It had been assumed for a century before that the deep-sea fauna was depauperate, and prior to then that the great depths were essentially sterile. These ideas were largely a carry over from Edward Forbes in the late 1800's, who proposed the azoic hypothesis for the deep sea. Interestingly, his idea was largely based on samples from the Aegean Sea now known to have relatively low densities of organisms compared to other deep-sea areas. The lack of deep-sea life was overturned by several later reports of deep-sea species attached to sounding lines and the dredging cruises of the H.M.S. Lighting, Porcupine, and Challenger. In 1968, Sander's comparative study of marine benthic diversity, showed that bathyal diversity exceeded coastal diversity in the temperate zone and approached shallow-water tropical diversity. The development and deployment of more effective sampling gear has indicated that diversity is probably even higher than Sanders estimated, potentially rivaling that of tropical rain forests. On relatively small scales, the number of species coexisting in the deep sea is surprisingly high, exceeding 300 macrofaunal species m-2 at bathyal depths in the western North Atlantic. The greater diversity in what appears to be a more homogeneous environment has long perplexed marine ecologists and remains a major theoretical challenge. Numerous hypotheses have been proposed including competition, facilitation, predation, disturbance, productivity, environmental heterogeneity and patch dynamics.
Further papers on deep-sea diversity (in no particular order)
Etter RJ, Grassle JF (1992) Patterns of species diversity in the deep sea as a function of sediment particle size diversity. Nature 360:576-578
Gray JS (1994) Is deep-sea species diversity really so high: species diversity of the Norwegian continental shelf. Marine Ecological Progress Series 112:205-209
Lambshead PJD, Boucher G (2003) Marien nematode deep-sea biodiversity-hyperdiverse or hype? Journal of Biogeography 30:475-485
Cronin TM, Raymo ME (1997) Orbital forcing of deep-sea benthic species diversity. Nature 385:624-627
Gage JD, Tyler PA (1991) Deep-Sea Biology: A Natural History of Organisms at the Deep-Sea Floor, Vol. Cambridge University Press, Cambridge
Gage J, Lambshead PJD, Bishop JDD, Stuart CT, Jones NS (2004) Large-scale biodiversity pattern of Cumacea (Percarida: Crustacean) in the deep Atlantic. Marine Ecological Progress Series 277:181-196
Gooday AJ, Bett BJ, Shire R, Lambshead PJD (1998) Deep-sea benthic foraminiferal species diversity in the NE Atlantic and NW Arabian sea: a synthesis. Deep-Sea Research II 45:165-201
Snelgrove PVR, Smith CR (2002) A riot of species in an environmental calm: the paradox of the species-rich deep-sea floor. Oceanography and Marine Biology 40:211-242
Rex MA (1981) Community structure in the deep-sea benthos. Annual Review of Ecology and Systematics 12:331
Rex MA (1983) Geographic patterns of species diversity in the deep-sea benthos. In: Rowe GT (ed) The Sea. Wiley, New York, p 453-472
Stuart CT, Rex MA, Etter RJ (2003) Large-scale spatial and temporal patterns of deep-sea benthic species diversity. In: Tyler PA (ed) Ecosystems of the World-Ecosystems of Deep Oceans. Elsevier, Amsterdam
Levin LA, Etter RJ, Rex MA, Gooday AJ, Smith CR, Pineda J, Stuart CT (2001) Environmental influences on regional deep-sea species diversity. Annual Review of Ecology and Systematics 32:51-93
Gage J (1998) Why are there so many species in deep-sea sediments. Journal of Experimental Marine Biology and Ecology 200:257-286
Sanders HL (1968) Marine benthic diversity: a comparative study. American Naturalist 102:243-282
Grassle JF (1989) Species diversity in deep-sea communities. Trends in Ecology and Evolution 4:12-15
Rex MA, McClain CR, Johnson NA, Etter RJ, Allen JA, Bouchet P, Waren A (2005) A source-sink hypothesis for abyssal biodiversity. American Naturalist 165:163-178
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