Introduction
Methods
Study area
Variable | Description |
---|---|
Period under urban pressure (PU) | VI: foundation territory, V: ~600 years, IV: ~100 years (before 1916), III: 1916–1931 (~90–80 years), II: 1954–1959 (~60–50 years), I: 1975–1989 (~40–20 years) |
Land-use and land-cover levels | F: forests and seminatural areas: broad-leaved (blf), coniferous (cf); A: agricultural areas: complex cultivation patterns (ccp), nonirrigated arable land (al), pastures (p), land principally occupied by agriculture with significant areas of natural vegetation (nv); U: urban areas: continuous urban fabric (cuf), discontinuous urban fabric (duf), green urban areas (gua), construction sites (cs), sport and leisure facilities (sandl) |
Communication network | Ro: roads (km × 1 km−2); Ra: railway areas (km × 1 km−2) |
Habitat heterogeneity (HH) | Amount of land-use and land-cover habitats |
Hemeroby level (HL) | 1: oligohemerobic—little anthropogenic influence, actual vegetation corresponds with natural vegetation, e.g., best preserved patches of forest, xerothermic grasslands; 2: mesohemerobic—anthropogenic factors in a low to moderate degree, e.g., meadows, pastures, bushes; 3: euhemerobic—anthropogenic factors act constantly to a high degree; ruderal, segetal, and transformed seminatural communities, e.g., field and garden habitats, fallow grounds, waste lands; 4: polyhemerobic—influence of anthropogenic factors is constant and very strong, e.g., roadsides, railway areas |
Flora indicator | Description |
---|---|
Species richness (SR) | Numbers of species in basic squares or entire transect |
Endangered species (ES) | Species from Regional and National Red Lists and legally protected |
Geographical-historical
elements
| |
Sponthaneophytes (N
SP) | Indigenous species, occupying natural and seminatural habitats |
Apophytes (N
AP) | Indigenous species, permanently occupying anthropogenic habitats |
Native species (N
NS) | Amount of apophytes and sponthaneophytes |
Archeophytes (N
AR) | Species of foreign origin, introduced before the end of fifteenth century, permanently naturalized |
Kenophytes (N
KN) | Species of foreign origin introduced after fifteenth century, permanently naturalized |
Diaphytes (N
DIA) | Species of foreign origin, occurring sporadically, introduced by human activity or growing wild from arable lands |
Alien species (N
AS) | Amount of archeo-, keno-, and diaphytes |
Synecological groups
| |
Forest species (N
F) | Species from syntaxa: Rhamno-Prunetea, Salicetea purpureae, Alnetea glutinosae, Erico-Pinetea, Vaccinio-Piceetea, Quercetea robori-petraeae, Querco-Fagetea
|
Cut-over and moorland species (N
CM) | Species from syntaxa: Nardo-Callunetea, Epilobion angustifolii
|
Meadow species (N
M) | Species from syntaxa: Molinio-Arrhenatheretea
|
Xerothermic species (N
X) | Species from syntaxa: Festuco-Brometea, Trifolio-Geranietea sanguinei
|
Water and waterside species (N
W) | Species from syntaxa: Lemnetea, Potametea, Utricularietea intermedio-minoris, Montio-Cardaminetea, Scheuchzerio-Caricetea fuscae, Bidentetea tripartiti, Isoëto-Nanojuncetea, Phragmitetea
|
Segetal species (N
S) | Species from syntaxa: Polygono-Chenopodietalia, Stellarietea mediae
|
Ruderal species (N
R) | Species from syntaxa: Eragrostietalia, Sisymbrietalia, Artemisietea vulgaris, Agropyretea intermedio-repentis
|
Undefined species (N
UN) | Species with undefined phytosociological affiliation |
Synanthropization indicators
| |
Anthropophytization indicator (I
AN) |
I
AN = (N
AR + N
KN) × 100/SR |
Archaeophytization indicator (I
AR) |
I
AR = N
AR × 100/SR |
Kenophytization indicator (I
KN) |
I
KN = N
KNz × 100/SR |
Modernization indicator (I
M) |
I
M = N
KN × 100/(N
KN + N
AR + N
DIA) |
Data collection
Square | Land-use and land-cover | ||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|
Agricultural areas (%) | Urban areas (%) | ||||||||||
ccp | al | p | nv | Sum | cuf | duf | gua | cs | sandl | Sum | |
A1 | 2 | 89 | 0 | 9 | 100 | 0 | 0 | 0 | 0 | 0 | 0 |
A2 | 4 | 62 | 0 | 34 | 100 | 0 | 0 | 0 | 0 | 0 | 0 |
A3 | 18 | 62 | 8 | 2 | 100 | 0 | 0 | 0 | 0 | 0 | 0 |
A4 | 30 | 4 | 36 | 0 | 72 | 0 | 14 | 0 | 0 | 15 | 28 |
A5 | 67 | 25 | 0 | 0 | 92 | 0 | 8 | 0 | 0 | 0 | 8 |
A20 | 2 | 54 | 0 | 0 | 57 | 0 | 0 | 0 | 0 | 0 | 0 |
F16 | 5 | 3 | 0 | 0 | 8 | 0 | 0 | 0 | 7 | 0 | 7 |
F17 | 0 | 14 | 0 | 0 | 14 | 0 | 0 | 0 | 0 | 0 | 0 |
F18 | 4 | 0 | 0 | 0 | 4 | 0 | 0 | 0 | 0 | 0 | 0 |
F19 | 20 | 0 | 0 | 0 | 20 | 0 | 0 | 0 | 0 | 0 | 0 |
U6 | 10 | 10 | 0 | 0 | 20 | 2 | 60 | 18 | 0 | 0 | 80 |
U7 | 0 | 0 | 0 | 0 | 0 | 0 | 32 | 29 | 0 | 39 | 100 |
U8 | 0 | 0 | 0 | 0 | 0 | 1 | 47 | 0 | 0 | 52 | 100 |
U9 | 0 | 0 | 0 | 0 | 0 | 36 | 35 | 28 | 1 | 100 | 100 |
U10 | 0 | 0 | 0 | 0 | 0 | 0 | 73 | 26 | 0 | 0 | 100 |
U11 | 0 | 0 | 0 | 0 | 0 | 34 | 20 | 46 | 0 | 0 | 100 |
U12 | 0 | 0 | 0 | 0 | 0 | 0 | 53 | 46 | 1 | 0 | 100 |
U13 | 0 | 0 | 0 | 0 | 0 | 43 | 13 | 44 | 0 | 0 | 100 |
U14 | 0 | 0 | 0 | 0 | 0 | 0 | 70 | 21 | 1 | 0 | 93 |
U15 | 0 | 0 | 0 | 0 | 0 | 0 | 17 | 0 | 15 | 62 | 79 |
Square | Land-use and land-cover | |||||||
---|---|---|---|---|---|---|---|---|
Forests (%) | Communication network in km, 1 km−2
| Urban indicators | ||||||
blf | cf | Sum | Ro | Ra | PU | HH | HL | |
A1 | 0 | 0 | 0 | 4.17 | 0 | 1 | 1 | 3.12 |
A2 | 0 | 0 | 0 | 4.76 | 0 | 1 | 1 | 2.85 |
A3 | 0 | 0 | 0 | 5.83 | 0 | 1 | 1 | 2.86 |
A4 | 0 | 0 | 0 | 12.42 | 2.11 | 1.5 | 2 | 2.93 |
A5 | 0 | 0 | 0 | 8.16 | 0 | 2 | 2 | 2.78 |
A20 | 43 | 0 | 43 | 4.73 | 0 | 1.5 | 2 | 3.1 |
F16 | 85 | 0 | 85 | 4.31 | 5.36 | 2 | 3 | 2.7 |
F17 | 53 | 33 | 86 | 0 | 2.06 | 2 | 2 | 2.8 |
F18 | 79 | 17 | 96 | 0 | 0 | 2 | 2 | 2.84 |
F19 | 80 | 0 | 80 | 0 | 0 | 2 | 2 | 2.78 |
U6 | 0 | 0 | 0 | 5.36 | 1.75 | 3 | 2 | 3.08 |
U7 | 0 | 0 | 0 | 28.86 | 0 | 3 | 1 | 2.98 |
U8 | 0 | 0 | 0 | 16.43 | 0 | 3.5 | 1 | 3.09 |
U9 | 0 | 0 | 0 | 18.62 | 0 | 4.5 | 1 | 2.96 |
U10 | 0 | 0 | 0 | 18.33 | 0.26 | 3.5 | 1 | 3.16 |
U11 | 0 | 0 | 0 | 18.35 | 0 | 3.5 | 1 | 3.01 |
U12 | 0 | 0 | 0 | 5.47 | 2.0 | 3 | 1 | 3.09 |
U13 | 0 | 0 | 0 | 17.98 | 0 | 2.5 | 1 | 2.97 |
U14 | 7 | 0 | 7 | 12.48 | 2.43 | 2.5 | 2 | 3.10 |
U15 | 6 | 0 | 21 | 17.07 | 0.24 | 2 | 2 | 2.85 |
Statistical analyses
Results
Value of habitat indicators
Variable | Explains (%) | Contribution (%) |
F value |
P value |
---|---|---|---|---|
HH | 8.2 | 21.9 | 1.6 | 0.002 |
A | 6.9 | 18.5 | 1.4 | 0.004 |
Ro | 5.9 | 15.9 | 1.2 | 0.054 |
PU | 5.6 | 15.0 | 1.1 | 0.208 |
Ra | 5.4 | 12.1 | 1.1 | 0.254 |
HL | 5.4 | 14.3 | 1.1 | 0.286 |
Variable | PU | HH | HL | A | U | F | Ro | Ra |
---|---|---|---|---|---|---|---|---|
SR | 0.26 | 0.11 | 0.40 | −0.28 | 0.47* | −0.39 | 0.40 | −0.22 |
N
Ap
| 0.18 | 0.16 | 0.32 | −0.20 | 0.43 | −0.39 | 0.37 | −0.16 |
N
Sp
| −0.30 | 0.55* | −0.28 | 0.27 | −0.27 | 0.30 | −0.04 | −0.03 |
N
NS
| 0.08 | 0.31 | 0.19 | −0.11 | 0.27 | −0.18 | 0.30 | −0.09 |
N
Ar
| 0.70** | −0.29 | 0.66** | −0.68* | 0.79** | −0.45* | 0.54* | −0.32 |
N
KN
| 0.28 | −0.10 | 0.12 | −0.34 | 0.32 | −0.04 | 0.24 | −0.56** |
N
Dia
| 0.17 | −0.04 | 0.46* | −0.16 | 0.30 | −0.39 | 0.22 | 0.33 |
N
AS
| 0.67** | −0.29 | 0.73** | −0.67** | 0.80** | −0.54* | 0.57** | −0.27 |
N
F
| 0.10 | 0.04 | −0.06 | −0.12 | 0.04 | 0.15 | 0.04 | −0.35 |
N
CM
| −0.11 | 0.17 | −0.28 | 0.04 | −0.04 | 0.09 | −0.11 | −0.10 |
N
M
| 0.35 | 0.33 | 0.41 | −0.46* | 0.55* | −0.19 | 0.32 | 0.07 |
N
X
| 0.31 | 0.41 | 0.42 | −0.37 | 0.46* | −0.23 | 0.30 | 0.05 |
N
W
| 0.28 | 0.09 | 0.37 | −0.42 | 0.52* | −0.26 | 0.30 | 0.06 |
N
R
| 0.70** | −0.04 | 0.63** | −0.80** | 0.75** | −0.12 | 0.59* | 0.03 |
N
S
| 0.72*** | 0.01 | 0.73*** | −0.78*** | 0.81*** | −0.33 | 0.59** | 0.08 |
N
UN
| 0.52* | −0.18 | 0.38 | −0.64** | 0.55* | −0.06 | 0.46* | −0.26 |
I
AN
| 0.72** | −0.47* | 0.78*** | −0.67** | 0.77** | −0.58** | 0.48* | −0.18 |
I
AR
| 0.33 | −0.28 | 0.48* | −0.26 | 0.38 | 0.46* | −0.35 | −0.53* |
I
KN
| 0.83*** | −0.50* | 0.80*** | −0.12 | 0.54* | 0.82*** | −0.83*** | −0.39 |
I
M
| 0.19 | −0.24 | 0.00 | −0.51* | 0.06 | 0.14 | −0.23 | 0.11 |
Value of flora indicators
Flora indicator | Type of square | |||||||||
---|---|---|---|---|---|---|---|---|---|---|
Agricultural areas | Urban areas | |||||||||
A1 | A2 | A3 | A4 | A5 | A20 | U6 | U7 | U8 | U9 | |
SR | 152 | 268 | 158 | 276 | 284 | 137 | 226 | 301 | 200 | 313 |
ES | 4 | 10 | 7 | 10 | 7 | 1 | 6 | 2 | 8 | 4 |
N
SP
| 9 | 29 | 17 | 21 | 36 | 4 | 15 | 20 | 17 | 23 |
N
AP
| 95 | 162 | 103 | 171 | 179 | 87 | 126 | 175 | 97 | 173 |
N
NS
| 104 | 191 | 120 | 192 | 215 | 91 | 141 | 195 | 114 | 196 |
N
AR
| 14 | 30 | 20 | 35 | 40 | 29 | 36 | 54 | 40 | 52 |
N
KN
| 1 | 4 | 17 | 4 | 28 | 17 | 7 | 45 | 44 | 13 |
N
DIA
| 33 | 43 | 1 | 45 | 1 | 0 | 42 | 7 | 2 | 52 |
N
AS
| 48 | 77 | 38 | 84 | 69 | 46 | 85 | 106 | 86 | 117 |
N
F
| 24 | 82 | 15 | 15 | 33 | 22 | 29 | 30 | 32 | 48 |
N
CM
| 6 | 7 | 4 | 3 | 11 | 4 | 6 | 10 | 3 | 4 |
N
M
| 31 | 35 | 39 | 32 | 62 | 27 | 55 | 58 | 32 | 50 |
N
X
| 16 | 23 | 35 | 21 | 54 | 13 | 59 | 43 | 23 | 51 |
N
W
| 5 | 6 | 5 | 8 | 21 | 1 | 10 | 19 | 8 | 1 |
N
S
| 21 | 22 | 21 | 30 | 32 | 24 | 39 | 41 | 25 | 42 |
N
R
| 41 | 38 | 34 | 38 | 57 | 41 | 55 | 79 | 63 | 77 |
N
UN
| 7 | 7 | 5 | 5 | 14 | 5 | 12 | 21 | 14 | 26 |
I
AN
| 31.6 | 28.7 | 24.1 | 30.4 | 24.3 | 33.6 | 37.6 | 35.2 | 43.0 | 37.4 |
I
AR
| 21.7 | 16 | 13 | 16.3 | 14 | 21 | 18.6 | 18 | 20 | 16.6 |
I
KN
| 9.2 | 11.2 | 11 | 12.7 | 10 | 12 | 15.9 | 15 | 22 | 16.6 |
I
M
| 2.1 | 5.2 | 44.7 | 4.8 | 40.6 | 37.0 | 8.2 | 42.5 | 51.2 | 11.1 |
Flora indicator | Type of square | |||||||||
---|---|---|---|---|---|---|---|---|---|---|
Urban areas | Forests and seminatural areas | |||||||||
U10 | U11 | U12 | U13 | U14 | U15 | F16 | F17 | F18 | F19 | |
SR | 219 | 240 | 257 | 369 | 150 | 321 | 219 | 220 | 222 | 171 |
ES | 5 | 2 | 7 | 2 | 11 | 10 | 4 | 10 | 7 | 6 |
N
SP
| 5 | 16 | 9 | 29 | 12 | 33 | 56 | 27 | 46 | 33 |
N
AP
| 127 | 136 | 146 | 211 | 92 | 187 | 117 | 139 | 114 | 97 |
N
NS
| 132 | 152 | 155 | 240 | 104 | 220 | 173 | 166 | 160 | 130 |
N
AR
| 43 | 49 | 51 | 60 | 20 | 48 | 33 | 28 | 20 | 23 |
N
KN
| 5 | 38 | 7 | 60 | 1 | 45 | 1 | 24 | 26 | 20 |
N
DIA
| 39 | 1 | 44 | 9 | 25 | 8 | 22 | 2 | 3 | 1 |
N
AS
| 87 | 88 | 102 | 129 | 46 | 101 | 54 | 62 | 46 | 41 |
N
F
| 30 | 36 | 30 | 47 | 31 | 49 | 26 | 47 | 64 | 51 |
N
CM
| 7 | 4 | 5 | 9 | 7 | 6 | 5 | 7 | 8 | 8 |
N
M
| 35 | 42 | 45 | 66 | 52 | 59 | 43 | 46 | 36 | 30 |
N
X
| 20 | 26 | 22 | 45 | 54 | 53 | 21 | 36 | 23 | 20 |
N
W
| 12 | 16 | 25 | 36 | 10 | 18 | 20 | 4 | 5 | 8 |
N
S
| 31 | 38 | 36 | 46 | 39 | 34 | 28 | 22 | 26 | 3 |
N
R
| 64 | 63 | 69 | 95 | 65 | 74 | 70 | 49 | 47 | 34 |
N
UN
| 19 | 17 | 23 | 27 | 17 | 28 | 13 | 9 | 13 | 30 |
I
AN
| 39.7 | 36.7 | 39.7 | 35.0 | 30.7 | 31.5 | 21.0 | 24.5 | 27.9 | 24.0 |
I
AR
| 17.8 | 20 | 17.1 | 35 | 16.7 | 15 | 10 | 13 | 15 | 11.7 |
I
KN
| 19.6 | 16 | 19.8 | 16 | 13.3 | 14 | 11 | 12 | 10.5 | 11.7 |
I
M
| 5.7 | 43.2 | 6.9 | 46.5 | 2.2 | 44.6 | 48.8 | 44.4 | 2.2 | 41.9 |
Variable | ||||||||
---|---|---|---|---|---|---|---|---|
PU | HH | HL | U | A | F | Ro | Ra | |
Axis 1 | 0.58*** | −0.33 | 0.77*** | 0.79*** | −0.60** | −0.74*** | 0.58** | −0.36 |
Axis 2 | −0.73** | 0.43 | −0.44 | −0.46* | 0.66** | −0.10 | −0.15 | −0.4 |
Relation between flora and urban indicators
Variable | Explains (%) | Contribution (%) |
F value |
P value |
---|---|---|---|---|
Ro | 18.9 | 35.6 | 4.2 | 0.014 |
Ra | 15.3 | 28.9 | 4.0 | 0.024 |
HL | 10.3 | 19.5 | 3.0 | 0.052 |
A | 5.0 | 9.5 | 1.5 | 0.252 |
HH | 2.9 | 5.4 | 0.8 | 0.468 |
PU | 0.6 | 1.1 | 0.2 | 0.902 |
Variable | Explains (%) | Contribution (%) |
F value |
P value |
---|---|---|---|---|
HH | 16.9 | 43.1 | 3.7 | 0.024 |
HL | 6.8 | 17.4 | 1.5 | 0.043 |
Ra | 6.0 | 15.4 | 1.4 | 0.244 |
A | 4.0 | 10.2 | 0.9 | 0.490 |
PU | 3.5 | 8.9 | 0.8 | 0.518 |
Ro | 1.9 | 5.0 | 0.4 | 0.816 |