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The ecology of rafting in the marine environment - Bedim

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MARTIN THIEL & LARS GUTOW<br />

that future comparisons should focus on biological traits <strong>of</strong> <strong>raft<strong>in</strong>g</strong> organisms and on ecological<br />

<strong>in</strong>teractions.<br />

Geographic and seasonal variations <strong>in</strong> <strong>raft<strong>in</strong>g</strong> communities<br />

<strong>The</strong> abundance and species composition <strong>of</strong> <strong>raft<strong>in</strong>g</strong> communities varies throughout <strong>the</strong> world’s<br />

oceans. This is partly due to <strong>the</strong> global variations <strong>in</strong> availability <strong>of</strong> different <strong>raft<strong>in</strong>g</strong> substrata (see<br />

Thiel & Gutow 2004). However, <strong>the</strong>re are also regional differences <strong>in</strong> o<strong>the</strong>r <strong>environment</strong>al variables<br />

(e.g., nutrient availability, primary productivity and temperature) that affect <strong>the</strong> presence and<br />

persistence <strong>of</strong> <strong>raft<strong>in</strong>g</strong> organisms on float<strong>in</strong>g items.<br />

Regional differences <strong>in</strong> <strong>the</strong> Sargassum community had been recognised early on. Timmermann<br />

(1932) emphasised that <strong>the</strong> nor<strong>the</strong>rn part <strong>of</strong> <strong>the</strong> Sargasso Sea, i.e., <strong>the</strong> area <strong>of</strong> <strong>the</strong> Gulf Stream,<br />

features a higher diversity <strong>of</strong> <strong>raft<strong>in</strong>g</strong> organisms than <strong>the</strong> sou<strong>the</strong>rn part. Conover & Sieburth (1964)<br />

revealed that bacterial population densities on float<strong>in</strong>g Sargassum <strong>in</strong> <strong>the</strong> North Atlantic varied<br />

geographically accord<strong>in</strong>g to <strong>the</strong> specific production <strong>of</strong> chemical antifoul<strong>in</strong>g mechanisms. Stoner &<br />

Green<strong>in</strong>g (1984) provided a multifactorial comparison <strong>of</strong> <strong>the</strong> fauna associated with float<strong>in</strong>g Sargassum<br />

from <strong>the</strong> Sargasso Sea and <strong>the</strong> Gulf Stream and found only low similarities between faunal<br />

assemblages from <strong>the</strong>se regions. Sargasso Sea samples showed a relatively even distribution <strong>of</strong><br />

species, with <strong>the</strong> gastropod Litiopa melanostoma dom<strong>in</strong>at<strong>in</strong>g <strong>the</strong> community with a mean proportion<br />

<strong>of</strong> about 25% <strong>of</strong> <strong>the</strong> total fauna. Gulf Stream samples, however, were slightly less diverse and<br />

strongly dom<strong>in</strong>ated by <strong>the</strong> shrimp Latreutes fucorum compris<strong>in</strong>g about 70% <strong>of</strong> <strong>the</strong> Sargassum fauna<br />

<strong>in</strong> that region. Consistently, <strong>the</strong> Gulf Stream fauna yielded a higher percentage <strong>of</strong> large omnivores<br />

(sensu Butler et al. 1983), while <strong>in</strong> <strong>the</strong> Sargasso Sea smaller omnivores consum<strong>in</strong>g primarily sessile<br />

prey organisms were more abundant. Similarly, Smith et al. (1973) found higher numbers <strong>of</strong> <strong>raft<strong>in</strong>g</strong><br />

species <strong>in</strong> <strong>the</strong> Slope Water (33 species) than <strong>in</strong> <strong>the</strong> Sargasso Sea (about 25 species). However,<br />

densities <strong>of</strong> <strong>the</strong> animals <strong>in</strong> <strong>the</strong> Sargasso Sea (~4200 <strong>in</strong>d. g –1 wet weight [WW] Sargassum) were<br />

about twice as high as <strong>in</strong> <strong>the</strong> Slope Water (~2000 <strong>in</strong>d. g –1 WW Sargassum). In <strong>the</strong> same region,<br />

Keller (1987) found that <strong>the</strong> biomass <strong>of</strong> <strong>the</strong> three most abundant <strong>in</strong>vertebrate species was l<strong>in</strong>ked<br />

to <strong>the</strong> nutrient content <strong>of</strong> <strong>the</strong> surround<strong>in</strong>g waters. High epifaunal biomass was observed <strong>in</strong> <strong>the</strong><br />

centre <strong>of</strong> <strong>the</strong> nutrient-rich, cool slope waters north <strong>of</strong> <strong>the</strong> Gulf Stream while productivity was low<br />

<strong>in</strong> <strong>the</strong> oligotrophic waters <strong>of</strong> <strong>the</strong> Sargasso Sea. Unexpectedly high productivity <strong>of</strong> epifauna was<br />

detected <strong>in</strong> <strong>the</strong> Sargasso Sea <strong>in</strong> <strong>the</strong> track <strong>of</strong> a cold water r<strong>in</strong>g from <strong>the</strong> Gulf Stream. <strong>The</strong> nutrientloaded<br />

subarctic slope water allowed for high primary production <strong>in</strong>side <strong>the</strong> r<strong>in</strong>g <strong>in</strong>fluenc<strong>in</strong>g even<br />

<strong>the</strong> productivity <strong>of</strong> higher trophic levels such as <strong>the</strong> motile carnivorous epifauna on float<strong>in</strong>g<br />

Sargassum. Niermann (1986) suggested that <strong>the</strong> low degree <strong>of</strong> epibiont coverage (primarily bryozoans<br />

and hydrozoans) on float<strong>in</strong>g Sargassum <strong>in</strong> <strong>the</strong> Sargasso Sea proper could also be due to food<br />

limitation.<br />

Data presented by W<strong>in</strong>ston et al. (1997) first h<strong>in</strong>ted towards a latitud<strong>in</strong>al decrease <strong>in</strong> colonisation<br />

<strong>of</strong> mar<strong>in</strong>e debris by sessile rafters, which was later confirmed by studies from Barnes & Sanderson<br />

(2000). In general, items from low latitudes are densely colonised while items from high latitudes<br />

are typically uncolonised (W<strong>in</strong>ston et al. 1997, Barnes & Sanderson 2000, Barnes 2002). While<br />

<strong>the</strong>re exist latitud<strong>in</strong>al variations <strong>in</strong> abundance and type <strong>of</strong> float<strong>in</strong>g substrata (Barnes 2002, Thiel &<br />

Gutow 2004), differences <strong>in</strong> colonisation appear to be primarily due to ecological factors. In polar<br />

regions sessile rafters (primarily bryozoans) were almost completely absent from float<strong>in</strong>g debris<br />

(Figure 22), and <strong>the</strong> authors discussed that this could be due to <strong>the</strong> harsh physical conditions at<br />

high latitudes (Barnes & Sanderson 2000, Barnes 2002).<br />

Seasonal variations <strong>of</strong> <strong>the</strong> abundance <strong>of</strong> <strong>raft<strong>in</strong>g</strong> organisms are also more pronounced at high<br />

latitudes (Figure 23). Dur<strong>in</strong>g <strong>the</strong> w<strong>in</strong>ter, many float<strong>in</strong>g items collected <strong>in</strong> Icelandic waters (64°N)<br />

were almost free <strong>of</strong> rafters. In particular, <strong>the</strong> occurrence <strong>of</strong> some colonisers on float<strong>in</strong>g seaweed<br />

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