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BSEP116B Biodiversity in the Baltic Sea - Helcom

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52<br />

Macoma baltica<br />

where <strong>the</strong> <strong>in</strong>fluence of periodic salt-water <strong>in</strong>flows<br />

is tangible. These patterns are more dist<strong>in</strong>ct closer<br />

to <strong>the</strong> Danish Straits and <strong>the</strong> proximity to <strong>the</strong><br />

<strong>in</strong>flows. S<strong>in</strong>ce <strong>the</strong> largest recorded <strong>in</strong>flow <strong>in</strong> <strong>the</strong><br />

1950s, sal<strong>in</strong>ity has decreased <strong>in</strong> <strong>the</strong> entire <strong>Baltic</strong><br />

Proper and <strong>the</strong> dom<strong>in</strong>ance has changed from<br />

mar<strong>in</strong>e species to typical brackish-water species<br />

<strong>in</strong> <strong>the</strong> sou<strong>the</strong>astern Gotland Bas<strong>in</strong> (Figure 3.4.2).<br />

Similarly, <strong>the</strong> ‘mar<strong>in</strong>e’ elements of <strong>the</strong> community<br />

<strong>in</strong> <strong>the</strong> Gulf of F<strong>in</strong>land, such as <strong>the</strong> polychaete<br />

Terebellides stroemi, have disappeared altoge<strong>the</strong>r.<br />

The nature and magnitude of <strong>the</strong> salt-water<br />

<strong>in</strong>flows have a direct <strong>in</strong>fluence on <strong>the</strong> strength of<br />

<strong>the</strong> halocl<strong>in</strong>e and, hence, <strong>the</strong> oxygen dynamics<br />

<strong>in</strong> <strong>the</strong> <strong>Baltic</strong>. Stratification of <strong>the</strong> water column <strong>in</strong><br />

comb<strong>in</strong>ation with an <strong>in</strong>creased organic load<strong>in</strong>g<br />

(which has been exacerbated by eutrophication)<br />

has resulted <strong>in</strong> widespread hypoxia and anoxia.<br />

This poses a severe threat to <strong>the</strong> biodiversity<br />

of benthic <strong>in</strong>vertebrate communities, which is<br />

illustrated <strong>in</strong> <strong>the</strong> periodically very poor and low<br />

diversity communities recorded <strong>in</strong> <strong>the</strong> sou<strong>the</strong>astern<br />

Gotland Bas<strong>in</strong> and Gulf of F<strong>in</strong>land (Figure<br />

3.4.2). Dur<strong>in</strong>g <strong>the</strong> mid-1990s, benthic communities<br />

recovered ow<strong>in</strong>g to <strong>the</strong> stagnation period<br />

that weakened stratification; however, with <strong>the</strong><br />

large salt-water <strong>in</strong>trusion <strong>in</strong> 1993, <strong>the</strong> halocl<strong>in</strong>e<br />

streng<strong>the</strong>ned and oxygen conditions deteriorated<br />

(La<strong>in</strong>e et al. 2007, Norkko et al. 2007). In contrast,<br />

changes <strong>in</strong> species composition are small<br />

<strong>in</strong> <strong>the</strong> sou<strong>the</strong>rn Bothnian <strong>Sea</strong> (SR5) and virtually<br />

absent <strong>in</strong> <strong>the</strong> Bothnian Bay (BO3). This is due to<br />

overall low species diversity. Never<strong>the</strong>less, <strong>in</strong>vasive<br />

species, such as <strong>the</strong> polychaete Marenzelleria<br />

spp., are conspicuous elements of <strong>the</strong> system<br />

and have established viable populations <strong>in</strong> many<br />

areas of <strong>the</strong> <strong>Baltic</strong> <strong>Sea</strong>, as exemplified <strong>in</strong> sou<strong>the</strong>rn<br />

Bothnian <strong>Sea</strong>. In <strong>the</strong> Bothnian <strong>Sea</strong>, large natural<br />

variations <strong>in</strong> abundance of <strong>the</strong> dom<strong>in</strong>ant species,<br />

<strong>the</strong> amphipod Monoporeia aff<strong>in</strong>is, are typical. In<br />

this area, <strong>the</strong> <strong>in</strong>fluence of salt-water <strong>in</strong>flows is<br />

m<strong>in</strong>imal and <strong>the</strong>re is not a strong halocl<strong>in</strong>e, which<br />

would promote <strong>the</strong> formation of hypoxia. In <strong>the</strong><br />

Bothnian <strong>Sea</strong>, <strong>the</strong> Monoporeia populations have<br />

been decl<strong>in</strong><strong>in</strong>g s<strong>in</strong>ce <strong>the</strong> mid-1990s for reasons<br />

that have not been clarified. However, recent<br />

monitor<strong>in</strong>g suggests that <strong>the</strong>se populations may<br />

be recover<strong>in</strong>g.<br />

When exam<strong>in</strong><strong>in</strong>g long-term trends from data collected<br />

<strong>in</strong> 1965–2006, it becomes immediately<br />

obvious that conditions were already disturbed<br />

<strong>in</strong> <strong>the</strong> mid-1960s. Benthic <strong>in</strong>vertebrate status <strong>in</strong><br />

<strong>the</strong> central parts of <strong>the</strong> <strong>Baltic</strong> <strong>Sea</strong>, <strong>in</strong> particular,<br />

is more or less entirely controlled by <strong>the</strong> presence<br />

or absence of hypoxia/anoxia. Already <strong>in</strong><br />

Hessle’s (1924) sem<strong>in</strong>al work, hypoxia/anoxia<br />

was reported <strong>in</strong> both coastal and open-sea areas.<br />

However, he also reported on <strong>the</strong> presence of<br />

species such as <strong>the</strong> polychaete Scoloplos armiger<br />

at over 140 m depth <strong>in</strong> <strong>the</strong> eastern Gotland Bas<strong>in</strong>,<br />

which <strong>in</strong>dicates that <strong>the</strong> spatial extent of hypoxic<br />

bottom waters was limited at that time. Current<br />

evidence suggests that <strong>the</strong> spatial and temporal<br />

extent of oxygen deficiency has <strong>in</strong>creased over<br />

<strong>the</strong> past decades. In <strong>the</strong> light of historical work<br />

(Hessle 1924), it is also likely that reference conditions<br />

def<strong>in</strong>ed for open-sea areas <strong>in</strong> this assessment<br />

are underestimates. Generally, <strong>Baltic</strong> benthic<br />

macrofauna are characterized by small, shallowdwell<strong>in</strong>g<br />

species ow<strong>in</strong>g to low sal<strong>in</strong>ity and<br />

transient hypoxia; historically, it was only <strong>in</strong> <strong>the</strong><br />

sou<strong>the</strong>rn <strong>Baltic</strong>, where more mature communities<br />

composed of deeper-dwell<strong>in</strong>g larger species, e.g.,<br />

some long-lived bivalves and large polychaetes,<br />

could have developed (Tulkki 1965, Rumohr et al.<br />

1996). However, currently macrobenthic communities<br />

are severely degraded and abundances are<br />

below a 40-year average <strong>in</strong> <strong>the</strong> entire <strong>Baltic</strong> <strong>Sea</strong><br />

(Norkko et al. 2007).<br />

Functional aspects of biodiversity<br />

The broad-scale patterns and changes <strong>in</strong> structural<br />

biodiversity also translate <strong>in</strong>to differences <strong>in</strong> functional<br />

biodiversity. While <strong>the</strong> actual relationship

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