The Natura 2000 network and the ranges of threatened species in Greece

Abstract

Global environmental goals mandate the expansion of the protected area network to halt biodiversity loss. The European Union’s Natura 2000 network covers 27.3% of the terrestrial area of Greece, one of the highest percentages in Europe. However, the extent to which this network protects Europe’s biodiversity, especially in a biodiverse country like Greece, is unknown. Here, we overlap the country’s Natura 2000 network with the ranges of the 424 species assessed as threatened on the IUCN Red List and present in Greece. Natura 2000 overlaps on average 47.6% of the mapped range of threatened species; this overlap far exceeds that expected by random networks (21.4%). Special Protection Areas and Special Areas of Conservation (non-exclusive subsets of Natura 2000 sites) overlap 33.4% and 38.1% respectively. Crete and Peloponnese are the two regions with the highest percentage of threatened species, with Natura 2000 sites overlapping on average 62.3% with the threatened species’ ranges for the former, but only 30.6% for the latter. The Greek ranges of all 62 threatened species listed in Annexes 1 and II to the Birds and Habitats Directives are at least partially overlapped by the network (52.0%), and 18.0% of these are fully overlapped. However, the ranges of 27 threatened species, all of which are endemic to Greece, are not overlapped at all. These results can inform national policies for the protection of biodiversity beyond current Natura 2000 sites.

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https://doi.org/10.1007/s10531-021-02125-7
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ORIGINAL PAPER
The Natura 2000 network andtheranges ofthreatened
species inGreece
KonstantinaSpiliopoulou1,2 · PanayiotisG.Dimitrakopoulos3 ·
ThomasM.Brooks4 · GabrielaKelaidi2· KaloustParagamian5 · VassilikiKati6 ·
AnthiOikonomou1 · DimitrisVavylis2· PanayiotisTrigas7 · PetrosLymberakis8 ·
WilliamDarwall9 · MariaTh.Stoumboudi1· KostasA.Triantis2
Received: 8 June 2020 / Revised: 13 January 2021 / Accepted: 20 January 2021 /
Published online: 13 February 2021
© The Author(s) 2021
Abstract
Global environmental goals mandate the expansion of the protected area network to halt
biodiversity loss. The European Union’s Natura 2000 network covers 27.3% of the ter-
restrial area of Greece, one of the highest percentages in Europe. However, the extent to
which this network protects Europe’s biodiversity, especially in a biodiverse country like
Greece, is unknown. Here, we overlap the country’s Natura 2000 network with the ranges
of the 424 species assessed as threatened on the IUCN Red List and present in Greece.
Natura 2000 overlaps on average 47.6% of the mapped range of threatened species; this
overlap far exceeds that expected by random networks (21.4%). Special Protection Areas
and Special Areas of Conservation (non-exclusive subsets of Natura 2000 sites) overlap
33.4% and 38.1% respectively. Crete and Peloponnese are the two regions with the high-
est percentage of threatened species, with Natura 2000 sites overlapping on average 62.3%
with the threatened species’ ranges for the former, but only 30.6% for the latter. The Greek
ranges of all 62 threatened species listed in Annexes 1 and II to the Birds and Habitats
Directives are at least partially overlapped by the network (52.0%), and 18.0% of these are
fully overlapped. However, the ranges of 27 threatened species, all of which are endemic to
Greece, are not overlapped at all. These results can inform national policies for the protec-
tion of biodiversity beyond current Natura 2000 sites.
Keywords Protected areas· IUCN red list· Threatened species· Greece· Natura 2000
Communicated by Stephen Garnett.
This article belongs to the Topical Collection: Biodiversity protection and reserves.
* Konstantina Spiliopoulou
k.spilio@hcmr.gr
* Kostas A. Triantis
ktriantis@biol.uoa.gr
Extended author information available on the last page of the article
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Introduction
Biodiversity is severely threatened and declining in many parts of the world (Joppa etal.
2016b; Maxwell etal. 2016; IPBES 2019), raising concerns that an era of mass extinction
is beginning (Dirzo etal. 2014; Lewis and Maslin 2015). According to the IUCN Red List
of Threatened Species, more than 27% of the 105,700 species assessed, face “a high risk of
extinction in the wild” (IUCN 2019). Over recent years, in response to this increasing bio-
diversity loss, great effort has been allocated in implementing strategies for the protection
of nature. Protected areas have long been regarded as one of the most valuable tools for the
protection of biodiversity (Chape etal. 2008; Watson etal. 2014), and so play a major role
in these strategies. The 2011–2020 Strategic Plan for Biodiversity’s Aichi Target 11 states
that, by 2020, at least 17% of terrestrial and inland water areas and 10% of coastal and
marine areas, should be effectively managed by protected areas and “other effective area-
based conservation measures” (OECMs; CBD 2010). According to the Protected Planet
live-report (www.prote ctedp lanet .net), 15% of terrestrial and freshwater environments and
7.8% of the marine environment are protected (UNEP-WCMC & IUCN 2020). In addition
to Aichi Target 11, the Sustainable Development Goals (SDGs) 14 and 15 call for protec-
tion of the planet’s marine, terrestrial and freshwater biota (United Nations 2015). These
global targets and goals, as well as the post-2020 agenda (CBD 2020), advocate expansion
of the protected area network at the regional and national level in order to halt biodiversity
loss.
Although protected areas are considered a critical tool to conserve biodiversity, there is
still no comprehensive answer to whether they actually deliver on this commitment. While
some analyses have revealed impacts of protected areas in reducing rates of habitat loss
(Andam etal. 2008; Joppa and Pfaff 2011; Geldmann etal. 2013) and reducing increases in
extinction risk for species (Butchart etal. 2012), for most taxa the conservation outcomes
of protected areas are unknown (Joppa etal. 2016a). Generally, the greater the overlap of
a species distribution by protected areas, the higher the chances for long term persistence
ought to be (e.g. Rodrigues etal. 2004a, b); but the overall conservation outcome is highly
dependent on the specific environmental context (e.g. hydrology), as well as the particular
protected area planning, management scheme, governance and budget allocation (Rodri-
gues etal. 2004a, b; Watson etal. 2014).
The European Union has the largest coordinated network of protected areas in the world
(European Commission 2020). The Natura 2000 is a network of protected areas that was
established in 1992, operating under the European Union’s Birds and Habitat Directives.
It is comprised of two non-mutually exclusive site types, Special Protection Areas (SPAs)
and Special Areas of Conservation (SACs) (European Commission 1992, 2009). By 2019,
one year before the end of the Strategic Plan for Biodiversity 2011–2020, Natura 2000 sites
had covered 18% of the terrestrial, and almost 9.5% of the marine, European Union terri-
tory. Terrestrial coverage of the Natura 2000 varies among the European Union countries
between 8.4% and 37.8% (European Environment Agency 2019).
While the Natura 2000 network aims to “ensure the long term survival of the most valu-
able and threatened species and habitats in Europe” (European Commission 2020), bio-
diversity is not evenly distributed throughout Europe. The southern European countries,
which belong to the Mediterranean biodiversity hotspot, are characterized by higher lev-
els of threat to biodiversity and have higher levels of endemism than the rest of Europe
(Médail & Quézel 1999; Myers etal. 2000). Greece is exceptionally diverse. Despite its
relatively small size (131,940 km2; 1.3% of Europe and 3% of the European Union’s area),
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it contributes significantly to the European biodiversity with almost 32% of the known
European species being present in Greece (Aravanopoulos 2010). Due to its high topo-
graphic heterogeneity, complex paleogeographic history, fragmented landscape, and loca-
tion at the crossroad of three continents, i.e. Europe, Asia and Africa, Greece hosts a very
high number of species and has high levels of endemism (Legakis and Maragkou 2009;
Sfenthourakis etal. 2018; Legakis etal. 2018). It is estimated that Greece has about 50,000
animal species, more than 20% being endemic (e.g. Legakis etal. 2018) and more than
5800 vascular plant species, more than 22% endemic (Flora of Greece Web 2018). The
degree of endemism for some taxonomic groups, especially those that have diversified in
insular systems, exceeds 50% (e.g. Sfenthourakis etal. 2018). However, Greece also has
the second highest number of threatened species in Europe as well as in the Mediterranean
biodiversity hotspot, after Spain (BirdLife International 2017; IUCN 2019).
In Greece, the Natura 2000 network covers 27.3% of the terrestrial area. This is one of
the highest levels of protected area coverage in Europe, and is far above the 17% coverage
mandated by Aichi Target 11. However, it remains unclear how well this represents the
threatened biodiversity of Greece. Two studies have evaluated representation of species by
protected areas in Greece. One, focused on 1624 native plant species in Crete, found that
SAC sites do not represent satisfactorily the regional plant biodiversity (Dimitrakopoulos
etal. 2004). The other, on 395 vascular plant species and subspecies endemic to Pelopon-
nese found low overlap with selected networks from complementarity analysis (Trigas
etal. 2012). Here, we assess the overlap between the 424 extant, native, resident species
assessed as threatened on the IUCN Red List and present in Greece, and the country’s
Natura 2000 network. We then compared our results against null models obtained by plac-
ing equivalent “Natura 2000” sites at random over the land area of Greece.
Materials andmethods
Natura 2000
Of 1288 (overlapping) protected areas in Greece (UNEP-WCMC 2020), 446 sites are part
of the Natura 2000 network, including 239 Special Areas of Conservation, 181 Special
Protection Areas, and 26 sites that are both (Fig.1). Special Areas of Conservation are des-
ignated to ensure the favorable conservation status of each habitat type and species listed
in the Annexes of the Habitats Directive (European Commission 1992), while Special Pro-
tection Areas are designated for 194 particularly threatened species and all migratory bird
species listed in the Annexes of the Birds Directive (European Commission 2009). These
sites together cover 36,000 km2 (27.3%) of the country’s land territory; SACs cover 16.6%
and SPAs cover 20.9% with overlaps between the two site types of 10.2%. Geographical
Information System (GIS) data on Natura 2000 sites were downloaded from the European
Environmental Agency (2019).
Threatened species
We considered all species with known presence in Greece, according to the IUCN Red
List (IUCN 2019). As of March 2019 there were 3280 species assessed for Greece, of
which 2809 are terrestrial and freshwater species. Of all species assessed, animals rep-
resent 73.1% (2055 species; ~ 4% of the entire Greek fauna) and plants represent 26.8%
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(753 species; ~ 13% of the entire Greek flora). For some taxonomic groups (e.g., tetrapod
vertebrates, freshwater fishes, land snails), almost all Greek species has been assessed for
the IUCN Red List; for others (including most invertebrate and plant groups), assessments
are not comprehensive, and may be biased (e.g., towards species a priori considered likely
to be threatened, or towards particular regions). Throughout, we used the global extinc-
tion risk status for each species, not the national one (e.g., Phitos etal. 1995; Legakis and
Maragkou 2009), consistent with our objective to evaluate the contribution of the Natura
2000 network of Greece to the protection of the global biodiversity, and thus the progress
of the country towards global environmental targets.
The threatened species, i.e. those assigned to Vulnerable (VU), Endangered (EN) and Crit-
ically Endangered (CR) categories, total 476 species (Table1). We followed the taxonomy
used by the IUCN Red List, but we grouped the plant classes of Liliopsida and Magnoliop-
sida into Magnoliopsida (following Euro + Med 2006). Four Greek species, all gastropods,
Fig. 1 The Natura 2000 network in Greece. Yellow represents Special Areas of Conservation (SACs), light
blue stripes represent Special Protection Areas (SPAs) and purple represents sites that are designated as
both SAC and SPA
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assessed as Extinct, Zonites santoriniensis, Zonites siphnicus, Vitrea storchi and Graecoana-
tolica macedonica, were excluded from the analyses.
Among species groups comprehensively assessed, Actinopterygii, Gastropoda, and
Amphibia have the highest threat prevalence of threatened species: 36.6%, 26.7%, and 21.7%
respectively (Table1).
For all threatened species, we downloaded the available range maps from the IUCN Red
List website (www.iucnr edlis t.org). Range maps represent the ‘current known limits of distri-
bution of a species, accounting for all known, inferred or projected sites of occurrence’ (IUCN
2016). We only considered species that have extant, resident and native distributions which
account for code 1 in Presence (extant), Origin (native) and Seasonality (resident) following
the IUCN Red List mapping standards (IUCN 2018), yielding 424 extant, native, resident
threatened species with mapped Greek ranges. Of these, 323 are terrestrial, 97 are freshwater
species and four are amphibious species. Out of the 424 threatened species, 303 (71.5%) are
endemic to Greece. According to Annexes 1 and II of the Birds and Habitats Directives, 62 of
the 424 threatened species in this study are formally protected by the Natura 2000 network.
For Aves, since we included only species with permanent presence in the country, we
excluded from the analysis the migrating species (summer, winter and passage visitors) and
vagrants.
Table 1 Numbers of terrestrial and freshwater species in each Class assessed on the IUCN Red List, in each
threat category, and the percentage (%) of threatened species
Classes/groups with the majority (> 90%) of their species assessed are highlighted in bold.
a All totals exclude Data Deficient (DD) species.
Class/group Number of
assessed speciesa
Number of threat-
ened species
VU EN CR % of threat-
ened species
Gastropoda 647 173 113 24 36 26.7
Insecta 423 124 66 45 13 29.3
Actinopterygii 123 45 13 15 17 36.6
Magnoliopsida 633 69 29 25 15 10.9
Aves 435 25 17 4 4 5.7
Reptilia 58 11 8 3 0 19.0
Mammalia 99 9 7 2 0 9.1
Amphibia 23 5 3 1 1 21.7
Bivalvia 19 5 2 3 0 26.3
Polypodiopsida 15 3 2 1 0 20.0
Malacostraca 8 3 3 0 0 37.5
Lycopodiopsida 4 2 0 1 1 50.0
Cephalaspidomorphi 1 1 0 0 1 100.0
Agaricomycetes 1 1 1 0 0 100.0
Pinopsida 17 0 – – – 0.0
Total 2506 476 264 124 88 19.0
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Measuring theactual andexpected overlap ofthreatened species ranges
withtheNatura 2000 network
Following existing approaches (Rodrigues etal. 2004b; Araújo et al. 2007; Watson etal.
2010; Beresford etal. 2011; Cantú-Salazar etal. 2013; Trochet and Schmeller 2013; Venter
etal. 2014; Abellán and Sánchez-Fernández 2015; Klein etal. 2015; Shanee etal. 2017),
we overlapped the Natura 2000 network with the species range maps, and we calculated the
percentage of overlap using the formula:
where areanatura is the size of the species range that overlaps with the Natura 2000 network
in Greece, and arearange is the size of the species range in Greece.
We calculated the overlap of the Greek ranges of threatened species by the whole of
the Natura 2000 network and the individual site types, SACs and SPAs separately. This is
because SPA and SAC sites have been designated to protect different taxa (SPAs solely for
the protection of bird species, while SAC sites for the rest of the species). For calculating
the overlap by the individual Natura 2000 site types, we used the sites designated as SAC/
SPA in the calculations for the overlap both with SACs and SPAs. For the overlap with
the whole of the Natura 2000 network we considered all overlapping areas of the sites as a
single area value. The above analysis was repeated only for the classes/groups comprehen-
sively assessed (see Table1), in order to test for potential biases due to the fact that some
classes have a low percentage of species assessed in the IUCN Red List.
We evaluated the overlap against random, simulated protected area systems (e.g. Guil-
haumon etal. 2015; Rosso etal. 2018) in order to explore whether the overlap of the Natura
2000 network in Greece with the ranges of threatened species differs from that expected by
chance. We used a null model to generate 999 random networks of protected areas across
Greece, with the same land coverage and configuration (i.e. shape) of existing Natura 2000
sites. The algorithm used, randomly changes the centroid—therefore the location—and
rotates each Natura 2000 site on the terrestrial part of Greece. The model is constrained to
ensure that each site has a location that does not overlap with marine areas, nor with other
Natura 2000 sites and big cities, and that the total land coverage of the random networks
is the same to the current Natura 2000 network (see Guilhaumon etal. 2015; Rosso etal.
2018). Probability values were estimated as the proportion of mean overlap values from
random systems that are equal or greater than the observed overlap value (P hereafter, with
P = P (Random ≥ Observed)), by inspecting the positions of the observed value in the cor-
responding null distributions.
The analysis was conducted using the R programming language version 3.6.1 (https ://
www.r-proje ct.org/) and the “sf” R package. The code for the analysis is available at https
://zenod o.org/depos it/44363 99.
Results
The mean percentage overlap between the ranges of threatened species in Greece and
the Natura 2000 network is 47.6% (recall that Natura 2000 sites cover 27.3% of Greece’s
land area; for the respective median values see Table S1). The individual site types that
area
natura
area
range
×
100
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comprise Natura 2000, SPAs and SACs, overlap the ranges of threatened species by 33.4%
and 38.2% respectively (compared to 20.9% and 16.6% coverage of land area, respectively;
Table2). For the percentage overlap for each individual species’ Greek range by the Natura
2000 network see supplementary materials (TableS2).
Among the most comprehensively assessed classes/groups (Table 1), Actinopterygii
(51.3%), Gastropoda (48.8%) and Reptilia (46.1%) have the highest overlap with the Natura
2000 network. Our focus on resident species may explain the relatively low overlap for
Aves (29.3%), given that 13 threatened bird species occur in Greece only as winter visitors,
migrants, or vagrants.
Among the 62 threatened species listed in the Annexes of the Birds and Habitats Direc-
tives, all have at least part of their range within the Natura 2000 network in Greece. The
majority of these species (51) are partially overlapped and 11 of them (18%) are fully over-
lapped (> 99%). The mean overlap for the 62 Annexed species is 52.0%.
Twenty-seven (6.4%) out of the 424 species in this analysis had no overlap with the
Natura 2000 network (overlap < 0.1%), while 46 (10.8%) species had less than 10% over-
lap. All of these species belong to the class Gastropoda, except for two Magnoliopsida and
nine Insecta. Almost half (19) of the species with < 10% overlap are present in Peloponnese
(12 species) and Crete (7 species). There are no threatened species with < 10% overlap pre-
sent in Western Macedonia, Central Macedonia and Eastern Macedonia and Thrace, while
species with 0% overlap are present in the islands, Epirus, Central Greece, Attica and Pelo-
ponnese (Fig.2). We estimate that 4.8% increase of the Natura 2000 network will result in
covering at least 10% of the ranges of all threatened species in Greece.
The mean overlap of species ranges among the threatened categories is 51.2% ± 4.1
(mean ± Standard Error) for Critically Endangered species, 46.5% ± 2.8 for Endangered
species, and 47.1% ± 2.3 for Vulnerable species. We found no significant difference on the
overlap among the three threat categories (one-way ANOVA, p-value = 0.59). Similarly,
we found no significant difference between the terrestrial and freshwater species (t-test,
p-value = 0.67), using data on habitat and ecology for each species from the IUCN Red
List to divide into “terrestrial” (n = 327 species) and “freshwater” (n = 101 species) sys-
tems (four species are coded as both). The overlap of their ranges with the Natura 2000
network, but also the SPAs and SACs, is generally similar, although SPAs overlap the
ranges of threatened freshwater species to a greater extent than they do for terrestrial spe-
cies (Table3). Analyses using only the comprehensively assessed classes/groups yield very
similar results (Tables S3–S5; Figs. S1, S2).
The percentage of overlap with the Natura 2000 for the ranges of threatened species in
Greece is higher than offered by random networks (P = 1;Fig.3). This is also the case for
all classes individually, except from Malacostraca (the largest crustacean Class, for which
only two threatened species have been documented in Greece), for which the overlap by
the Natura 2000 is lower than expected considering random networks (P = 0.969; Fig. S3).
One-hundred and twelve species (26.4%) are expected by random networks to have < 0.1%
overlap with the Natura 2000 network (compared to 27 or 6.4% observed).
The percentage of overlap with the Natura 2000 network for each one of the threat cate-
gories (Vulnerable, Endangered and Critically Endangered) is also higher to that offered by
random networks (P = 1). For terrestrial and freshwater species, the percentage of overlap
with the Natura 2000 is again higher than that offered by random networks (P = 1).
The 62 threatened species listed in Annexes 1 and II to the Birds and Habitats Direc-
tives are expected by random networks to have 22.9% (compared to 52.0% observed) over-
lap with the Natura 2000 network. No annexed species (compared to 11 species or 18%
observed) is expected by random networks to have 100% overlap with the network.
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Table 2 Mean percentage (%) overlap of species ranges per class with the Natura 2000 network, the Special Protection Areas (SPAs) sites and the Special Areas of Conserva-
tion (SACs) sites separately
Percentages are not directly comparable between classes which have not been comprehensively assessed.
Class/Group Threatened
species
Species in the
Annexes
Mean range (km2) SPAs (%) SACs (%) Natura 2000 (%) Species with 0%
overlap
Species with
10% overlap
All species 424 62 – 33.4 38.1 47.6 27 46
Bivalvia 2 0 1558.8 61.9 58.9 65.7 0 0
Magnoliopsida 60 23 3819.2 34.9 51.8 58.4 2 2
Actinopterygii 43 26 4469.2 40.7 41.4 51.3 0 0
Gastropoda 166 0 1067.5 37.2 39.1 48.8 24 35
Reptilia 9 2 4014.1 34.0 38.6 46.1 0 0
Polypodiopsida 3 0 4284.8 12.6 40.4 45.3 0 0
Insecta 119 2 4822.9 26.9 30.3 41.2 1 9
Amphibia 5 1 2750.3 13.9 39.1 41.1 0 0
Mammalia 9 7 25,152.8 28.4 27.3 39.8 0 0
Aves 3 1 5739.7 18.3 22.4 29.3 0 0
Malacostraca 2 0 61,626.7 23.3 17.0 28.8 0 0
Agaricomycetes 1 0 93,338.5 19.5 14.4 24.2 0 0
Cephalaspidomorphi 1 0 9702.8 21.7 11.1 23.9 0 0
Lycopodiopsida 1 0 239.6 6.0 20.4 22.3 0 0
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The percentage of threatened species in Greece across the 13 administrative regions
increases southwards. Crete and Peloponnese are the two regions with the highest per-
centage of threatened species (12.0% and 9.7% respectively). To the north, although
species richness increases, the percentage of threatened species per region decreases
(Fig.4; TableS6). Eastern Macedonia and Thrace (5.5%) and North Aegean (4.9%) are
the two regions with the lowest percentage of threatened species. The mean overlap per
Fig. 2 The location of threatened species ranges (Red) that have no overlap with the Natura 2000 network
and the Natura 2000 sites (Green) in Greece
Table 3 Mean percentage (%) overlap of threatened species ranges with the Natura 2000 network, SPAs
and SACs for terrestrial and freshwater species
System Natura 2000 (%) SPAs (%) SACs (%)
Terrestrial 47.7 31.6 38.3
Freshwater 46.0 38.3 36.4
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Class with the Natura 2000 in each administrative region can be found at the supple-
mentary materials (TableS7).
Regions contribute at different levels to the total overlap for threatened species ranges
in Greece (Fig.4). Crete provides the highest mean overlap of threatened species’ ranges
(62.3%), followed by South Aegean (54.3%). The regions with the lowest percentages are
Western Greece (29.4%) and Attica (18.6%).
Discussion
Protected areas are a critical tool for the protection of nature. However, the effectiveness
of protected areas still remains a highly debated topic (Watson et al. 2014; Joppa etal.
2016a; Acreman etal. 2019). Several indices have been used to evaluate protected areas,
e.g. governance, budget allocation and management plans. Although these parameters pro-
vide an indirect evaluation of protected area performance, some correlation with favorable
0.0
0.1
0.2
0.3
0.4
0.5
10 12 14 16 18 20 22 24 26 28 30 32 34 36 38 40 42 44 46 48 50
Overlap (%)
Density
Null model
distribution
Mean overlap
null model
Mean observed
overlap
Observed Value: 47.6
P (Random = Observed): 1
Fig. 3 The distribution of the percentages of overlap between the Greek ranges of threatened species (424)
and the 999 random networks obtained by the null model (light grey), the mean percentage overlap obtained
by the null model (grey dashed line) and the mean percentage overlap observed between the Greek ranges
of threatened species (424) and the current Natura 2000 network in Greece (black dashed line)
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conservation outcomes has been shown (Bruner etal. 2001; Leverington etal. 2010; Geld-
mann etal. 2018; Coad etal. 2019). Other approaches use representation targets for indi-
vidual species, based on the area they occupy and on the percentage of the global distribu-
tion range occurring in the focal regions (Rodrigues etal. 2004a; Maiorano etal. 2015), or
calculate an index of representation for Annex II species of the Habitats Directive in the
Natura 2000 network (Gruber etal. 2012).
The percentage of overlap of species ranges is another metric used to evaluate protected
areas. In comparison to other metrics, this uses distribution data for multiple species. The
percentage of overlap of species ranges with protected areas has been used in studies at the
global scale (e.g. Rodrigues etal. 2004a, b; Cantú-Salazar etal. 2013; Venter etal. 2014),
regional scale (e.g. Watson etal. 2010; Trochet and Schmeller 2013; Abellán and Sánchez-
Fernández 2015) and national scale (Araújo et al. 2007; Shanee et al. 2017). However,
these studies mainly focused on the better known chordate taxa (birds, mammals, reptiles
and amphibians).
Our study evaluates the whole of the Natura 2000 network in Greece—which pro-
vides a better coverage than provided by random networks (Fig.3)—using all threatened
species data available, for 424 species encompassing both vertebrate and invertebrate,
Fig. 4 Percentage (%) of threatened species per administrative region in Greece. Darker hue of purple rep-
resents higher percentage of threatened species. Numbers under the names of the regions represent the total
number of species (first number in brackets; derived from IUCN Red List range maps), the total number
of threatened species (second number in brackets; again derived from IUCN Red List range maps) and the
mean percentage (%) overlap of threatened species’ ranges with the Natura 2000 per region (TableS7)
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as well as plant, taxa; thus we have not focused solely on the species listed in the
Annexes of the Habitats and Birds Directives, but all threatened species and not to spe-
cific regions but the whole country. No difference on the overlap between terrestrial and
freshwater species was observed. A similar study at the European scale, using data from
300 species overall, with the most being fish, showed 35% overlap between the ranges
of 76 threatened species and the Natura 2000 network in Greece (Trochet and Schmel-
ler 2013). Our results show a higher degree of overlap of species ranges with the Natura
2000 network probably due to many more IUCN Red List assessments being conducted
since 2013, and due to the Natura 2000 network in Greece being expanded in 2017.
Our finding that the SPA and SAC sites separately overlap the ranges of threatened
species in Greece at 33.4% and 38.2% respectively, is consistent with expectations based
on the fact that these two categories have been established based on different criteria and
with different aims. Specifically, SACs should have a higher overlap of threatened spe-
cies ranges compared to SPAs, which are designed for the protection of birds only. The
fact that there is no significant difference in the overlap among the three threat catego-
ries (CR, EN and VU) with the Natura 2000 network (one-way ANOVA, p-value = 0.59)
is also interesting. Re-analyzing the percentages of overlap for each species using the
data provided by Trochet and Schmeller (2013; Appendix1), the pattern remains the
same (one-way ANOVA, p-value = 0.33). The overlap of all threatened species ranges
with the Natura 2000 network—regardless of the level of threat they face—is important
to the conservation of threatened biodiversity. That said, the expansion of the network,
in order to increase the overlap with the ranges of species facing imminent extinction,
could be considered as a policy response. Additionally, as we saw an increase of 4.8%
would result to the coverage of at least 10% of all the threatened species.
Greece has higher species richness in the northern part of the country and higher
endemism in the south. Endemic species are more sensitive to changes (e.g. Gaston
1994) which could explain the higher number of threatened species in southern admin-
istrative regions (e.g. Crete and Peloponnese). Although the overlap between threatened
species and the Natura 2000 network in Crete is quite high (62.3%), that in Peloponnese
is only 30.6% (the third lowest in Greece), highlighting the need to expand and poten-
tially re-structure the conservation network in this part of the country. Western Greece
and Attica regions provide the lowest overlap between threatened species and Natura
2000. Increasing the conservation efforts in these regions, and also in regions that have
high percentages of threatened species but relatively low overlap with the Natura 2000
network (e.g. Peloponnese) will contribute substantially to the protection of threatened
biodiversity in Greece. Such efforts could be achieved through multiple pathways: for
example, through expansion of the Natura 2000 network, as was achieved in 2017 (Joint
Ministerial Decision 50743/2017), or by implementation of complementary conserva-
tion approaches such as community protected areas (Dudley 2008), or “other effective
area-based conservation measures” (IUCN WCPA 2019), or by restoration (Carrizo
etal. 2017).
For Greece, the protection of threatened species—the majority of which are endemics
(71.5%)—offers significant contribution to the global biodiversity. The results presented
herein can inform national policies for the protection of biodiversity. Five aspects can be
identified: (a) determine the threatened species that have zero or low overlap with the Nat-
ura 2000 network in Greece; (b) increase the levels of protection for the species with the
most imminent threat of extinction; (c) restore critical ecosystems; (d) pinpoint the regions
that are of critical importance for the biodiversity of Greece, as they contain the highest
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Biodiversity and Conservation (2021) 30:945–961
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percentages of threatened species; (e) highlight the regions with low mean Natura 2000
overlap with their threatened species ranges.
In order to accomplish more efficient protected area networks, several prioritiza-
tion and planning tools are available. One such tool is Key Biodiversity Areas (KBAs),
which implements the long-standing concept of important areas for the persistence
of biodiversity, across all taxonomic groups. The Standard for the Identification of
KBAs was recently released (IUCN 2016) and it could be used to identify sites that
will strengthen the protected area network in Greece, possibly also through harnessing
“other effective area-based conservation measures” as a complement to the Natura 2000
network. KBAs themselves can also provide benefits to threatened species—even if they
are not within protected areas—as they can stimulate environmental safeguards.
Conclusion
Greece has met the percentage coverage of area specified by Aichi target 11 and has
one of the most extensive Natura 2000 networks in the European Union. Moreover, the
current network is demonstrably superior to a random placement of the sites. However,
it fails to adequately represent all threatened species that are of priority for protection
globally, with 27 endemic species wholly unrepresented. Expansion of the network to
encompass populations of these species would put Greece at the forefront of countries
fulfilling their EU’s Biodiversity Strategy for 2030, and their responsibility to safe-
guard global biodiversity, which would be a remarkable result given its concentration
of endemic and threatened biodiversity. Results herein should be complemented with
other available approaches such as habitat restoration and approaches that evaluate pro-
tected areas such as governance, institutional framework and stability, budget allocation,
management plans and gap analysis for species listed in the annexes of the Habitats and
Birds Directives, in the case of Europe.
Supplementary Information The online version contains supplementary material available at https ://doi.
org/10.1007/s1053 1-021-02125 -7.
Acknowledgements We would like to thank Francois Guilhaumonand Stephen Garnett for theirvalua-
ble contribution to this paper. This project has received funding from the European Union’s Horizon 2020
research and innovation programme under the Marie Skłodowska-Curie Grant Agreement No. 766417. This
communication reflects only the authors’ view and the Research Executive Agency of the European Union is
not responsible for any use that may be made of the information it contains
Funding This project has received funding from the European Union’s Horizon 2020 research and innova-
tion programme under the Marie Skłodowska-Curie Grant Agreement No. 766417.
Data availability The datasets analysed during the current study are available in the Euro-
pean Environmental Agency, https ://www.eea.europ a.eu/data-and-maps/data/natur a-11 and
the IUCN Red List of Threatened Species, https ://www.iucnr edlis t.org/searc h.
Code availability The code developed for the purpose of this study is available at https ://
zenod o.org/depos it/44363 99.
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Compliance with ethical standards
Conflict of interests The authors have no conflicts of interest to declare.
Open Access This article is licensed under a Creative Commons Attribution 4.0 International License,
which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long
as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Com-
mons licence, and indicate if changes were made. The images or other third party material in this article
are included in the article’s Creative Commons licence, unless indicated otherwise in a credit line to the
material. If material is not included in the article’s Creative Commons licence and your intended use is not
permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly
from the copyright holder. To view a copy of this licence, visit http://creat iveco mmons .org/licen ses/by/4.0/.
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Authors and Aliations
KonstantinaSpiliopoulou1,2 · PanayiotisG.Dimitrakopoulos3 ·
ThomasM.Brooks4 · GabrielaKelaidi2· KaloustParagamian5 · VassilikiKati6 ·
AnthiOikonomou1 · DimitrisVavylis2· PanayiotisTrigas7 · PetrosLymberakis8 ·
WilliamDarwall9 · MariaTh.Stoumboudi1· KostasA.Triantis2
1 Institute ofMarine Biological Resources andInland Waters, Hellenic Centre forMarine Research,
19013Anavissos, Greece
2 Department ofEcology andTaxonomy, Faculty ofBiology, National andKapodistrian University
ofAthens, 15784Athens, Greece
3 Biodiversity Conservation Laboratory, Department ofEnvironment, University oftheAegean,
81100Mytilene, Greece
4 Science andKnowledge Unit, International Union forConservation ofNature (IUCN),
1196Gland, Switzerland
5 Hellenic Institute ofSpeleological Research, 71409Irakleio, Crete, Greece
6 Department ofBiological Applications andTechnology, University ofIoannina, 45110Ioannina,
Greece
7 Faculty ofCrop Science, Agricultural University ofAthens, 11855Athens, Greece
8 Natural History Museum ofCrete, University ofCrete, 71409Irakleio, Greece
9 Freshwater Biodiversity Unit, IUCN Global Species Programme, The David Attenborough
Building, CambridgeCB23QZ, UK
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