The role of the oceanic oxygen minima in generating biodiversity in the deep sea

Many studies on the deep-sea benthic biota have shown that the most species-rich areas lie on the continental margins between 500 and 2500 m, which coincides with the present oxygen-minimum in the world's oceans. Some species have adapted to hypoxic conditions in oxygen-minimum zones, and some can even fulfil all their energy requirements through anaerobic metabolism for at least short periods of time. It is, however, apparent that the geographic and vertical distribution of many species is restricted by the presence of oxygen-minimum zones. Historically, cycles of global warming and cooling have led to periods of expansion and contraction of oxygen-minimum layers throughout the world's oceans. Such shifts in the global distribution of oxygen-minimum zones have presented many opportunities for allopatric speciation in organisms inhabiting slope habitats associated with continental margins, oceanic islands and seamounts. On a smaller scale, oxygen-minimum zones can be seen today as providing a barrier to gene-flow between allopatric populations. Recent studies of the Arabian Sea and in other regions of upwelling also have shown that the presence of an oxygen-minimum layer creates a strong vertical gradient in physical and biological parameters. The reduced utilisation of the downward flux of organic material in the oxygen-minimum zone results in an abundant supply of food for organisms immediately below it. The occupation of this area by species exploiting abundant food supplies may lead to strong vertical gradients in selective pressures for optimal rates of growth, modes of reproduction and development and in other aspects of species biology. The presence of such strong selective gradients may have led to an increase in habitat specialisation in the lower reaches of oxygen-minimum zones and an increased rate of speciation