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Gooday, AJ, Bett BJ, Escobar E, Ingole B, Levin LA, Neira C, Raman AV, Sellanes J.  2010.  Habitat heterogeneity and its influence on benthic biodiversity in oxygen minimum zones. Marine Ecology-an Evolutionary Perspective. 31:125-147.   10.1111/j.1439-0485.2009.00348.x   AbstractWebsite

Oxygen minimum zones (OMZs; midwater regions with O(2) concentrations <0.5 ml l(-1)) are mid-water features that intercept continental margins at bathyal depths (100-1000 m). They are particularly well developed in the Eastern Pacific Ocean, the Arabian Sea and the Bay of Bengal. Based on analyses of data from these regions, we consider (i) how benthic habitat heterogeneity is manifested within OMZs, (ii) which aspects of this heterogeneity exert the greatest influence on alpha and beta diversity within particular OMZs and (iii) how heterogeneity associated with OMZs influences regional (gamma) diversity on continental margins. Sources of sea-floor habitat heterogeneity within OMZs include bottom-water oxygen and sulphide gradients, substratum characteristics, bacterial mats, and variations in the organic matter content of the sediment and pH. On some margins, hard grounds, formed of phosphorites, carbonates or biotic substrata, represent distinct subhabitats colonized by encrusting faunas. Most of the heterogeneity associated with OMZs, however, is created by strong sea-floor oxygen gradients, reinforced by changes in sediment characteristics and organic matter content. For the Pakistan margin, combining these parameters revealed clear environmental and faunal differences between the OMZ core and the upper and lower boundary regions. In all Pacific and Arabian Sea OMZs examined, oxygen appears to be the master driver of alpha and beta diversity in all benthic faunal groups for which data exist, as well as macrofaunal assemblage composition, particularly in the OMZ core. However, other factors, notably organic matter quantity and quality and sediment characteristics, come into play as oxygen concentrations begin to rise. The influence of OMZs on meiofaunal, macrofaunal and megafaunal regional (gamma) diversity is difficult to assess. Hypoxia is associated with a reduction in species richness in all benthic faunal groups, but there is also evidence for endemism in OMZ settings. We conclude that, on balance, OMZs probably enhance regional diversity, particularly in taxa such as Foraminifera, which are more tolerant of hypoxia than others. Over evolutionary timescales, they may promote speciation by creating strong gradients in selective pressures and barriers to gene flow.

Grosholz, ED, Levin LA, Tyler AC, Neira C.  2009.  Changes in community structure and ecosystem function following Spartina alterniflora invasion of Pacific estuaries. Human impacts on salt marshes : a global perspective. ( Silliman BR, Grosholz E, Bertness MD, Eds.).:23-40., Berkeley: University of California Press Abstract
Gutierrez, D, Gallardo VA, Mayor S, Neira C, Vasquez C, Sellanes J, Rivas M, Soto A, Carrasco F, Baltazar M.  2000.  Effects of dissolved oxygen and fresh organic matter on the bioturbation potential of macrofauna in sublittoral sediments off Central Chile during the 1997/1998 El Nino. Marine Ecology-Progress Series. 202:81-99.   10.3354/meps202081   AbstractWebsite

A study off Concepcion, central Chile, during the 1997/1998 El Nino (EN) revealed that the concentration of dissolved oxygen and the organic content and quality of the sediment control the vertical distribution of macrofauna in the sediment and bioturbation potential. The study area, characterized by organic-rich, silty sediments, lies within the most intense upwelling center off the coast of Chile, and is subject to the seasonal influx of hypoxic subsurface waters. Five stations (28 to 120 m depth) were sampled seasonally. The vertical distribution and integrated biomass and abundance of macrofauna (> 0.5 mm) were determined, as well as the dissolved oxygen content of the bottom water (BWDO) and sediment parameters such as total organic carbon (TOC), the C/N ratio, sulphide content, chi a content, and the thickness of the oxidized zone. Chi a proved to be a good indicator of fresh (high-quality) organic matter. Major components contributing to variation in the macrofauna feeding guilds, bioturbation categories, and their vertical position in the sediment were: (1) the relative bioturbation potential (contribution of bioturbating taxa to the assemblage) and (2) the vertical distribution and ratio of surface-to subsurface deposit-feeders. Higher levels of BWDO and a lower quality of organic matter at the sediment surface tended to provide better conditions for potentially strong bioturbators, while lower BWDO levels and higher-quality organic matter were accompanied by the dominance of tube-dwelling, surface-defecating land hence weakly bioturbating) species. Higher TOC levels and lower-quality organic matter at the surface resulted in deeper vertical distributions of animals and a higher relative abundance of subsurface deposit-feeders. During the study period, BWDO levels increased, while the total organic carbon and the quality of organic matter decreased. These conditions encouraged the vertical penetration of macrofauna into the sediment column and the relatively larger contribution of stronger bioturbators to the assemblage. The most drastic changes in faunal lifestyles and vertical distribution during the 1997/1998 EN were observed within the Bay of Concepcion, an area usually characterized by sulphidic sediments under the conditions of severe seasonal hypoxia or anoxia obtaining during 'normal' (i.e. non-EN) years; and in the deepest shelf site, which usually experiences permanent hypoxia because of the influence of the 'oxygen minimum zone'.