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Maloney, JM, Grupe BM, Pasulka AL, Dawson KS, Case DH, Frieder CA, Levin LA, Driscoll NW.  2015.  Transpressional segment boundaries in strike-slip fault systems offshore southern California: Implications for fluid expulsion and cold seep habitats. Geophysical Research Letters. 42:4080-4088.   10.1002/2015gl063778   AbstractWebsite

The importance of tectonics and fluid flow in controlling cold seep habitats has long been appreciated at convergent margins but remains poorly understood in strike-slip systems. Here we present geophysical, geochemical, and biological data from an active methane seep offshore from Del Mar, California, in the inner California borderlands (ICB). The location of this seep appears controlled by localized transpression associated with a step in the San Diego Trough fault zone and provides an opportunity to examine the interplay between fluid expulsion and restraining step overs along strike-slip fault systems. These segment boundaries may have important controls on seep locations in the ICB and other margins characterized by strike-slip faulting (e.g., Greece, Sea of Marmara, and Caribbean). The strike-slip fault systems offshore southern California appear to have a limited distribution of seep sites compared to a wider distribution at convergent plate boundaries, which may influence seep habitat diversity and connectivity.

Marlow, JJ, Steele JA, Ziebis W, Thurber AR, Levin LA, Orphan VJ.  2014.  Carbonate-hosted methanotrophy represents an unrecognized methane sink in the deep sea. Nature Communications. 5   10.1038/ncomms6094   AbstractWebsite

The atmospheric flux of methane from the oceans is largely mitigated through microbially mediated sulphate-coupled methane oxidation, resulting in the precipitation of authigenic carbonates. Deep-sea carbonates are common around active and palaeo-methane seepage, and have primarily been viewed as passive recorders of methane oxidation; their role as active and unique microbial habitats capable of continued methane consumption has not been examined. Here we show that seep-associated carbonates harbour active microbial communities, serving as dynamic methane sinks. Microbial aggregate abundance within the carbonate interior exceeds that of seep sediments, and molecular diversity surveys reveal methanotrophic communities within protolithic nodules and well-lithified carbonate pavements. Aggregations of microbial cells within the carbonate matrix actively oxidize methane as indicated by stable isotope FISH-nanoSIMS experiments and (CH4)-C-14 radiotracer rate measurements. Carbonate-hosted methanotrophy extends the known ecological niche of these important methane consumers and represents a previously unrecognized methane sink that warrants consideration in global methane budgets.

McCann, LD, Levin LA.  1989.  Oligochaete influence on settlement, growth and reproduction in a surface-deposit-feeding polychaete. Journal of Experimental Marine Biology and Ecology. 131:233-253.   10.1016/0022-0981(89)90115-9   AbstractWebsite

The functional-group hypothesis predicting that a mobile subsurface-deposit feeder would inhibit the recruitment and subsequent development of a more sedentary tubiculous surface-deposit feeder was tested through a series of laboratory and field manipulations of the oligochaete Monopylephorus evertus Brinkhurst and the polychaete Streblospio benedicti Webster. Laboratory and field studies indicated that settlement of S. benedicti was unaffected by the presence of M. evertus. Settlement of syllid, capitellid and other spionid polychaetes, gastropods and Gemma gemma Totten also appeared to be unaffected. In laboratory cultures, M. evertus caused decreased survivorship of S. benedicti from age 11 wk through senescence and decreased growth (addition of setigers) at age 6–11 wk. Growth of newly settled and older worms may be decreased by the presence of M. evertus, however, the experimental design had insufficient power to determine significance when treatment differences were small. Reduced growth of juveniles, leading to smaller size, was predicted to cause decreased reproductive output in S. benedicti although size-specific reproductive capacity of individual worms was unaffected. Results are consistent with functional-group theory predicting mobile burrowing forms to have negative effects on more stationary tubiculous forms. However, significant decreases in growth rate were limited to the juvenile (recruitment) stages of S. benedicti, emphasizing the importance of examining the entire life cycle of the species of interest. This study also highlights the need for future investigations to determine the role of marine oligochaetes in the dynamics of salt-marsh and estuarine communities.

McCormick, LR, Levin LA, Oesch NW.  2019.  Vision is highly sensitive to oxygen availability in marine invertebrate larvae. Journal of Experimental Biology. 222   10.1242/jeb.200899   AbstractWebsite

For many animals, evolution has selected for complex visual systems despite the high energetic demands associated with maintaining eyes and their processing structures. Therefore, the metabolic demands of visual systems make them highly sensitive to fluctuations in available oxygen. In the marine environment, oxygen changes over daily, seasonal and inter-annual time scales, and there are large gradients of oxygen with depth. Vision is linked to survival in many marine animals, particularly among the crustaceans, cephalopods and fish, and early life stages of these groups rely on vision for prey capture, predator detection and their distribution in the water column. Using in vivo electroretinogram recordings, we show that there is a decrease in retinal sensitivity to light in marine invertebrates when exposed to reduced oxygen availability. We found a 60-100% reduction in retinal responses in the larvae of cephalopods and crustaceans: the market squid (Doryteuthis opalescens), the two-spot octopus (Octopus bimaculatus), the tuna crab (Pleuroncodes planipes) and the graceful rock crab (Metacarcinus gracilis). A decline in oxygen also decreases the temporal resolution of vision in D. opalescens. These results are the first demonstration that vision in marine invertebrates is highly sensitive to oxygen availability and that the thresholds for visual impairment from reduced oxygen are species-specific. Oxygen-impaired retinal function may change the visual behaviors crucial to survival in these marine larvae. These findings may impact our understanding of species' vulnerability to ocean oxygen loss and suggest that researchers conducting electrophysiology experiments should monitor oxygen levels, as even small changes in oxygen may affect the results.

McCormick, LR, Levin LA.  2017.  Physiological and ecological implications of ocean deoxygenation for vision in marine organisms. Philosophical Transactions of the Royal Society a-Mathematical Physical and Engineering Sciences. 375   10.1098/rsta.2016.0322   AbstractWebsite

Climate change has induced ocean deoxygenation and exacerbated eutrophication-driven hypoxia in recent decades, affecting the physiology, behaviour and ecology of marine organisms. The high oxygen demand of visual tissues and the known inhibitory effects of hypoxia on human vision raise the questions if and how ocean deoxygenation alters vision in marine organisms. This is particularly important given the rapid loss of oxygen and strong vertical gradients in oxygen concentration in many areas of the ocean. This review evaluates the potential effects of low oxygen (hypoxia) on visual function in marine animals and their implications for marine biota under current and future ocean deoxygenation based on evidence from terrestrial and a few marine organisms. Evolutionary history shows radiation of eye designs during a period of increasing ocean oxygenation. Physiological effects of hypoxia on photoreceptor function and light sensitivity, in combination with morphological changes that may occur throughout ontogeny, have the potential to alter visual behaviour and, subsequently, the ecology of marine organisms, particularly for fish, cephalopods and arthropods with `fast' vision. Visual responses to hypoxia, including greater light requirements, offer an alternative hypothesis for observed habitat compression and shoaling vertical distributions in visual marine species subject to ocean deoxygenation, which merits further investigation. This article is part of the themed issue 'Ocean ventilation and deoxygenation in a warming world'.

Mehring, AS, Hatt BE, Kraikittikun D, Orelo BD, Rippy MA, Grant SB, Gonzalez JP, Jiang SC, Ambrose RF, Levin LA.  2016.  Soil invertebrates in Australian rain gardens and their potential roles in storage and processing of nitrogen. Ecological Engineering. 97:138-143.   10.1016/j.ecoleng.2016.09.005   AbstractWebsite

Research on rain gardens generally focuses on hydrology, geochemistry, and vegetation. The role of soil invertebrates has largely been overlooked, despite their well-known impacts on soil nutrient storage, removal, and processing. Surveys of three rain gardens in Melbourne, Australia, revealed a soil invertebrate community structure that differed significantly among sites but was stable across sampling dates (July 2013 and April 2014). Megadrilacea (earthworms), Enchytraeidae (potworms), and Collembola (springtails) were abundant in all sites, and together accounted for a median of 80% of total soil invertebrate abundance. Earthworms were positively correlated to soil organic matter content, but the abundances of other taxonomic groups were not strongly related to organic matter content, plant cover, or root biomass across sites. While less than 5% of total soil N was estimated to be stored in the body tissues of these three taxa, and estimated N gas emissions from earthworms (N2O and N-2) were low, ingestion and processing of soil was high (e.g., up to 417% of the upper 5 cm of soil ingested by earthworms annually in one site), suggesting that the contribution of these organisms to N cycling in rain gardens may be substantial. Thus, invertebrate communities represent an overlooked feature of rain garden design that can play an important role in the structure and function of these systems. (C) 2016 Elsevier B.V. All rights reserved.

Mehring, AS, Levin LA.  2015.  Potential roles of soil fauna in improving the efficiency of rain gardens used as natural stormwater treatment systems. Journal of Applied Ecology. 52:1445-1454.   10.1111/1365-2664.12525   AbstractWebsite

Natural treatment systems such as rain gardens aim to overcome the negative effects of urbanization on water quality, availability, and freshwater and marine ecosystem integrity by mimicking the natural water cycle in urban planning and design. While soils in these systems are inhabited by a diverse array of invertebrates, the soil macrofauna is ignored in the vast majority of studies on new or existing rain gardens. Here, we review the functional roles of invertebrates commonly found within soils of rain gardens. Soil fauna have the potential to substantially alter plant growth, water infiltration rates, and the retention and removal of pathogens, nutrients, heavy metals and other contaminants. Their lack of inclusion in controlled laboratory or greenhouse studies may contribute to differences in observed function in field and laboratory settings. Promising future research directions include the following: (i) the use of controlled experiments to study invertebrate effects on rain garden function; (ii) determining the factors affecting variability in organismal abundance among and within sites; and (iii) the design of rain gardens to facilitate development of fauna that promote desired functions.Synthesis and applications. Soil fauna may substantially alter the function of rain gardens as natural stormwater treatment systems in urban areas. Therefore, incorporat-ing animal effects into design and testing may better enable managers and researchers tounderstand and optimize rain garden functioning, and forecast the longevity of rain gardens. Soil fauna may substantially alter the function of rain gardens as natural stormwater treatment systems in urban areas. Therefore, incorporat-ing animal effects into design and testing may better enable managers and researchers tounderstand and optimize rain garden functioning, and forecast the longevity of rain gardens.

Mengerink, KJ, Vandover CL, Ardron J, Baker M, Escobar-Briones E, Gjerde K, Koslow J, Ramirez-Llodra E, Lara-Lopez A, Squires D, Sutton T, Sweetman A, Levin LA.  2014.  A Call For Deep-Ocean Stewardship. Science. 344:696-698.
Middelburg, JJ, Levin LA.  2009.  Coastal hypoxia and sediment biogeochemistry. Biogeosciences. 6:1273-1293. AbstractWebsite

The intensity, duration and frequency of coastal hypoxia (oxygen concentration < 63 mu M) are increasing due to human alteration of coastal ecosystems and changes in oceanographic conditions due to global warming. Here we provide a concise review of the consequences of coastal hypoxia for sediment biogeochemistry. Changes in bottom-water oxygen levels have consequences for early diagenetic pathways (more anaerobic at expense of aerobic pathways), the efficiency of re-oxidation of reduced metabolites and the nature, direction and magnitude of sediment-water exchange fluxes. Hypoxia may also lead to more organic matter accumulation and burial and the organic matter eventually buried is also of higher quality, i.e. less degraded. Bottom-water oxygen levels also affect the organisms involved in organic matter processing with the contribution of metazoans decreasing as oxygen levels drop. Hypoxia has a significant effect on benthic animals with the consequences that ecosystem functions related to macrofauna such as bio-irrigation and bioturbation are significantly affected by hypoxia as well. Since many microbes and microbial-mediated biogeochemical processes depend on animal-induced transport processes (e.g. re-oxidation of particulate reduced sulphur and denitrification), there are indirect hypoxia effects on biogeochemistry via the benthos. Severe long-lasting hypoxia and anoxia may result in the accumulation of reduced compounds in sediments and elimination of macrobenthic communities with the consequences that biogeochemical properties during trajectories of decreasing and increasing oxygen may be different (hysteresis) with consequences for coastal ecosystem dynamics.

Mora, C, Wei CL, Rollo A, Amaro T, Baco AR, Billett D, Bopp L, Chen Q, Collier M, Danovaro R, Gooday AJ, Grupe BM, Halloran PR, Ingels J, Jones DOB, Levin LA, Nakano H, Norling K, Ramirez-Llodra E, Rex M, Ruhl HA, Smith CR, Sweetman AK, Thurber AR, Tjiputra JF, Usseglio P, Watling L, Wu TW, Yasuhara M.  2013.  Biotic and human vulnerability to projected changes in ocean biogeochemistry over the 21st century. Plos Biology. 11   10.1371/journal.pbio.1001682   AbstractWebsite

Ongoing greenhouse gas emissions can modify climate processes and induce shifts in ocean temperature, pH, oxygen concentration, and productivity, which in turn could alter biological and social systems. Here, we provide a synoptic global assessment of the simultaneous changes in future ocean biogeochemical variables over marine biota and their broader implications for people. We analyzed modern Earth System Models forced by greenhouse gas concentration pathways until 2100 and showed that the entire world's ocean surface will be simultaneously impacted by varying intensities of ocean warming, acidification, oxygen depletion, or shortfalls in productivity. In contrast, only a small fraction of the world's ocean surface, mostly in polar regions, will experience increased oxygenation and productivity, while almost nowhere will there be ocean cooling or pH elevation. We compiled the global distribution of 32 marine habitats and biodiversity hotspots and found that they would all experience simultaneous exposure to changes in multiple biogeochemical variables. This superposition highlights the high risk for synergistic ecosystem responses, the suite of physiological adaptations needed to cope with future climate change, and the potential for reorganization of global biodiversity patterns. If co-occurring biogeochemical changes influence the delivery of ocean goods and services, then they could also have a considerable effect on human welfare. Approximately 470 to 870 million of the poorest people in the world rely heavily on the ocean for food, jobs, and revenues and live in countries that will be most affected by simultaneous changes in ocean biogeochemistry. These results highlight the high risk of degradation of marine ecosystems and associated human hardship expected in a future following current trends in anthropogenic greenhouse gas emissions.

Moseman, SM, Levin LA, Currin C, Forder C.  2004.  Colonization, succession, and nutrition of macrobenthic assemblages in a restored wetland at Tijuana Estuary, California. Estuarine Coastal and Shelf Science. 60:755-770.   10.1016/j.ecss.2004.03.013   AbstractWebsite

Modes of colonization, the successional trajectory, and trophic recovery of a macrofaunal community were analyzed over 19 months in the Friendship marsh, a 20-acre restored wetland in Tijuana Estuary, California. Traditional techniques for quantifying macrofaunal communities were combined with emerging stable isotopic approaches for evaluation of trophic recovery, making comparisons with a nearby natural Spartina foliosa habitat. Life history-based predictions successfully identified major colonization modes, although most taxa employed a variety of tactics for colonizing the restored marsh. The presence of S.foliosa did not seem to affect macrofaunal colonization or succession at the scale of this study. However, soil organic matter content in the restored marsh was positively correlated with insect densities, and high initial salinities may have limited the success of early colonists. Total macrofaunal densities recovered to natural marsh levels after 14 months and diversity, measured as species richness and the Shannon index (H'), was comparable to the natural marsh by 19 months. Some compositional disparities between the natural and created communities persisted after 19 months, including lower percentages of surface-feeding polychaetes (Polydora spp.) and higher percentages of dipteran insects and turbellarians in the Friendship marsh. As surficial structural similarity of infaunal communities between the Friendship and natural habitat was achieved, isotopic analyses revealed a simultaneous trajectory towards recovery of trophic structure. Enriched delta(13)C signatures of benthic microalgae and infauna, observed in the restored marsh shortly after establishment compared to natural Spartina habitat, recovered after 19 months. However, the depletion in delta(15)N signatures of macrofauna in the Friendship marsh indicated consumption of microalgae, particularly nitrogen-fixing cyanobacteria, while macroalgae and Spartina made a larger contribution to macrofaunal diets in the natural habitat. Future successional studies must continue to develop and employ novel combinations of techniques for evaluating structural and functional recovery of disturbed and created habitats. (C) 2004 Elsevier Ltd. All rights reserved.

Moseman, SM, Zhang R, Qian PY, Levin LA.  2009.  Diversity and functional responses of nitrogen-fixing microbes to three wetland invasions. Biological Invasions. 11:225-239.   10.1007/s10530-008-9227-0   AbstractWebsite

Impacts of invasive species on microbial components of wetland ecosystems can reveal insights regarding functional consequences of biological invasions. Nitrogen fixation (acetylene reduction) rates and diversity of nitrogen fixers, determined by genetic fingerprinting (T-RFLP) of the nifH gene, were compared between native and invaded sediments in three systems. Variable responses of nitrogen fixing microbes to invasion by a non-native mussel, Musculista senhousia, and mangrove, Avicennia marina, in Kendall Frost-Northern Wildlife Preserve (Mission Bay) and salt cedar, Tamarisk (Tamarix spp.) in Tijuana Estuary suggest microbes respond to both species- and site-specific influences. Structurally similar invaders (the mangrove and salt cedar) produced different effects on activity and diversity of nitrogen fixers, reflecting distinct environmental contexts. Despite relative robustness of microbial community composition, subtle differences in total diversity or activity of nitrogen fixers reveal that microbes are not immune to impacts of biological invasions, and that functional redundancy of microbial diversity is limited, with significant consequences for functional dynamics of wetlands.

Moseman-Valtierra, S, Levin LA, Martin RM.  2016.  Anthropogenic impacts on nitrogen fixation rates between restored and natural Mediterranean salt marshes. Marine Ecology-an Evolutionary Perspective. 37:370-379.   10.1111/maec.12289   AbstractWebsite

To test the effects of site and successional stage on nitrogen fixation rates in salt marshes of the Venice Lagoon, Italy, acetylene reduction assays were performed with Salicornia veneta- and Spartina townsendii-vegetated sediments from three restored (6-14years) and two natural marshes. Average nitrogen fixation (acetylene reduction) rates ranged from 31 to 343 mu mol C2H4.m(-2.)h(-1) among all marshes, with the greatest average rates being from one natural marsh (Tezze Fonde). These high rates are up to six times greater than those reported from Southern California Spartina marshes of similar Mediterranean climate, but substantially lower than those found in moister climates of the Atlantic US coast. Nitrogen fixation rates did not consistently vary between natural and restored marshes within a site (Fossei Est, Tezze Fonde, Cenesa) but were negatively related to assayed plant biomass within the acetylene reduction samples collected among all marshes. Highest nitrogen fixation rates were found at Tezze Fonde, the location closest to the city of Venice, in both natural and restored marshes, suggesting possible site-specific impacts of anthropogenic stress on marsh succession.

Moseman-Valtierra, SM, Armaiz-Nolla K, Levin LA.  2010.  Wetland response to sedimentation and nitrogen loading: diversification and inhibition of nitrogen-fixing microbes. Ecological Applications. 20:1556-1568.   10.1890/08-1881.1   AbstractWebsite

Anthropogenic inputs of nutrients and sediment simultaneously impact coastal ecosystems, such as wetlands, especially during storms. Independent and combined effects of sediment and ammonium nitrate loading on nitrogen fixation rates and diversity of microbes that fix nitrogen (diazotrophs) were tested via field manipulations in Spartina foliosa and unvegetated zones at Tijuana Estuary (California, USA). This estuary is subject to episodic nitrogen enrichment and sedimentation associated with rain-driven flooding and slope instabilities, the latter of which may worsen as the Triple Border Fence is constructed along the U.S.-Mexico border. Responses of diazotrophs were assessed over 17 days using acetylene reduction assays and genetic fingerprinting (terminal restriction fragment length polymorphism [T-RFLP]) of nifH, which codes for dinitrogenase reductase. Sulfate-reducing bacteria performed similar to 70% of nitrogen fixation in Spartina foliosa rhizospheres in the absence of nitrogen loading, based on sodium molybdate inhibitions in the laboratory. Following nutrient additions, richness (number of T-RFs [terminal restriction fragments]) and evenness (relative T-RF fluorescence) of diazotrophs in surface sediments increased, but nitrogen fixation rates decreased significantly within 17 days. These responses illustrate, within a microbial community, conformance to a more general ecological pattern of high function among assemblages of low diversity. Diazotroph community composition (T-RF profiles) and rhizosphere diversity were not affected. Pore water ammonium concentrations were higher and more persistent for 17 days in plots receiving sediment additions (1 cm deep), suggesting that recovery of diazotroph functions may be delayed by the combination of sediment and nutrient inputs. Nitrogen fixation constitutes a mechanism for rapid transfer of fixed N to S. foliosa roots and a variety of primary consumers (within 3 and 8 days, respectively), as determined via (15)N(2) enrichment studies with in situ microcosms of intact marsh sediment. Thus, long-term declines in nitrogen fixation rates in response to increasingly frequent nutrient loading and sedimentation may potentially alter nitrogen sources for vascular plants as well as trophic pathways in wetland ecosystems.

Moy, LD, Levin LA.  1991.  Are Spartina marshes a replaceable resource? A functional approach to evaluation of marsh creation efforts Estuaries. 14:1-16.   10.2307/1351977   AbstractWebsite

Marsh creation has come into increasing use as a measure to mitigate loss of valuable wetlands. However, few programs have addressed the functional ecological equivalence of man-made marshes and their natural counterparts. This study addresses structural and functional interactions in a man-made and two natural marshes. This was done by integrating substrate characteristics and marsh utilization by organisms of two trophic levels. Sediment properties, infaunal community composition, and Fundulus heteroclitus marsh utilization were compared for a man-made Spartina salt marsh (between ages 1 to 3 yr) in Dills Creek, North Carolina, and adjacent natural marshes to the east and west. East natural marsh and planted marsh sediment grain-size distributions were more similar to each other than to the west natural marsh due to shared drainage systems, but sediment organic content of the planted marsh was much lower than in either natural marsh. This difference was reflected in macrofaunal composition. Natural marsh sediments were inhabited primarily by subsurface, deposit-feeding oligochaetes whereas planted marsh sediments were dominated by the tube-building, surface-deposit feeding polychaetes Streblospio benedicti and Manayunkia aestuarina. Infaunal differences were mirrored in Fundulus diets. Natural marsh diets contained more detritus and insects because oligochaetes, though abundant, were relatively inaccessible. Polychaetes and algae were major constituents of the planted marsh Fundulus diet. Though natural-marsh fish may acquire a potentially less nutritive, detritus-based diet relative to the higher animal protein diet of the planted marsh fish, Fundulus abundances were markedly lower in the planted marsh than in the natural marshes, indicating fewer fish were being supported. Lower Spartina stem densities in the planted marsh may have provided inadequate protection from predation or insufficient spawning sites for the fundulids. After three years, the planted marsh remained functionally distinct from the adjacent natural marshes. Mitigation success at Dills Creek could have been improved by increasing tidal flushing, thereby enhancing access to marine organisms and by mulching with Spartina wrack to increase sediment organic-matter content and porosity. Results from this study indicate that salt marshes should not be treated as a replaceable resource in the short term. The extreme spatial and temporal variability inherent to salt marshes make it virtually impossible to exactly replace a marsh by planting one on another site.

Mullineaux, LS, Metaxas A, Beaulieu SE, Bright M, Gollner S, Grupe BM, Herrera S, Kellner JB, Levin LA, Mitarai S, Neubert MG, Thurnherr AM, Tunnicliffe V, Watanabe HK, Won YJ.  2018.  Exploring the ecology of deep-sea hydrothermal vents in a metacommunity framework. Frontiers in Marine Science. 5   10.3389/fmars.2018.00049   AbstractWebsite

Species inhabiting deep-sea hydrothermal vents are strongly influenced by the geological setting, as it provides the chemical-rich fluids supporting the food web, creates the patchwork of seafloor habitat, and generates catastrophic disturbances that can eradicate entire communities. The patches of vent habitat host a network of communities (a metacommunity) connected by dispersal of planktonic larvae. The dynamics of the metacommunity are influenced not only by birth rates, death rates and interactions of populations at the local site, but also by regional influences on dispersal from different sites. The connections to other communities provide a mechanism for dynamics at a local site to affect features of the regional biota. In this paper, we explore the challenges and potential benefits of applying metacommunity theory to vent communities, with a particular focus on effects of disturbance. We synthesize field observations to inform models and identify data gaps that need to be addressed to answer key questions including: (1) what is the influence of the magnitude and rate of disturbance on ecological attributes, such as time to extinction or resilience in ametacommunity; (2) what interactions between local and regional processes control species diversity, and (3) which communities are "hot spots" of key ecological significance. We conclude by assessing our ability to evaluate resilience of vent metacommunities to human disturbance (e.g., deep-sea mining). Although the resilience of a few highly disturbed vent systems in the eastern Pacific has been quantified, these values cannot be generalized to remote locales in the western Pacific ormid Atlantic where disturbance rates are different and information on local controls is missing.