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  1. The deep sea (below 200 m depth) is the largest carbon sink on Earth. It hosts abundant biodiversity that underpins the carbon cycle and provides provisioning, supporting, regulating and cultural ecosystem services. There is growing attention to climate-regulating ocean ecosystem services from the scientific, business and political sectors. In this essay we synthesize the unique biophysical, socioeconomic and governance characteristics of the deep sea to critically assess opportunities for deep-sea blue carbon to mitigate climate change. Deep-sea blue carbon consists of carbon fluxes and storage including carbon transferred from the atmosphere by the inorganic and organic carbon pumps to deep water, carbon sequestered in the skeletons and bodies of deep-sea organisms, carbon buried within sediments or captured in carbonate rock. However, mitigating climate change through deep-sea blue carbon enhancement suffers from lack of scientific knowledge and verification, technological limitations, potential environmental impacts, a lack of cooperation and collaboration, and underdeveloped governance. Together, these issues suggest that deep-sea climate change mitigation is limited. Thus, we suggest that a strong focus on blue carbon is too limited a framework for managing the deep sea to contribute to international goals, including the Sustainable Development Goals (SDGs), the Paris Agreement and the post-2020 Biodiversity Goals. Instead, the deep sea can be viewed as a more holistic nature-based solution, including many ecosystem services and biodiversity in addition to climate. Environmental impact assessments (EIAs), area-based management, pollution reduction, moratoria, carbon accounting and fisheries management are tools in international treaties that could help realize benefits from deep-sea, nature-based solutions. 
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    Free, publicly-accessible full text available July 6, 2024
  2. Organisms in coastal waters experience naturally high oxygen variability and steep oxygen gradients with depth, in addition to ocean deoxygenation. They often undergo diel vertical migration involving a change in irradiance that initiates a visual behavior. Retinal function has been shown to be highly sensitive to oxygen loss; here we assess whether visual behavior (photobehavior) in paralarvae of the squid Doryteuthis opalescens and the octopus Octopus bimaculatus is affected by low oxygen conditions, using a novel behavioral paradigm. Larvae showed an irradiance-dependent, descending photobehavior after extinction of the light stimulus, measured through the change in vertical position of larvae in the chamber. The magnitude of photobehavior was decreased as oxygen was reduced, and the response was entirely gone at <6.4 kPa partial pressure of oxygen (<74.7 umol kg-1 at 15.3 7C) in D. opalescens paralarvae. Oxygen also affected photobehavior in O. bimaculatus paralarvae. The mean vertical velocity of paralarvae was unaffected by exposure to reduced oxygen, indicating that oxygen deficits selectively affect vision prior to locomotion. These findings suggest that variable and declining oxygen conditions in coastal upwelling areas and elsewhere will impair photobehavior and likely affect the distribution, migration behavior, and survival of highly visual marine species. 
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  3. Vulnerability of marine species to climate change (including ocean acidification, deoxygenation, and associated changes in food supply) depends on species’ ecological and biological characteristics. Most existing assessments focus on coastal species but systematic analysis of climate vulnerability for the deep sea is lacking. Here, we combine a fuzzy logic expert system with species biogeographical data to assess the risks of climate impacts to the population viability of 32 species of exploited demersal deep-sea species across the global ocean. Climatic hazards are projected to emerge from historical variabilities in all the recorded habitats of the studied species by the mid-twenty-first century. Species that are both at very high risk of climate impacts and highly vulnerable to fishing include Antarctic toothfish (Dissostichus mawsoni), rose fish (Sebastes norvegicus), roughhead grenadier (Macrourus berglax), Baird’s slickhead (Alepocephalus bairdii), cusk (Brosme brosme), and Portuguese dogfish (Centroscymnus coelepis). Most exploited deep-sea fishes are likely to be at higher risk of local, or even global, extinction than previously assessed because of their high vulnerability to both climate change and fishing. Spatially, a high concentration of deep-sea species that are climate vulnerable is predicted in the northern Atlantic Ocean and the Indo-Pacific region. Aligning carbon mitigation with improved fisheries management offers opportunities for overall risk reduction in the coming decades. Regional fisheries management organizations (RFMOs) have an obligation to incorporate climate change in their deliberations. In addition, deep-sea areas that are not currently managed by RFMOs should be included in existing or new international governance institutions or arrangements. 
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  4. Deep-sea chemosynthesis-based ecosystems support unique biological communities, but human impacts are an increasing threat. Understanding the life-history traits of species from deep-sea chemosynthesis-based ecosystems can help to develop adequate management strategies, as these can have impacts on ecological responses to changes in the environment. Here we examined the occurrence of sexual dimorphism in the yeti crab Kiwa puravida , an endemic species from the Costa Rican Pacific margin that aggregates at active methane seeps and depends on chemosynthetic bacteria for nutrition. The two morphological features examined included the claws, suspected to be under sexual selection and used for defense, and the carpus of the second pereopod not suspected to be under sexual selection. A total of 258 specimens, 161 males, 81 females, 16 juveniles, were collected from Mound 12 at 1,000-1,040 m depth in 2017 and 2018 and analyzed. We found that males have larger and wider claws than females, while there were no differences in carpus length. These results suggest that claw weaponry is under sexual selection in K . puravida , which is probably related to the mating system of this deep-sea species. This is the first attempt to study the reproductive biology of K . puravida , and additional observations will be necessary to shed more light on this matter. 
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  5. Global policy goals for halting biodiversity loss and climate change depend on each other to be successful. Marine biodiversity and climate change are intertwined through foodwebs that cycle and transport carbon and contribute to carbon sequestration. Yet, biodiversity conservation and fisheries management seldom explicitly include ocean carbon transport and sequestration. In order to effectively manage and govern human activities that affect carbon cycling and sequestration, international biodiversity and climate agreements need to address both biodiversity and climate issues. International agreements that address issues for climate and biodiversity are best poised to facilitate the protection of ocean carbon with existing policies. The degree to which the main international biodiversity and climate agreements make reference to multiple issues has however not been documented. Here, we used a text mining analysis of over 2,700 binding and non-binding policy documents from ten global ocean-related agreements to identify keywords related to biodiversity, climate, and ocean carbon. While climate references were mostly siloed within climate agreements, biodiversity references were included in most agreements. Further, we found that six percent of policy documents (n=166) included ocean carbon keywords. In light of our results, we highlight opportunities to strengthen the protection of ocean carbon in upcoming negotiations of international agreements, and via area-based management, environmental impact assessment and strategic environmental assessment. 
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  6. Organisms in coastal waters experience naturally high oxygen variability and steep oxygen gradients with depth, in addition to ocean deoxygenation. They often undergo diel vertical migration involving a change in irradiance that initiates a visual behavior. Retinal function has been shown to be highly sensitive to oxygen loss; here we assess whether visual behavior (photobehavior) in paralarvae of the squid Doryteuthis opalescens and the octopus Octopus bimaculatus is affected by low oxygen conditions, using a novel behavioral paradigm. Larvae showed an irradiance-dependent, descending photobehavior after extinction of the light stimulus, measured through the change in vertical position of larvae in the chamber. The magnitude of photobehavior was decreased as oxygen was reduced, and the response was entirely gone at <6.4 kPa partial pressure of oxygen (<74.7 mmol kg21 at 15.3 7C) in D. opalescens paralarvae. Oxygen also affected photobehavior in O. bimaculatus paralarvae. The mean vertical velocity of paralarvae was unaffected by exposure to reduced oxygen, indicating that oxygen deficits selectively affect vision prior to locomotion. These findings suggest that variable and declining oxygen conditions in coastal upwelling areas and elsewhere will impair photobehavior and likely affect the distribution, migration behavior, and survival of highly visual marine species. 
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  7. Vermeij, Geerat J. (Ed.)
    Continental margins host methane seeps, animal falls and wood falls, with chemosynthetic communities that may share or exchange species. The goal of this study was to examine the existence and nature of linkages among chemosynthesis-based ecosystems by deploying organic fall mimics (bone and wood) alongside defaunated carbonate rocks within high and lesser levels of seepage activity for 7.4 years. We compared community composition, density, and trophic structure of invertebrates on these hard substrates at active methane seepage and transition (less seepage) sites at Mound 12 at ~1,000 m depth, a methane seep off the Pacific coast of Costa Rica. At transition sites, the community composition on wood and bone was characteristic of natural wood- and whale-fall community composition, which rely on decay of the organic substrates. However, at active sites, seepage activity modified the relationship between fauna and substrate, seepage activity had a stronger effect in defining and homogenizing these communities and they depend less on organic decay. In contrast to community structure, macrofaunal trophic niche overlap between substrates, based on standard ellipse areas, was greater at transition sites than at active sites, except between rock and wood. Our observations suggest that whale- and wood-fall substrates can function as stepping stones for seep fauna even at later successional stages, providing hard substrate for attachment and chemosynthetic food. 
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