The vessel as a vector - biofouling, ballast water and sediments
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Hewitt, CL and Gollasch, S and Minchin, D, The vessel as a vector - biofouling, ballast water and sediments, Biological Invasions in Marine Ecosystems: Ecological, Management and Geographic Perspectives, Springer, Gil Rilov and Jeffrey A. Crooks (ed), Heidelberg, Germany, pp. 117-131. ISBN 978-3-540-79235-2 (2009) [Research Book Chapter]
Human-mediated marine bioinvasions have altered the way we view the marine
environment – virtually all regions of the global oceans have experienced the introduction
of marine species (e.g., Carlton 1979; Coles et al. 1999; Cranfield et al.
1998; Cohen and Carlton 1998; Hewitt et al. 1999, 2004; Orensanz et al. 2002;
Leppäkoski et al. 2002; Lewis et al. 2003; Castilla et al. 2005; Wolff 2005; Gollasch
and Nehring 2006; Minchin 2006), placing marine and coastal resources under
increased threat. Humans have almost certainly transported marine species since
early attempts to voyage by sea. These ancient transport vectors were slow, and for
the most part restricted to small spatial scales. The beginning of significant exploration
and subsequent expansion by Europeans (post 1500 AD) has resulted in the
transport of many thousands of species across all world oceans (Crosby 1986;
diCastri 1989; Carlton 2001).
The transport of species by human vectors was recognized by early workers
(Ostenfeld 1908; Elton 1958), but it is only in the last few decades that significant
progress on identifying patterns and processes has been made (e.g., Carlton 1985,
1996, 2001; Ruiz et al. 2000; Hewitt et al. 2004; Castilla et al. 2005; Minchin
2006). Numerous transport vectors have been identified and described (Carlton
2001; Chap. 5, Minchin et al.); however the majority of species appear to have been
associated with vessel movements, either as exploratory, military, commercial or
recreational vessels (e.g., Carlton 1985, 2001; Cohen and Carlton 1998; Hewitt
et al. 1999; Gollasch et al. 2002, Minchin and Gollasch 2003).
The ship as a transport vector is comprised of several sub-vectors. These include
(1) the hull and other ‘niche’ areas, such as the propeller, rudder, on exposed surfaces
of water piping, seachests, and thruster tunnels, where accumulations of
growths of organisms develop (typically known as hull fouling), (2) the boring of
organisms into the structure of the vessel (primarily limited to wooden hulled
vessels), and (3) the uptake of organisms in association with wet or dry ballast
(Carlton 1985, 1996; Ruiz et al. 2000). Several of these ship sub-vectors are no
longer active. Hull boring for example, virtually ceased to exist with the use of steel
G. Rilov, J.A. Crooks (eds.) Biological Invasions in Marine Ecosystems. 117
Ecological Studies 204,
© Springer-Verlag Berlin Heidelberg 2009
118 C.L. Hewitt et al.
as the primary ship-building material in merchant and naval vessels. However,
many pleasure boats and fishing craft are still constructed of wood (Nagabhushanam
and Sarojini 1997). Similarly, dry-ballast made up of sand, gravel and rock taken
from littoral environments was replaced with water as ballast beginning in the late
1800s and had become phased out by 1950.
None of these sub-vectors is species-specific, and each is likely to transport entire
assemblages of species. Each may also facilitate the transport of a differing suite of
species with different physiological and ecological characteristics (see Table 6.1).
Biofouling primarily transports species that have attached sedentary or sessile, benthic
habits, or species associated with these communities (e.g., living in, between or
on other organisms) (Minchin and Gollasch 2003). In contrast, ballast water transports
species associated with the plankton either as holo-plankton (species that have
their whole life-cycle in the water column), mero-plankton (species with a portion
of their life-cycle in the water column), or tycho-plankton (species accidentally
swept into the water column), and often include pelagic species. It is difficult to
establish a firm link between an already established introduced species and the vector
(or sub-vector) by which it arrived in the new location (Minchin 2007).
Nevertheless, attempts at assigning linkages to sub-vectors based on life history
modes, timing of invasions, and association between l
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