The species associated with this article are major components of the successionary communities arising around bathyal whale carcasses (though by no means the only whale-fall associated species).
A whale carcass arriving on the bathyal sea-floor (roughly 700-1000m depth) represents a massive influx of nutrients to an otherwise nutrient-poor ecosystem (Lundsten et al 2010a; Lundsten et al 2010b; Smith and Baco 2003). The background rate of carbon deposition to the deep-sea floor is on the order of tens of kilograms per hectare per year (Smith and Baco 2003); an adult whale can weigh up to 160 tons. Consequently, it has long been thought that whale carcasses must represent a significant source of nutrients for sea-bed communities. Additionally, since the discovery of deep-sea hydrothermal vents and cold seeps, it has been hypothesized that whale-falls may serve as stepping stones for the dispersal of organisms between chemosynthesis-dominated bottom communities (Smith and Baco 2003).
The community observed to spring up around whale carcasses has been characterized as having three major successionary stages (Danise et al 2012):
-Mobile scavenger stage: large, mobile detritivores consume the flesh of the whale.
-Enrichment opportunist stage: slow-moving or sessile organisms colonize the nutrient-enriched area in and around the carcass.
-Sulphophilic stage: a chemosynthesis-dominated system based on the sulfides released by anaerobic decomposition of bone lipids.
The duration of the first stage depends largely on the mass of the whale, ranging from a few months to up to one and a half years. Initially the community is dominated by large detritivores such as sleeper sharks and hagfish, but as the amount of flesh available decreases, smaller scavengers such as rattails, amphipods, and and lithodid crabs begin to replace them. Once the bulk of the tissue is removed from the skeleton, the community begins to shift to phase two. At this point, extremely dense populations of dorvilleid worms and other polychaetes, as well as crustaceans and gastropods colonize the area around the carcass, exploiting the rich organic material in the surrounding sediments. The rapid recruitment of these organisms suggests they may be opportunistic whale-fall specialists. Over time, without a discrete boundary, sulphide emission from anaerobic decay of bone lipids in the whale skeleton begins to support a chemosynthetic fauna similar to that found around cold seeps and hydrothermal vents, including bacteria, organisms with endosymbiotic bacteria, bacterial grazers, and small predators. This community may linger for up to several decades (Smith and Baco 2003). Fossil evidence suggests that a similar pattern of succession has been evolving since the late Miocene, and may even have operated on the carcasses of Cretaceous plesiosaurs (Danise et al 2012).
As always in ecology, this picture is somewhat oversimplified. In two 2010 articles, Lundsten et al observe that in addition to chemosynthetic fauna and whale-fall specialists, whale carcasses are often characterized by increased density of the background sea-floor organisms, particularly as time passes since the fall of the whale. Lundsten et al and Glover (2010) additionally found that there is a notable depth gradient in community structure, with fully sulphophilic ecosystems only developing on large, deep carcasses.
The function of whale-falls as stepping stones between cold seeps and hydrothermal vents remains unproven, but there is evidence for relatively large numbers of whale-fall specialist species, especially in the enrichment opportunist and sulphophilic stages (Smith and Baco 2003). Nearest-neighbor analyses of whale falls based on whale populations and the probability of a carcass sinking suggest that carcasses are distributed such that most organisms found in the latter two stages could easily disperse larvae between whale-fall sites. Unfortunately, this ecosystem may be endangered by declining whale populations and may even have already lost a great deal of diversity, as 19th century whale-fall density was likely up to six times higher than that in the present day (Smith and Baco 2003).
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