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% change<br />

200<br />

150<br />

100<br />

50<br />

0<br />

populations. Although the overall size of the<br />

European population of Harbour seal is twice<br />

that of 1960 by the end of the time series <strong>in</strong> 2005,<br />

abundance at this po<strong>in</strong>t is decreas<strong>in</strong>g. The trend<br />

for Harbour seal is based on 20 populations from<br />

Denmark, Germany, the Netherlands, Sweden and<br />

the United K<strong>in</strong>gdom, which represent an average<br />

of 47,000 <strong>in</strong>dividuals or 58% of the total European<br />

population of 2008, and cover 33% of the countries<br />

of occurrence.<br />

50<br />

1960s 1970s 1980s 1990s 2000–05 1960–2005<br />

Drivers of recovery<br />

Figure 2.<br />

Change <strong>in</strong> Harbour<br />

seal population<br />

abundance by<br />

decade and overall<br />

change between<br />

1960 and 2005. Please<br />

note that due to the<br />

way change was<br />

calculated, decadal<br />

change does not sum<br />

to overall change.<br />

Table 3.<br />

Major reasons for<br />

positive change <strong>in</strong> the<br />

status of the Harbour<br />

seal <strong>in</strong> Europe.<br />

Rank Reason for change Description<br />

1 Species management –<br />

Reduced hunt<strong>in</strong>g pressure<br />

(1599) and 1955 <strong>in</strong> the English Channel, North Sea<br />

and northern Russian coast is likely an artefact of<br />

<strong>in</strong>complete distributional <strong>in</strong>formation <strong>in</strong> historic<br />

times. It seems plausible that seals were, <strong>in</strong> fact,<br />

only documented <strong>in</strong> areas that were of <strong>in</strong>terest to<br />

humans [19] . In addition, the 1955 map may suffer<br />

from lack of detail, giv<strong>in</strong>g rise to over- or underestimations<br />

of change to the present day. Accord<strong>in</strong>g<br />

to the <strong>in</strong>formation <strong>in</strong> Figures 1a and b, the species<br />

lost territory <strong>in</strong> the Baltic Sea while it expanded<br />

around the United K<strong>in</strong>gdom and the Russian coast.<br />

This represents a recovery from range decreases<br />

prior to the middle of the 20 th century, leav<strong>in</strong>g the<br />

species to occupy an area that is 6% larger than <strong>in</strong><br />

historical times.<br />

The <strong>in</strong>creas<strong>in</strong>g trend <strong>in</strong> distribution is <strong>in</strong> l<strong>in</strong>e<br />

with the change <strong>in</strong> population size, as European<br />

Harbour seals doubled <strong>in</strong> abundance over the<br />

45-year study period between 1960 and 2005<br />

(Figure 2). Decl<strong>in</strong>es occurred <strong>in</strong> the 1960s and<br />

2000s, the latter co<strong>in</strong>cid<strong>in</strong>g with the second<br />

outbreak of PDV [12] . However, the trend <strong>in</strong> Figure 2<br />

also suggests <strong>in</strong>creases <strong>in</strong> the 1970s and 1980s,<br />

which are unexpected due to the PDV outbreak<br />

<strong>in</strong> 1988. It is therefore possible that the expected<br />

decl<strong>in</strong>es are masked by <strong>in</strong>creases <strong>in</strong> unaffected<br />

Follow<strong>in</strong>g a reduction <strong>in</strong> hunt<strong>in</strong>g pressure, the<br />

Harbour seal recovered from 3,000 <strong>in</strong>dividuals <strong>in</strong><br />

1974 to more than 15,000 <strong>in</strong> 2003 <strong>in</strong> the Wadden<br />

Sea [13] .<br />

2 Legislation Increased European-wide and country-level legal<br />

protection of the species and its haul-out sites<br />

from the late 1970s helped recovery throughout its<br />

range [15, 16] .<br />

The Kalmarsund population recovered from 50 <strong>in</strong><br />

the 1970s [34] to 630 <strong>in</strong> 2007 [40] due to protection.<br />

Hunt<strong>in</strong>g is now prohibited <strong>in</strong> all countries except<br />

Norway and Iceland [2] .<br />

3 Land/water protection &<br />

management<br />

4 Other – Collaborative<br />

management<br />

Habitat protection also contributed to the recovery<br />

of the Harbour seal <strong>in</strong> the Wadden Sea [13] .<br />

Some countries are collaborat<strong>in</strong>g on management<br />

plans, e.g. the Netherlands, Germany and Denmark<br />

<strong>in</strong> the Wadden Sea [30–33] .<br />

As discussed above, disease has had a profound<br />

impact on the abundance trend of the Harbour<br />

seal <strong>in</strong> Europe. Along with exploitation, disease<br />

emerged as one of the predict<strong>in</strong>g factors of<br />

population decl<strong>in</strong>e from our analysis, while habitat<br />

degradation and unknown threats were generally<br />

associated with positive trends. In addition, the<br />

presence of management <strong>in</strong>tervention, specifically<br />

the cessation of hunt<strong>in</strong>g, was beneficial, although<br />

change was also positive for those populations<br />

which received no attention. As expected, non-utilised<br />

populations <strong>in</strong>creased, while those with<br />

unknown utilisation status decl<strong>in</strong>ed.<br />

The importance of the factors identified <strong>in</strong> our<br />

data set is also confirmed by the literature (Table 3).<br />

Increased legal protection was implemented from<br />

the late 1970s both <strong>in</strong> <strong>in</strong>dividual countries at a<br />

European scale [15, 16] , and this undoubtedly helped<br />

the Harbour seal recover <strong>in</strong> number and range. For<br />

example, protection was key <strong>in</strong> the recovery of the<br />

Kalmarsund population <strong>in</strong> the Baltic from a mere<br />

50 <strong>in</strong>dividuals <strong>in</strong> the 1970s [34] to 630 <strong>in</strong> 2007 [40] . In<br />

the Wadden Sea, the species <strong>in</strong>creased from 3,000<br />

<strong>in</strong>dividuals <strong>in</strong> 1974 to more than 15,000 <strong>in</strong> 2003 due<br />

to a reduction <strong>in</strong> hunt<strong>in</strong>g pressure and <strong>in</strong>creased<br />

habitat protection [13] , made possible by collaborative<br />

management between the Netherlands,<br />

Germany and Denmark [30–33] . At present, hunt<strong>in</strong>g<br />

is prohibited <strong>in</strong> all countries except Norway and<br />

Iceland [2] . Other, non-governmental conservation<br />

measures <strong>in</strong>clude the rescue and rehabilitation of<br />

seal pups [43] , and while these efforts were unable to<br />

save <strong>in</strong>fected seals dur<strong>in</strong>g the two epidemics, they<br />

may have made some contribution to population<br />

recovery. However, there is some debate as to<br />

whether this <strong>in</strong>tervention is beneficial for the<br />

species <strong>in</strong> the long-term, as it side-steps natural<br />

selection by keep<strong>in</strong>g alive <strong>in</strong>dividuals that may<br />

otherwise perish, and also <strong>in</strong>creases the risk of<br />

<strong>in</strong>fection if healthy seals are brought to a centre<br />

where the disease is present.<br />

136

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