wolf drift primarily towards Malpelo and Cocos Islands, some reaching Costa Rica and Colombia. Mean temperature, pH, and PO43- declined from N to S. Strong thermocline shoaling, especially in the warm season, was observed at most sites. A single environmental factor could not explain the variability in observed coral community characteristics, with minimum temperature, pH and nutrient levels the strongest determinants. Thus, complex environmental determinants combined with larval connectivity patterns may explain why the northern Galápagos Islands (Darwin, Wolf) have higher coral richness and cover and also recover more rapidly than central/southern islands after region-wide disturbances. These northern islands are therefore potentially of critical conservation importance as important reservoirs of regional coral biodiversity and source of larvae.

State of corals and coral reefs of the Galápagos Islands (Ecuador): Past, present and future.

Coral populations and structural coral reefs have undergone severe reductions and losses respectively over large parts of the Galápagos Islands during and following the 1982-83 El Niño event. Coral tissue loss amounted to 95% across the Archipelago. Also at that time, all coral reefs in the central and southern islands disappeared following severe degradation and eventual collapse due primarily to intense bioerosion and low recruitment. Six sites in the southern islands have demonstrated low to moderate coral community (scattered colonies, but no carbonate framework) recovery. The iconic pocilloporid reef at Devil's Crown (Floreana Island) experienced recovery to 2007, then severe mortality during a La Niña cooling event, and is again (as of 2017) undergoing rapid recovery. Notable recovery has occurred at the central (Marchena) and northern islands (Darwin and Wolf). Of the 17 structural reefs first observed in the mid-1970s, the single surviving reef (Wellington Reef) at Darwin Island remains in a positive growth mode. The remainder either degraded to a coral community or was lost. Retrospective analyses of the age structure of corals killed in 1983, and isotopic signatures of the skeletal growth record of massive corals suggest the occurrence of robust coral populations during at least a 500-year period before 1983. The greatest potential threats to the recovery and persistence of coral reefs include: ocean warming and acidification, bioerosion, coral diseases, human population growth (increasing numbers of residents and tourists), overfishing, invasive species, pollution, and habitat destruction. Such a diverse spectrum of disturbances, acting alone or in combination, are expected to continue to cause local and archipelago-wide mortality and degradation of the coral reef ecosystem.

SeaWiFS satellite monitoring of oil spill impact on primary production in the Galápagos Marine Reserve.

Near daily satellite monitoring of ocean colour using sea viewing wide angle of field viewing sensor (SeaWiFS) allowed the oceanic and near coastal chlorophyll-a distributions to be followed across the Galápagos Marine Reserve (GMR) from space. In the aftermath of the Jessica spill early indications suggested that, compared to the three preceding years 1998-2000, local chlorophyll concentrations over January 2001 were elevated across the Galápagos Marine Reserve [Biological Impacts of the Jessica Oil Spill on the Galápagos Environment: Preliminary Report. Charles Darwin Foundation, Puerto Ayora, Galápagos, Ecuador, 2001]. At the time of the spill the central and eastern extent of the archipelago was experiencing a spatially extensive moderate bloom event (0.5-2.5 mgm(-3) chl-a) extending over the central islands, including the source of the spill and areas of known impact such as the islands of Santa Fé, eastern Santa Cruz and Floreana directly in the advection path.Further investigation shows that chlorophyll across the affected regions of western San Cristóbal, Santa Fé, southeast Santa Cruz, eastern Floreana and eastern Isabela declined in the week directly following the spill event, yet rose in the successive month to levels analogous to preceding years. Although there may have been a localised effect of the spill upon near coast phytoplankton primary production in the short term, the observed variance in the weeks following the spill was not significant in comparison to the normal high variation between years and within the El Niño/Southern Oscillation signal.

One-third of reef-building corals face elevated extinction risk from climate change and local impacts.

The conservation status of 845 zooxanthellate reef-building coral species was assessed by using International Union for Conservation of Nature Red List Criteria. Of the 704 species that could be assigned conservation status, 32.8% are in categories with elevated risk of extinction. Declines in abundance are associated with bleaching and diseases driven by elevated sea surface temperatures, with extinction risk further exacerbated by local-scale anthropogenic disturbances. The proportion of corals threatened with extinction has increased dramatically in recent decades and exceeds that of most terrestrial groups. The Caribbean has the largest proportion of corals in high extinction risk categories, whereas the Coral Triangle (western Pacific) has the highest proportion of species in all categories of elevated extinction risk. Our results emphasize the widespread plight of coral reefs and the urgent need to enact conservation measures.

Deep-water kelp refugia as potential hotspots of tropical marine diversity and productivity.

Classic marine ecological paradigms view kelp forests as inherently temperate-boreal phenomena replaced by coral reefs in tropical waters. These paradigms hinge on the notion that tropical surface waters are too warm and nutrient-depleted to support kelp productivity and survival. We present a synthetic oceanographic and ecophysiological model that accurately identifies all known kelp populations and, by using the same criteria, predicts the existence of >23,500 km(2) unexplored submerged (30- to 200-m depth) tropical kelp habitats. Predicted tropical kelp habitats were most probable in regions where bathymetry and upwelling resulted in mixed-layer shoaling above the depth of minimum annual irradiance dose for kelp survival. Using model predictions, we discovered extensive new deep-water Eisenia galapagensis populations in the Galápagos that increased in abundance with increasing depth to >60 m, complete with cold-water flora and fauna of temperate affinities. The predictability of deep-water kelp habitat and the discovery of expansive deep-water Galápagos kelp forests validate the extent of deep-water tropical kelp refugia, with potential implications for regional productivity and biodiversity, tropical food web ecology, and understanding of the resilience of tropical marine systems to climate change.