Supporting information

This is the R Markdown document to run the analyses of the paper Lenoir, J. et al. Historical continuity and spatial connectivity ensure hedgerows are effective corridors for forest plants: evidence from the species-time-area relationship. Journal of Vegetation Science. To access the paper, go to the journal website.

List of packages used

#
library(knitr)
library(kableExtra)
library(sf)
library(leaflet)
library(mapview)
library(viridis)
library(Hmisc)
library(corrplot)
library(MuMIn)
library(jtools)
library(lme4)
library(gridExtra)
library(tidyverse)
#

A template for figures

Here is the code for the figure template used in the paper.

#
fig <- theme_classic() +
  theme(legend.position=c(1, 1), 
        legend.direction="vertical", 
        legend.title=element_text(face="bold"),
        legend.justification=c(1, 1),
        legend.title.align=0) +
  theme(axis.title.x=element_text(face="bold"), 
        axis.title.y=element_text(face="bold"),
        plot.tag=element_text(face="bold"),
        plot.title=element_text(face="bold"))
#

Import hedgerows’ data

The first file to import is entitled “data_carac_haie.txt”. It contains information on each of the 99 sampled hedgerows, such as hedgerow age, length, width, etc.

#
carac <- read.table("data_carac_haie.txt", header=TRUE, sep="\t")
carac <- as_tibble(carac)
#
Table 1: Characteristics of the 99 sampled hedgerows
id_haie l93_x l93_y annee_inf annee_sup annee_med annee_amp type h_max h_min h_moy h_amp longueur largeur multi_bande nb_bande connection dist_for_500 surf_for_500 dist_for surf_for_1000 elem_adj1 elem_adj2 elem_adj ipa_adj1 ipa_adj2 ipa_adj
beaum_h1 637497 7004925 1829 1829 1829 0 3 15.0 6.0 11.0 9 153 2.0 0 1 0 90 22444 90 22444 culture pature culture_pature 5 4 9
beaum_h2 637108 7004805 1829 1829 1829 0 3 8.0 4.0 6.0 4 81 2.0 0 1 0 204 17677 204 17677 chemin pature chemin_autre 1 4 5
berna_h1 640312 7003394 1807 1807 1807 0 3 14.0 4.0 9.0 10 204 2.0 0 1 0 60 20197 60 29003 pature chemin chemin_autre 4 1 5
berna_h2 640555 7003541 1807 1835 1821 28 3 16.0 6.0 12.0 10 131 2.0 0 1 0 134 19386 134 19386 pature pature pature_pature 4 4 8
berna_h3 640784 7003623 1807 1807 1807 0 1 1.5 1.5 1.5 0 60 1.0 0 1 0 181 12624 181 12624 route pature pature_route 2 4 6
berna_h4 639543 7003989 1835 1835 1835 0 3 18.0 10.0 14.0 8 219 2.0 0 1 0 114 64048 114 150281 chemin pature chemin_autre 1 4 5
boutav_h1 610531 6949644 1700 1800 1750 100 2 2.0 2.0 2.0 0 247 1.0 1 44 0 6 105008 6 116814 pature pature pature_pature 4 4 8
boutav_h2 610703 6949575 1800 1947 1874 147 3 12.0 4.0 8.0 8 80 2.0 1 14 0 58 141963 58 163351 pature pature pature_pature 4 4 8
boutav_h3 610445 6949433 1800 1947 1874 147 3 8.0 3.0 6.0 5 144 2.0 0 1 0 168 90538 168 100909 culture culture culture_culture 5 5 10
boutav_h4 610575 6949246 1947 1975 1961 28 2 8.0 8.0 8.0 0 192 2.0 0 1 0 233 160520 233 170890 pature route pature_route 4 2 6
boutav_h5 610030 6949613 1700 1800 1750 100 1 2.0 2.0 2.0 0 187 1.0 0 1 0 293 6471 293 6471 route pature pature_route 2 4 6
boutav_h6 610061 6949330 1800 1947 1874 147 3 15.0 3.0 9.0 12 150 2.0 0 1 0 430 5667 430 7987 culture pature culture_pature 5 4 9
boutav_h7 611780 6949236 1800 1947 1874 147 2 6.0 6.0 6.0 0 310 1.5 0 1 0 0 79926 0 449585 pature culture culture_pature 4 5 9
boutav_h8 610652 6950209 1992 1999 1996 7 2 5.0 5.0 5.0 0 68 0.5 0 1 0 158 42423 158 42423 culture pature culture_pature 5 4 9
boutav_h9 610458 6949962 1700 1800 1750 100 3 12.0 6.0 9.0 6 54 2.0 0 1 0 0 46630 0 48949 culture pature culture_pature 5 4 9
dom_h1 623172 7011575 1999 2001 2000 2 2 12.0 12.0 12.0 0 71 1.0 0 1 0 209 24261 209 127249 culture pature culture_pature 5 4 9
dom_h2 623221 7012065 1790 1947 1869 157 3 18.0 2.0 10.0 16 175 1.0 0 1 0 151 86840 151 221240 culture prairie culture_prairie 5 4 9
dom_h3 622878 7012112 1947 1975 1961 28 2 8.0 8.0 8.0 0 88 1.0 0 1 0 149 6970 149 38750 pature pature pature_pature 4 4 8
dom_h4 622768 7012215 1790 1947 1869 157 3 16.0 10.0 13.0 6 132 2.0 0 1 0 278 6970 278 36977 prairie chemin chemin_autre 4 1 5
dom_h5 622856 7012381 1947 1975 1961 28 1 3.0 3.0 3.0 0 60 3.0 0 1 0 447 918 447 131569 culture pature culture_pature 5 4 9
dom_h6 622223 7012289 1790 1947 1869 157 3 20.0 6.0 13.0 14 224 2.0 0 1 0 6 94572 6 133970 chemin pature chemin_autre 1 4 5
domq_h1 630719 7002291 1807 1807 1807 0 3 15.0 6.0 11.0 9 215 2.0 0 1 0 11 263161 11 323155 pature pature pature_pature 4 4 8
fla_h1 765820 6988883 1940 1975 1958 35 1 6.0 6.0 6.0 0 187 1.5 0 1 0 NA 0 2608 0 pature pature pature_pature 4 4 8
fla_h10 763193 6989754 1700 1750 1725 50 4 20.0 8.0 15.0 12 156 3.0 1 20 1 0 334948 0 910788 pature pature pature_pature 4 4 8
fla_h11 762850 6988595 1700 1750 1725 50 4 7.0 7.0 7.0 0 206 2.0 1 11 1 0 334979 0 1310236 culture pature culture_pature 5 4 9
fla_h12 762856 6989280 1700 1750 1725 50 3 15.0 5.0 10.0 10 92 3.0 1 17 1 0 444876 0 1266259 pature pature pature_pature 4 4 8
fla_h13 763182 6989488 1700 1750 1725 50 3 16.0 5.0 10.0 11 229 2.0 0 1 0 160 258495 160 727683 pature pature pature_pature 4 4 8
fla_h14 762904 6989114 1700 1750 1725 50 2 15.0 10.0 12.0 5 142 2.0 1 18 0 93 404284 93 1144963 pature pature pature_pature 4 4 8
fla_h15 763000 6989051 1700 1750 1725 50 3 18.0 4.0 11.0 14 239 3.0 1 41 0 95 308440 95 922246 pature pature pature_pature 4 4 8
fla_h16 763729 6989623 1700 1750 1725 50 2 8.0 6.0 7.0 2 121 3.0 1 19 0 NA 0 949 2446 pature pature pature_pature 4 4 8
fla_h17 763582 6989623 1700 1750 1725 50 3 25.0 6.0 15.0 19 82 4.0 1 15 0 451 9392 451 53011 pature pature pature_pature 4 4 8
fla_h18 762973 6989435 1700 1750 1725 50 3 20.0 8.0 15.0 12 152 3.0 1 34 1 0 420894 0 1185741 pature pature pature_pature 4 4 8
fla_h19 765320 6991616 1700 1750 1725 50 3 16.0 8.0 12.0 8 103 3.5 1 19 1 0 371888 0 371888 chemin pature chemin_autre 1 4 5
fla_h2 765851 6988804 1991 1996 1994 5 1 6.0 6.0 6.0 0 142 1.0 0 1 0 NA 0 2559 0 prairie prairie prairie_prairie 4 4 8
fla_h20 763217 6989818 1700 1750 1725 50 2 18.0 10.0 14.0 8 166 1.5 1 28 1 0 324147 0 895300 pature pature pature_pature 4 4 8
fla_h3 765795 6988742 1940 1975 1958 35 1 6.0 6.0 6.0 0 124 1.0 0 1 0 NA 0 2578 0 prairie culture culture_prairie 4 5 9
fla_h4 765554 6988485 1836 1940 1888 104 1 2.0 2.0 2.0 0 211 2.0 0 1 0 NA 0 2677 0 pature pature pature_pature 4 4 8
fla_h5 765298 6988797 1700 1750 1725 50 3 16.0 3.0 10.0 13 226 2.5 0 1 0 NA 0 2470 0 pature prairie pature_prairie 4 4 8
fla_h6 765651 6989098 1750 1836 1793 86 1 6.0 6.0 6.0 0 187 2.0 0 1 0 NA 0 2852 0 pature pature pature_pature 4 4 8
fla_h7 764854 6990244 2006 2010 2008 4 1 4.0 4.0 4.0 0 197 0.5 0 1 0 NA 0 1815 0 pature pature pature_pature 4 4 8
fla_h8 765007 6989954 1750 1836 1793 86 3 15.0 6.0 12.0 9 137 1.0 0 1 0 NA 0 1955 0 pature pature pature_pature 4 4 8
fla_h9 764508 6990472 1836 1940 1888 104 2 12.0 12.0 12.0 0 206 1.0 0 1 0 NA 0 1433 0 pature pature pature_pature 4 4 8
fless_h1 648602 6984518 1997 2001 1999 4 1 2.0 2.0 2.0 0 470 2.0 0 1 0 43 77307 43 190568 culture culture culture_culture 5 5 10
la_bou_h1 770902 6976530 1700 1750 1725 50 3 16.0 8.0 12.0 8 206 1.0 0 1 0 15 91503 15 353562 pature culture culture_pature 4 5 9
la_bou_h2 771021 6976764 1700 1750 1725 50 3 18.0 6.0 12.0 12 225 2.0 0 1 0 203 92112 203 380555 culture pature culture_pature 5 4 9
la_bou_h3 769881 6974040 1750 1939 1845 189 3 16.0 6.0 11.0 10 123 2.0 0 1 0 NA 0 978 674 pature pature pature_pature 4 4 8
la_bou_h4 768995 6974206 1996 2001 1999 5 1 6.0 6.0 6.0 0 473 1.0 0 1 0 NA 0 1375 0 pature pature pature_pature 4 4 8
la_bou_h5 768727 6975647 1750 1939 1845 189 2 15.0 15.0 15.0 0 224 1.5 0 1 0 360 5141 360 5249 pature pature pature_pature 4 4 8
la_bou_h6 768673 6975734 1700 1750 1725 50 3 20.0 8.0 14.0 12 198 3.5 0 1 0 332 5754 332 9200 prairie chemin chemin_autre 4 1 5
la_chap_h1 618320 6937712 1700 1800 1750 100 3 10.0 3.0 7.0 7 98 4.0 0 1 0 178 7998 178 7998 pature chemin chemin_autre 4 1 5
la_chap_h2 617900 6937367 1975 1986 1981 11 3 8.0 3.0 6.0 5 202 1.0 1 5 1 0 110759 0 112064 culture culture culture_culture 5 5 10
la_chap_h3 617818 6937180 1800 1947 1874 147 1 15.0 15.0 15.0 0 105 2.0 1 16 1 0 110759 0 170193 culture culture culture_culture 5 5 10
la_chap_h4 617539 6937109 1800 1947 1874 147 2 10.0 4.0 7.0 6 208 2.0 0 1 0 36 118809 36 271837 pature pature pature_pature 4 4 8
lam_h1 616266 7010452 1806 1806 1806 0 3 12.0 3.0 7.0 9 178 3.0 0 1 0 228 815 228 815 pature pature pature_pature 4 4 8
lam_h2 616142 7010432 1806 1806 1806 0 3 18.0 6.0 12.0 12 117 2.0 0 1 0 386 815 386 815 pature culture culture_pature 4 5 9
lam_h3 616863 7010203 1997 2001 1999 4 2 16.0 16.0 16.0 0 107 2.0 0 1 0 236 5916 236 5916 pature culture culture_pature 4 5 9
lam_h4 616935 7010434 1806 1947 1877 141 1 2.0 2.0 2.0 0 65 1.0 0 1 0 179 5916 179 5916 pature pature pature_pature 4 4 8
lam_h5 617057 7010440 1806 1947 1877 141 3 16.0 4.0 10.0 12 119 2.0 0 1 0 76 5916 76 5916 pature pature pature_pature 4 4 8
lam_h6 615987 7010836 1806 1947 1877 141 3 20.0 6.0 13.0 14 132 2.0 0 1 0 429 815 429 815 pature pature pature_pature 4 4 8
le_boi_h1 627968 7019264 1773 1947 1860 174 3 4.0 20.0 12.0 -16 78 3.0 1 18 1 0 262807 0 392824 chemin pature chemin_autre 1 4 5
le_boi_h2 627573 7020272 1772 1774 1773 2 2 15.0 15.0 15.0 0 180 2.0 1 40 0 176 16529 176 192690 pature culture culture_pature 4 5 9
marc_h1 670124 6972320 2002 2002 2002 0 2 8.0 5.0 7.0 3 433 2.0 1 42 0 NA 0 929 4802 culture culture culture_culture 5 5 10
marc_h2 670219 6971161 2002 2002 2002 0 2 10.0 4.0 7.0 6 279 2.0 1 28 0 36 177556 36 392874 culture culture culture_culture 5 5 10
marc_h3 667914 6974131 2002 2002 2002 0 2 13.0 8.0 10.0 5 259 2.0 1 19 0 NA 0 2248 0 culture culture culture_culture 5 5 10
marc_h4 670717 6972214 2002 2002 2002 0 2 12.0 8.0 10.0 4 368 2.0 1 28 0 169 170282 169 219868 culture culture culture_culture 5 5 10
marc_h5 670552 6971058 2002 2002 2002 0 2 8.0 3.0 6.0 5 215 2.0 1 16 0 117 202315 117 410141 culture culture culture_culture 5 5 10
marc_h6 669539 6973809 2002 2002 2002 0 2 15.0 9.0 12.0 6 275 2.0 1 9 0 NA 0 2221 0 culture culture culture_culture 5 5 10
mon_h1 612404 6950594 1700 1800 1750 100 1 1.8 1.8 1.8 0 171 0.8 1 34 1 0 324580 0 717664 pature pature pature_pature 4 4 8
mon_h2 612954 6951606 1800 1947 1874 147 2 12.0 12.0 12.0 0 119 3.0 1 23 1 0 82743 0 137675 culture pature culture_pature 5 4 9
mon_h3 611879 6950951 1986 1989 1988 3 1 6.0 6.0 6.0 0 71 1.0 1 16 0 454 2764 454 121258 culture culture culture_culture 5 5 10
mon_h4 612295 6951134 1947 1975 1961 28 1 4.0 4.0 4.0 0 51 1.0 0 1 0 395 10099 395 58813 pature pature pature_pature 4 4 8
ons_h1 621065 6924502 1936 1950 1943 14 3 15.0 4.0 10.0 11 104 2.0 0 1 0 156 7035 156 7035 pature pature pature_pature 4 4 8
ons_h2 621279 6924498 2001 2006 2004 5 1 4.0 4.0 4.0 0 103 2.0 0 1 0 261 7714 261 7714 pature pature pature_pature 4 4 8
ons_h3 621199 6924720 1936 1950 1943 14 3 15.0 3.0 9.0 12 164 3.0 0 1 0 128 12828 128 12828 pature pature pature_pature 4 4 8
ons_h4 621878 6925117 1950 1975 1963 25 3 15.0 5.0 10.0 10 75 5.0 0 1 0 80 66149 80 83743 pature pature pature_pature 4 4 8
ons_h5 622559 6925398 1950 1975 1963 25 1 6.0 6.0 6.0 0 84 1.5 0 1 0 337 47290 337 64884 pature pature pature_pature 4 4 8
ons_h6 621679 6923862 1936 1950 1943 14 3 18.0 10.0 14.0 8 168 3.0 0 1 0 NA 0 637 110727 pature culture culture_pature 4 5 9
ons_h7 622215 6923592 1936 1950 1943 14 2 8.0 8.0 8.0 0 105 4.0 0 1 0 NA 0 891 25164 pature pature pature_pature 4 4 8
prou_h1 638065 7005849 1835 1835 1835 0 2 14.0 4.0 9.0 10 111 1.0 0 1 0 170 10470 170 10470 culture pature culture_pature 5 4 9
sor_h1 764375 6978041 1800 1866 1833 66 1 6.0 6.0 6.0 0 317 2.5 0 1 0 11 70552 11 133952 pature pature pature_pature 4 4 8
sor_h2 763566 6977695 1700 1800 1750 100 3 12.0 6.0 9.0 6 101 2.0 0 1 0 377 23427 377 339802 pature culture culture_pature 4 5 9
sor_h3 763793 6977684 1700 1800 1750 100 3 15.0 2.0 8.0 13 52 2.0 0 1 0 176 39263 176 227900 pature chemin chemin_autre 4 1 5
sor_h4 763874 6977829 1700 1800 1750 100 3 16.0 6.0 11.0 10 83 3.0 0 1 0 119 43418 119 233117 pature chemin chemin_autre 4 1 5
sor_h5 763987 6976777 1800 1826 1813 26 3 16.0 4.0 10.0 12 163 3.0 0 1 0 NA 0 1011 0 pature culture culture_pature 4 5 9
st_ar_h1 613807 6948769 1992 1997 1995 5 1 4.0 4.0 4.0 0 99 2.0 0 1 0 175 62143 175 81077 pature chemin chemin_autre 4 1 5
st_ar_h2 613641 6948837 1800 1947 1874 147 1 8.0 8.0 8.0 0 116 4.0 1 20 1 0 101136 0 101136 pature culture culture_pature 4 5 9
st_ar_h3 614013 6948722 1800 1947 1874 147 1 6.0 6.0 6.0 0 143 3.0 0 1 0 248 39514 248 130426 pature chemin chemin_autre 4 1 5
st_ar_h4 614071 6948739 1800 1947 1874 147 2 14.0 14.0 14.0 0 56 2.0 0 1 0 245 28246 245 132579 pature pature pature_pature 4 4 8
st_ar_h5 614002 6948624 1947 1975 1961 28 1 8.0 8.0 8.0 0 83 2.0 0 1 0 305 20825 305 62639 pature pature pature_pature 4 4 8
st_ar_h6 614468 6948722 1700 1800 1750 100 3 12.0 3.0 8.0 9 135 3.0 0 1 0 195 8440 195 188286 pature prairie pature_prairie 4 4 8
st_ar_h7 613047 6949441 1800 1947 1874 147 3 10.0 3.0 6.0 7 119 1.5 1 18 1 0 158222 0 386874 pature prairie pature_prairie 4 4 8
st_ar_h8 613173 6949590 1800 1947 1874 147 1 1.8 1.8 1.8 0 90 1.0 0 1 1 0 129735 0 333762 pature pature pature_pature 4 4 8
st_ar_h9 612384 6948563 1800 1947 1874 147 2 14.0 14.0 14.0 0 104 2.0 0 1 0 NA 0 989 823 pature pature pature_pature 4 4 8
st_germ_h1 611497 6928305 1782 1782 1782 0 3 18.0 6.0 12.0 12 97 2.0 0 1 0 254 93297 254 377991 pature pature pature_pature 4 4 8
st_germ_h2 610841 6927898 1782 1947 1865 165 3 8.0 3.0 6.0 5 269 2.0 0 1 0 163 35548 163 188341 pature pature pature_pature 4 4 8
st_germ_h3 610819 6928159 1782 1782 1782 0 3 7.0 3.0 5.0 4 196 1.0 0 1 0 401 11388 401 198464 pature pature pature_pature 4 4 8
vsver_h1 608349 6941967 1945 1975 1960 30 2 20.0 4.0 12.0 16 54 2.0 0 1 0 242 3485 242 29676 pature pature pature_pature 4 4 8
vsver_h2 608036 6943127 1784 1947 1866 163 2 10.0 2.0 6.0 8 162 1.0 1 14 1 0 394114 0 1141877 pature pature pature_pature 4 4 8
vsver_h3 609061 6943100 1784 1947 1866 163 2 18.0 4.0 11.0 14 144 3.0 1 12 1 0 258870 0 620342 pature pature pature_pature 4 4 8

Import the floristic data

The second file to import is entitled “data_flo_haie.txt”. It contains the floristic composition of each hedgerow and is displayed as a species-by-hedgerow matrix. Note that some hedgerows are divided into several and consecutive 5-m segments providing the species composition of each 5-m segment.

#
flo <- read.table("data_flo_haie_pa.txt", header=TRUE, sep="\t")
flo <- as_tibble(flo)
#
Table 2: Species composition matrix of the 99 sampled hedgerows
id_haie id_bande multi_bande xl93_bande yl93_bande dist_bande num_bande largeur_bande rich_bande A_Acer_campestre A_Acer_platanoides A_Acer_pseudoplatanus A_Alnus_glutinosa A_Betula_pendula A_Bryonia_dioica A_Buxus_sempervirens A_Calystegia_sepium A_Carpinus_betulus A_Clematis_vitalba A_Cornus_mas A_Cornus_sanguinea_.Swida_sanguinea. A_Corylus_avellana A_Crataegus_laevigata A_Crataegus_monogyna A_Euonymus_europaeus A_Fagus_sylvatica A_Frangula_dodonei_.Frangula_alnus. A_Fraxinus_excelsior A_Hedera_helix A_Humulus_lupulus A_Ilex_aquifolium A_Juglans_regia A_Ligustrum_vulgare A_Lonicera_caprifolium A_Lonicera_periclymenum A_Malus_sp. A_Malus_sylvestris A_Mespilus_germanica A_Populus_nigra A_Populus_tremula A_Prunus_avium A_Prunus_domestica_ssp._insititia A_Prunus_spinosa A_Pyrus_sp. A_Quercus_pedunculata A_Quercus_petraea A_Quercus_robur A_Rhamnus_cathartica A_Ribes_rubrum A_Ribes_uva.crispa A_Robinia_pseudoacacia A_Rosa_arvensis A_Rosa_canina A_Rosa_sp. A_Rosa_stylosa A_Rosa_tomentosa A_Rubus_caesius A_Rubus_fruticosus_agg. A_Rubus_idaeus A_Rubus_sp. A_Salix_alba A_Salix_caprea A_Salix_cinerea A_Salix_fragilis A_Salix_triandra A_Sambucus_ebulus A_Sambucus_nigra A_Solanum_dulcamara A_Sorbus_aucuparia A_Stachys_sylvatica A_Staphylea_pinnata A_Taxus_baccata A_Tilia_cordata A_Tilia_platyphyllos A_Ulmus_glabra A_Ulmus_minor A_Viburnum_lantana A_Viburnum_opulus A_Viscum_album H_Acer_campestre H_Acer_pseudoplatanus H_Adoxa_moschatellina H_Agrimonia_eupatoria H_Ajuga_reptans H_Alliaria_petiolata H_Allium_ursinum H_Anemone_nemorosa H_Anthriscus_sylvestris H_Arctium_lappa H_Arum_maculatum H_Athyrium_filix.femina H_Aegopodium_podagraria H_Balota_fetida H_Betula_pubescens H_Brachypodium_pinnatum H_Brachypodium_sylvaticum H_Bromus_ramosus H_Bryonia_dioica H_Calystegia_sepium H_Campanula_trachelium H_Cardamine_hirsuta H_Cardamine_pratensis H_Carex_spicata_agg. H_Carex_sylvatica H_Carpinus_betulus H_Castanea_sativa H_Cerastium_fontanum H_Cerastium_sp. H_Chaerophyllum_temulum H_Chelidonium_majus H_Circaea_lutetiana H_Cornus_mas H_Cornus_sanguinea_.Swida_sanguinea. H_Corylus_avellana H_Crataegus_laevigata H_Crataegus_monogyna H_Deschampsia_cespitosa H_Dryopteris_affinis H_Dryopteris_carthusiana H_Dryopteris_filix.mas H_Elymus_caninus H_Epilobium_angustifolium H_Epilobium_hirsutum H_Epilobium_montanum H_Epipactis_helleborine H_Equisetum_arvense H_Euonymus_europaeus H_Euphorbia_amygdaloides H_Fagus_sylvatica H_Festuca_gigantea H_Filipendula_ulmaria H_Fraxinus_excelsior H_Galanthus_nivalis H_Galeopsis_tetrahit H_Galium_aparine H_Galium_odoratum_.Asperula_odorata. H_Geranium_molle H_Geranium_robertianum H_Geum_urbanum H_Glechoma_hederacea H_Hedera_helix H_Helleborus_viridis H_Heracleum_sphondylium H_Hieracium_murorum_.Hieracium_sylvaticum. H_Holcus_lanatus H_Holcus_mollis H_Humulus_lupulus H_Hyacinthoides_non.scripta H_Hypericum_hirsutum H_Hypericum_pulchrum H_Hypericum_perforatum H_Ilex_aquifolium H_Lamium_galeobdolon_.Galeobdolon_luteum. H_Lamium_purpureum H_Lapsana_communis H_Lonicera_periclymenum H_Luzula_multiflora H_Luzula_pilosa H_Lysimachia_nemorum H_Lysimachia_vulgaris H_Melica_uniflora H_Mercurialis_perennis H_Milium_effusum H_Moehringia_trinervia H_Oxalis_acetosella H_Paris_quadrifolia H_Philadelphus_coronarius H_Poa_nemoralis H_Poa_trivialis H_Polygonatum_multiflorum H_Polypodium_interjectum H_Potentilla_sterilis H_Primula_elatior H_Primula_veris H_Prunella_vulgaris H_Prunus_avium H_Prunus_domestica_ssp._insititia H_Prunus_spinosa H_Pteridium_aquilinum H_Quercus_robur H_Ranunculus_auricomus_agg. H_Ranunculus_ficaria H_Ranunculus_repens H_Ribes_rubrum H_Ribes_uva.crispa H_Rosa_arvensis H_Rosa_canina H_Rosa_sp. H_Rosa_tomentosa H_Rubus_fruticosus_agg. H_Rubus_idaeus H_Rumex_obtusi H_Rumex_sanguineus H_Sambucus_nigra H_Sambucus_racemosa H_Sonchus_oleraceus H_Scilla_bifolia H_Scrophularia_nodosa H_Senecio_ovatus_ssp._ovatus H_Silene_dioica_.Melandrium_dioicum. H_Solanum_dulcamara H_Sorbus_aucuparia H_Stachys_alpina H_Stachys_sylvatica H_Stellaria_holostea H_Symphytum_officinale H_Syringa_vulgaris H_Tamus_communis H_Taraxacum_sp. H_Taxus_baccata H_Teucrium_scorodonia H_Ulmus_minor H_Urtica_dioica H_Valeriana_officinalis_.Valeriana_repens. H_Veronica_chamaedrys H_Veronica_hederifolia H_Veronica_montana H_Veronica_officinalis H_Vicia_cracca H_Vicia_sepium H_Vinca_minor H_Viola_hirta H_Viola_odorata H_Viola_reichenbachiana H_Viola_riviniana H_Viscum_album
marc_h1 marc_h1-1 1 670081.4 6972110 772.163822 1 3.0 3 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0
marc_h1 marc_h1-2 1 670082.4 6972115 771.911277 2 3.0 4 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
marc_h1 marc_h1-3 1 670083.4 6972120 771.691130 3 3.0 5 0 0 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
marc_h1 marc_h1-4 1 670084.4 6972125 771.503127 4 3.0 3 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
marc_h1 marc_h1-5 1 670085.4 6972130 771.347606 5 3.0 7 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
marc_h1 marc_h1-6 1 670086.4 6972135 771.224453 6 2.0 3 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
marc_h1 marc_h1-7 1 670087.4 6972139 771.133700 7 2.0 3 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
marc_h1 marc_h1-8 1 670088.4 6972144 771.075378 8 2.0 3 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
marc_h1 marc_h1-9 1 670089.4 6972149 771.049457 9 2.0 3 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
marc_h1 marc_h1-10 1 670090.4 6972154 771.046583 10 2.0 3 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
marc_h1 marc_h1-11 1 670091.4 6972159 771.044638 11 2.0 3 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
marc_h1 marc_h1-12 1 670092.4 6972164 771.042713 12 2.0 5 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
marc_h1 marc_h1-13 1 670093.4 6972169 771.040768 13 2.0 5 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
marc_h1 marc_h1-14 1 670094.4 6972174 771.038823 14 2.0 6 0 0 0 0 0 0 0 0 1 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
marc_h1 marc_h1-15 1 670095.4 6972179 771.036898 15 2.0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
marc_h1 marc_h1-16 1 670096.4 6972184 771.034953 16 2.0 5 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0
marc_h1 marc_h1-17 1 670097.4 6972188 771.033008 17 2.0 6 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0
marc_h1 marc_h1-18 1 670098.4 6972193 771.031181 18 2.0 5 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
marc_h1 marc_h1-19 1 670099.4 6972198 771.029139 19 2.0 5 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
marc_h1 marc_h1-20 1 670100.4 6972203 771.027194 20 2.0 4 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
marc_h1 marc_h1-21 1 670101.4 6972208 771.025269 21 2.0 4 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
marc_h1 marc_h1-22 1 670102.4 6972213 771.023324 22 2.0 6 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0
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vsver_h3 vsver_h3-1 1 609127.4 6943119 2.499993 1 5.0 19 0 0 0 0 0 0 0 0 1 0 0 0 1 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0
vsver_h3 vsver_h3-2 1 609122.7 6943117 7.500015 2 5.0 24 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 0 0 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0
vsver_h3 vsver_h3-3 1 609118.0 6943115 12.500001 3 5.0 32 0 0 0 0 0 0 0 0 1 0 0 0 1 1 0 0 1 0 1 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 1 0 0 0 1 0 1 0 0 0 0 1 0 0 0 1 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 1 1 0
vsver_h3 vsver_h3-4 1 609113.3 6943113 17.500023 4 5.0 25 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 1 0 1 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 1 0 0 0 0 0 0 0 0 1 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 1 0
vsver_h3 vsver_h3-5 1 609108.7 6943112 22.500009 5 5.0 28 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 1 0 0 1 0 1 0 0 0 1 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 1 0 0 0 1 0 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 1 0
vsver_h3 vsver_h3-6 1 609104.0 6943110 27.500031 6 5.0 18 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 1 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0
vsver_h3 vsver_h3-7 1 609099.3 6943108 32.480366 7 5.0 23 0 0 0 0 0 0 0 0 1 0 0 0 1 0 1 0 1 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 1 0
vsver_h3 vsver_h3-8 1 609094.4 6943107 37.425941 8 5.0 22 0 0 0 0 0 0 0 0 1 0 0 0 1 0 1 0 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 0 0
vsver_h3 vsver_h3-9 1 609089.5 6943106 42.364779 9 5.0 25 0 0 0 0 0 0 0 0 1 0 0 0 1 1 1 0 0 0 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 0 0 1 0 0 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 1 0
vsver_h3 vsver_h3-10 1 609084.6 6943105 47.316475 10 5.0 23 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 0 1 0 1 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 0 1 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0
vsver_h3 vsver_h3-11 1 609079.6 6943104 51.083897 11 5.0 39 1 0 0 0 0 0 0 0 1 0 0 0 1 0 0 1 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0 0 1 0 1 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 1 0 0 1 1 0 1 0 0 0 0 0 0 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 1 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 1 1 0 0 1 0 0 0 1 1 0
vsver_h3 vsver_h3-12 1 609074.7 6943104 53.267266 12 5.0 35 1 0 0 0 0 0 0 0 1 0 0 0 1 0 1 0 0 0 1 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 1 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 1 0 0 1 1 0 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 1 0 1 0 1 0 0 1 0 0 0 1 1 0
ons_h1 ons_h1-1 0 NA NA NA NA NA NA 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 1 1 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 1 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 1 0 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 1 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 0 0 0 0 0 0 0 0 0 1
ons_h2 ons_h2-1 0 NA NA NA NA NA NA 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 1 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 0 0 0 0 1
ons_h3 ons_h3-1 0 NA NA NA NA NA NA 0 0 0 0 0 0 0 0 0 1 0 0 0 1 1 1 0 0 1 1 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 0 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 1 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 0 1 0 0 0
ons_h4 ons_h4-1 0 NA NA NA NA NA NA 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 0 0 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 1 0 0 0 0 1 0 0 0 0 1 0 0 0 1 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0
ons_h5 ons_h5-1 0 NA NA NA NA NA NA 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 1 0 0 0 0 1 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 1 1 0 1 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 1 0 1 0 1 0 0 0 0 0 0 0 0 0
ons_h6 ons_h6-1 0 NA NA NA NA NA NA 1 0 0 0 0 0 0 0 1 1 0 1 0 1 1 1 0 0 1 1 0 0 0 0 0 0 0 0 0 0 1 0 1 1 0 0 0 0 0 0 1 0 1 1 0 0 0 0 1 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 1 0 0 0 0 0 0 0 0 0 0
ons_h7 ons_h7-1 0 NA NA NA NA NA NA 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 1 0 0 0 1 0 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 1 1 1 0 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 1 0 0 0 0 0 0 0 0 0 0

Species list

Using the floristic composition of each hedgerow, we compiled a list of species occurring across the 99 sampled hedgerows. For each species, we also computed its frequency across the 99 sampled hedgerows in both the herbaceous (\(\le 1\)-m height) and tree (\(> 1\)-m height) vegetation layers.

#
flo %>%
  select(-c(multi_bande,
            xl93_bande,
            yl93_bande,
            dist_bande,
            num_bande, 
            largeur_bande, 
            rich_bande)) %>%
  gather(key="id_taxon", value="pa", -c(id_bande, id_haie)) %>%
  filter(pa==1) %>%
  select(-c(id_bande)) %>%
  distinct() %>% 
  group_by(id_taxon) %>%
  summarize(n_tot=n_distinct(id_haie)) %>% 
  separate(id_taxon, into=c("veg_layer", "sp_name"), sep=2) %>% 
  spread(key="veg_layer", value="n_tot") %>% 
  rename(occ_tree=A_, occ_herb=H_) %>% 
  replace_na(list(occ_tree=0, occ_herb=0)) %>% 
  arrange(desc(occ_herb, occ_tree)) %>% 
  mutate_if(is.character, str_replace_all, pattern="_\\.", replacement=" ") %>% 
  mutate_if(is.character, str_remove_all, pattern=" .+") %>% 
  mutate_if(is.character, str_replace_all, pattern="([a-z])\\.([a-z])", replacement="\\1-\\2") %>% 
  mutate_if(is.character, str_replace_all, pattern="_", replacement=" ") -> species_list
#
Table 3: List of forest plant species occurring in the 99 sampled hedgerows
sp_name occ_tree occ_herb
Urtica dioica 0 89
Galium aparine 0 87
Hedera helix 51 86
Geum urbanum 0 67
Glechoma hederacea 0 65
Anthriscus sylvestris 0 64
Arum maculatum 0 64
Rubus fruticosus agg. 77 57
Lamium galeobdolon 0 56
Heracleum sphondylium 0 50
Geranium robertianum 0 49
Stachys sylvatica 1 48
Galeopsis tetrahit 0 39
Stellaria holostea 0 37
Lapsana communis 0 35
Solanum dulcamara 17 34
Chaerophyllum temulum 0 31
Veronica chamaedrys 0 31
Quercus robur 18 28
Poa trivialis 0 26
Ranunculus ficaria 0 25
Polygonatum multiflorum 0 24
Silene dioica 0 24
Viola reichenbachiana 0 24
Moehringia trinervia 0 23
Veronica hederifolia 0 23
Scrophularia nodosa 0 18
Adoxa moschatellina 0 17
Prunus spinosa 75 16
Aegopodium podagraria 0 15
Ajuga reptans 0 14
Ribes uva-crispa 29 14
Viola odorata 0 14
Alliaria petiolata 0 13
Poa nemoralis 0 13
Rosa arvensis 48 12
Vinca minor 0 12
Brachypodium sylvaticum 0 11
Holcus lanatus 0 11
Carex sylvatica 0 10
Dryopteris filix-mas 0 10
Fraxinus excelsior 69 10
Milium effusum 0 10
Sambucus nigra 78 10
Bryonia dioica 13 9
Potentilla sterilis 0 9
Rumex obtusi 0 9
Taraxacum sp. 0 9
Anemone nemorosa 0 8
Primula elatior 0 8
Prunus avium 14 8
Calystegia sepium 4 7
Carpinus betulus 63 7
Crataegus monogyna 85 7
Euonymus europaeus 45 7
Mercurialis perennis 0 7
Ranunculus repens 0 7
Veronica montana 0 7
Veronica officinalis 0 7
Chelidonium majus 0 6
Corylus avellana 54 6
Deschampsia cespitosa 0 6
Lonicera periclymenum 36 6
Tamus communis 0 6
Vicia sepium 0 6
Acer pseudoplatanus 20 5
Dryopteris carthusiana 0 5
Fagus sylvatica 12 5
Holcus mollis 0 5
Hyacinthoides non-scripta 0 5
Ilex aquifolium 41 5
Lysimachia nemorum 0 5
Melica uniflora 0 5
Ranunculus auricomus agg. 0 5
Rosa canina 62 5
Rumex sanguineus 0 5
Sorbus aucuparia 5 5
Circaea lutetiana 0 4
Crataegus laevigata 61 4
Festuca gigantea 0 4
Galanthus nivalis 0 4
Hypericum perforatum 0 4
Primula veris 0 4
Rubus idaeus 4 4
Acer campestre 40 3
Bromus ramosus 0 3
Campanula trachelium 0 3
Cerastium fontanum 0 3
Cornus sanguinea 23 3
Elymus caninus 0 3
Euphorbia amygdaloides 0 3
Filipendula ulmaria 0 3
Ribes rubrum 5 3
Symphytum officinale 0 3
Viola hirta 0 3
Viscum album 9 3
Athyrium filix-femina 0 2
Balota fetida 0 2
Carex spicata agg. 0 2
Cornus mas 9 2
Epilobium angustifolium 0 2
Epilobium montanum 0 2
Galium odoratum 0 2
Hypericum hirsutum 0 2
Oxalis acetosella 0 2
Pteridium aquilinum 0 2
Rosa tomentosa 5 2
Senecio ovatus ssp. ovatus 0 2
Stachys alpina 0 2
Valeriana officinalis 0 2
Agrimonia eupatoria 0 1
Allium ursinum 0 1
Arctium lappa 0 1
Betula pubescens 0 1
Brachypodium pinnatum 0 1
Cardamine hirsuta 0 1
Cardamine pratensis 0 1
Castanea sativa 0 1
Cerastium sp. 0 1
Dryopteris affinis 0 1
Epilobium hirsutum 0 1
Epipactis helleborine 0 1
Equisetum arvense 0 1
Geranium molle 0 1
Helleborus viridis 0 1
Hieracium murorum 0 1
Humulus lupulus 3 1
Hypericum pulchrum 0 1
Lamium purpureum 0 1
Luzula multiflora 0 1
Luzula pilosa 0 1
Lysimachia vulgaris 0 1
Paris quadrifolia 0 1
Philadelphus coronarius 0 1
Polypodium interjectum 0 1
Prunella vulgaris 0 1
Prunus domestica ssp. insititia 5 1
Rosa sp. 3 1
Sambucus racemosa 0 1
Scilla bifolia 0 1
Sonchus oleraceus 0 1
Syringa vulgaris 0 1
Taxus baccata 3 1
Teucrium scorodonia 0 1
Ulmus minor 22 1
Vicia cracca 0 1
Viola riviniana 0 1
Acer platanoides 2 0
Alnus glutinosa 2 0
Betula pendula 2 0
Buxus sempervirens 1 0
Clematis vitalba 15 0
Frangula dodonei 6 0
Juglans regia 3 0
Ligustrum vulgare 39 0
Lonicera caprifolium 3 0
Malus sp. 6 0
Malus sylvestris 1 0
Mespilus germanica 11 0
Populus nigra 2 0
Populus tremula 5 0
Pyrus sp. 2 0
Quercus pedunculata 1 0
Quercus petraea 1 0
Rhamnus cathartica 1 0
Robinia pseudoacacia 2 0
Rosa stylosa 1 0
Rubus caesius 1 0
Rubus sp. 1 0
Salix alba 6 0
Salix caprea 9 0
Salix cinerea 2 0
Salix fragilis 1 0
Salix triandra 2 0
Sambucus ebulus 1 0
Staphylea pinnata 2 0
Tilia cordata 3 0
Tilia platyphyllos 2 0
Ulmus glabra 1 0
Viburnum lantana 5 0
Viburnum opulus 4 0

Where are the hedgerows?

We built an interactive leaflet map to display the geographic location of each of the 99 hedgerows that we sampled across the Picardy region in northern France (Hauts-de-France).

#
crs_data <- rgdal::make_EPSG()
# 
tibble(id=carac$id_haie, 
       lon=carac$l93_x, 
       lat=carac$l93_y, 
       yr=carac$annee_med) %>%
  st_as_sf(coords=c("lon", "lat")) %>%
  st_set_crs(2154) -> hedgerow_coord_l93
#
hedgerow_coord_l93 %>%
  st_transform(4326) -> hedgerow_coord_lonlat
#
map_hedgerow <- leaflet() %>%
  addProviderTiles("Esri.WorldImagery") %>% 
  addProviderTiles(providers$Stamen.TonerLines, 
                   options=providerTileOptions(opacity=0.35)) %>%
  addProviderTiles(providers$Stamen.TonerLabels) %>%
  addMarkers(data=hedgerow_coord_lonlat, 
             clusterOptions=markerClusterOptions(), 
             popup=~as.character(yr), 
             label=~as.character(yr))
#
map_hedgerow

Figure 1: Distribution of the 99 hedgerows sampled across the study area in Picardy (France)

How many forest plants do we find in hedgerows?

Using the species-by-hedgerow matrix, we computed species richness (cf. total number of forest plant species) for each hedgerow, distinguishing between the herbaceous (\(\le 1\)-m height) and tree (\(> 1\)-m height) vegetation layers.

#
flo %>%
  select(-c(multi_bande,
            xl93_bande,
            yl93_bande,
            dist_bande,
            num_bande, 
            largeur_bande, 
            rich_bande)) %>%
  gather(key="id_taxon", value="pa", -c(id_bande, id_haie)) %>%
  filter(pa==1) %>%
  separate(id_taxon, into=c("layer", "name"), sep=1) %>%
  group_by(id_haie) %>%
  summarize(n_tot=n_distinct(name)) -> rich_all
#  
flo %>%
  select(-c(multi_bande, 
            num_bande, 
            largeur_bande, 
            rich_bande)) %>%
  gather(key="id_taxon", value="pa", -c(id_bande, id_haie)) %>%
  filter(pa==1) %>%
  separate(id_taxon, into=c("layer", "name"), sep=1) %>%  
  group_by(id_haie, layer) %>%
  summarize(rich=n_distinct(name)) %>%
  spread(key=layer, value=rich) %>%
  rename(n_tree=A, n_herb=H) -> rich_layer
#  
carac %>%  
  left_join(rich_all, by="id_haie") %>%
  left_join(rich_layer, by="id_haie") -> carac
#
carac %>%
  select(c(id_haie, 
           n_tree, 
           n_herb)) %>%
  rename(T=n_tree, 
         H=n_herb) %>%
  gather(key=layer, value=n, -id_haie) %>%
  mutate(layer=factor(layer)) -> richness
#
plot_r <- ggplot(data=richness, 
                 aes(x=n, 
                     fill=layer)) +
          geom_histogram(breaks=seq(2, 50, 2), 
                         alpha=0.5, 
                         col="white", 
                         position="identity") + 
          scale_y_continuous(limits=c(0, 20), 
                             breaks=seq(0, 20, 5), 
                             labels=c("0.0", "5.0", "10.0", "15.0", "20.0")) +
          scale_fill_manual(values=c("green4", "purple4"), 
                            name="Vegetation layer", 
                            labels=c("Below 1-m height", "Above 1-m height")) +
          labs(x="Species richness", 
               y="Number of hedgerows") +
          fig
#
plot_r

Figure 2: Histogram of the total number of forest plant species occurring in both the herbaceous (\(\le 1\)-m height) and tree (\(> 1\)-m height) vegetation layers of a given hedgerow (n = 99)

What are the main determinants of the number of herbaceous forest plant species we can find in hedgerows?

First of all, we computed the pairwise correlation matrix among all quantitative variables to detect variables that are too much correlated among each others and to assess which variables are positively and negatively correlated with species richness in the herbecous layer: our main response variable.

#
carac %>%
  select(c(id_haie,
           annee_med, 
           longueur, 
           largeur, 
           h_moy, 
           h_amp, 
           dist_for, 
           surf_for_500, 
           ipa_adj,
           connection,
           n_tree,
           n_herb)) %>%
  rename(birth_yr=annee_med, 
         tot_length=longueur, 
         max_width=largeur, 
         mean_height=h_moy, 
         range_height=h_amp, 
         for_dist=dist_for, 
         for_cov=surf_for_500, 
         adj_lui=ipa_adj, 
         for_con=connection) %>%
  mutate(for_con=factor(for_con)) -> richness
#
richness %>%
  select(-c(id_haie, 
            for_con)) %>%
  as.matrix() %>%
  rcorr(type="spearman") -> tab_cor
#
rval_cor <- as.data.frame(tab_cor$r)
pval_cor <- as.data.frame(tab_cor$P)
#
#
corrplot(tab_cor$r, 
         method="pie", 
         type="upper", 
         order="hclust", 
         p.mat=tab_cor$P, 
         sig.level=0.001/45,
         insig="label_sig")

Figure 3: Correlation matrix of the variables used to analyse the main determinants of forest plant species richness in the herbaceous layer of hedgerows (n_herb). Potential predictor variables are: maximum width of the hedgerow (max_width); average height of the hedgerow (mean_height); height range of the focal hedgerow (range_height); cumulated amount of forest cover within a buffer area of 500 m around the hedgerow (for_cov); total number of tree and shrub species occurring in the hedgerow overstorey (n_tree); total length of the hedgerow (tot_length); distance to the closest woodland (for_dist); year of first appearance of a hedgerow in the historical archives as a proxy for hedgerow age (birth_yr); and adjacentland-use intensity (adj_lui). Asterisks shows significant (P \(<\) 0.001) correlation values after accounting for multiple pairwise comparisons (10 × 9 / 2 = 45) using a Bonferroni correction (P / 45)

List of candidate models

Here is a list of candidate models we tested to explain species richness in the herbaceous layer of hedgerows. We used generalised linear models (GLMs) with a Poisson family as the response variable is a count data (i.e. species richness in the herbaceous layer of hedgerows).

#
M_rich_1 <- glm(n_herb ~ for_con + 
                         birth_yr +
                         tot_length +
                         max_width +
                         mean_height +
                         for_cov +
                         adj_lui +
                         n_tree +
                         birth_yr:tot_length,
                family=poisson, 
                data=richness)
#
M_rich_2 <- glm(n_herb ~ for_con + 
                         birth_yr +
                         tot_length +
                         max_width +
                         mean_height +
                         for_cov +
                         adj_lui +
                         n_tree +
                         birth_yr:max_width,
                family=poisson, 
                data=richness)
#
M_rich_3 <- glm(n_herb ~ for_con + 
                         birth_yr +
                         tot_length +
                         max_width +
                         mean_height +
                         for_cov +
                         adj_lui +
                         n_tree +
                         birth_yr:mean_height,
                family=poisson, 
                data=richness)
#
M_rich_4 <- glm(n_herb ~ for_con + 
                         birth_yr +
                         tot_length +
                         max_width +
                         mean_height +
                         for_cov +
                         adj_lui +
                         n_tree +
                         birth_yr:for_cov,
                family=poisson, 
                data=richness)
#
M_rich_5 <- glm(n_herb ~ for_con + 
                         birth_yr +
                         tot_length +
                         max_width +
                         mean_height +
                         for_cov +
                         adj_lui +
                         n_tree +
                         birth_yr:for_con,
                family=poisson, 
                data=richness)
#

Best model

We computed the Aikaike information criteria (AIC) to select the best model with the lowest AIC value.

#
AICs <- c(AIC(M_rich_1), 
          AIC(M_rich_2), 
          AIC(M_rich_3), 
          AIC(M_rich_4), 
          AIC(M_rich_5))
#
AICs <- as.data.frame(AICs)
#
names(AICs) <- c("aic_val")
#
AICs$mod_id <- c("M_rich_1", 
                 "M_rich_2", 
                 "M_rich_3", 
                 "M_rich_4", 
                 "M_rich_5")
#
AICs$delta_aic <- AICs$aic_val-min(AICs$aic_val)
#
Table 4: List of candidate models and their Aikaike information criteria (AIC) values
aic_val mod_id delta_aic
666.3170 M_rich_1 0.00000
679.1921 M_rich_2 12.87508
683.1957 M_rich_3 16.87873
683.2423 M_rich_4 16.92532
679.9825 M_rich_5 13.66548

Outputs from the best model

Once the best model was selected, we reran it with rescaled (using the gscale function from the jtools package) predictor variables to be able to compare coefficient estimates as effect sizes.

#
best_M_rich_scaled <- glm(n_herb ~ for_con + 
                                   gscale(birth_yr) +
                                   gscale(tot_length) +
                                   gscale(max_width) +
                                   gscale(mean_height) +
                                   gscale(for_cov) +
                                   gscale(adj_lui) +
                                   gscale(n_tree) +
                                   gscale(birth_yr):gscale(tot_length),
                          family=poisson, 
                          data=richness)
#
table_rich <- round(summary(best_M_rich_scaled)$coeff, digits=5)
table_rich <- as.data.frame(table_rich)
names(table_rich) <- c("Coefficient", "Standard error", "Z-value", "P-value")
row.names(table_rich)[1] <- "Intercept"
row.names(table_rich)[2] <- "Attached"
row.names(table_rich)[3] <- "Year"
row.names(table_rich)[4] <- "Length"
row.names(table_rich)[5] <- "Width"
row.names(table_rich)[6] <- "Height"
row.names(table_rich)[7] <- "Forest cover"
row.names(table_rich)[8] <- "Land-use"
row.names(table_rich)[9] <- "N tree"
row.names(table_rich)[10] <- "Year:Length"
#
Table 5: Best candidate model of the main determinants of the number of herbaceous forest species found in hedgerows. All continuous variables were z-transformed using the gscale function from the jtools package in R so that coefficient estimates can be interpreted as effect sizes. We used a generalized linear model with a logit link (family = Poisson). The intercept gives the effect size with all other continuous variables set to zero and the connectivity factor variable set to the non-connected level
Coefficient Standard error Z-value P-value
Intercept 2.69533 0.03091 87.19292 0.00000
Attached 0.56159 0.08651 6.49191 0.00000
Year -0.32865 0.06050 -5.43192 0.00000
Length 0.12376 0.06741 1.83598 0.06636
Width 0.12799 0.05325 2.40339 0.01624
Height 0.16056 0.05237 3.06576 0.00217
Forest cover 0.02524 0.06665 0.37868 0.70493
Land-use -0.30746 0.04965 -6.19260 0.00000
N tree 0.13676 0.05581 2.45072 0.01426
Year:Length -0.47051 0.11674 -4.03029 0.00006

Species-time-area relationship (STAR)

#
best_M_rich <- M_rich_1
#
ilink <- family(best_M_rich)$linkinv
#
pred <- as_tibble(expand.grid(birth_yr=seq(1720, 2020, 10), 
                              tot_length=seq(10, 500, 10), 
                              for_con=c(0, 1)))
#
pred %>%
  mutate(for_con=factor(for_con),
         adj_lui=rep(median(richness$adj_lui)),
         max_width=rep(median(richness$max_width)),
         mean_height=rep(median(richness$mean_height)),
         n_tree=rep(median(richness$n_tree)),
         for_cov=rep(median(richness$for_cov))) -> pred
#         
pred %>% 
  mutate(fit=predict(best_M_rich, pred, type="link", se.fit=TRUE)[[1]],
         se_fit=predict(best_M_rich, pred, type="link", se.fit=TRUE)[[2]],
         n_herb_fit=ilink(fit),
         ci_upper=ilink(fit+(2*se_fit)),
         ci_lower=ilink(fit-(2*se_fit))) -> pred
#
pred %>%
  filter(for_con==0) -> con_no
#
pred %>%
  filter(for_con==1) -> con_yes
#
plot_inte_a <- ggplot(con_no, 
                      aes(x=birth_yr, 
                          y=n_herb_fit)) +
               geom_line(aes(colour=tot_length, 
                             group=tot_length)) +
               scale_color_viridis(begin=1, 
                                   end=0, 
                                   name="Length\n(m)",
                                   breaks=seq(100, 500, 100),
                                   labels=c("  100", "  200", "  300", "  400", "  500")) +
               scale_y_continuous(limits=c(0, 100), 
                                  breaks=seq(0, 100, 20)) +
               labs(x="Year of hedgerow\nestablishment", 
                    y="Species richness", 
                    tag="(a)") +
               ggtitle("Not attached") +
               fig
#
plot_inte_b <- ggplot(con_yes, 
                       aes(x=birth_yr, 
                           y=n_herb_fit)) +
                geom_line(aes(colour=tot_length, 
                              group=tot_length)) +
                scale_color_viridis(begin=1, 
                                    end=0, 
                                    name="Length\n(m)") +
                scale_y_continuous(limits=c(0, 100), 
                                   breaks=seq(0, 100, 20)) +
                labs(x="Year of hedgerow\nestablishment", 
                     y="Species richness", 
                     tag="(b)") +
                ggtitle("Attached to a woodland") +
                fig
#
plot_inte_c <- ggplot(con_no, 
                      aes(x=tot_length, 
                          y=n_herb_fit)) +
               geom_line(aes(colour=birth_yr, 
                             group=birth_yr)) +
               scale_color_viridis(option="magma", 
                                   name="Year") +
               scale_y_continuous(limits=c(0, 100), 
                                  breaks=seq(0, 100, 20)) +
               labs(x="Total length of\nhedgerow (m)", 
                    y="Species richness", 
                    tag="(c)") +
               ggtitle("Not attached") +
               fig
#
plot_inte_d <- ggplot(con_yes, 
                       aes(x=tot_length, 
                           y=n_herb_fit)) +
                geom_line(aes(colour=birth_yr, 
                              group=birth_yr)) +
                scale_color_viridis(option="magma", 
                                    name="Year") +
                scale_y_continuous(limits=c(0, 100), 
                                   breaks=seq(0, 100, 20)) +
                labs(x="Total length of\nhedgerow (m)", 
                     y="Species richness", 
                     tag="(d)") +
                ggtitle("Attached to a woodland") +
                fig
#
get_legend<-function(myggplot){
     tmp <- ggplot_gtable(ggplot_build(myggplot))
     leg <- which(sapply(tmp$grobs, function(x) x$name)=="guide-box")
     legend <- tmp$grobs[[leg]]
     return(legend)
     }
#
legend_ab <- get_legend(plot_inte_a)
#
legend_cd <- get_legend(plot_inte_c)
#
plot_inte_a <- plot_inte_a + 
  theme(legend.position="none")
#
plot_inte_b <- plot_inte_b + 
  theme(legend.position="none")
#
plot_inte_c <- plot_inte_c + 
  theme(legend.position="none")
#
plot_inte_d <- plot_inte_d + 
  theme(legend.position="none")
#
grid.arrange(plot_inte_a, 
             plot_inte_b, 
             legend_ab, 
             plot_inte_c, 
             plot_inte_d, 
             legend_cd, 
             ncol=3,
             nrow=2, 
             widths=c(8, 8, 2), 
             heights=c(8, 8))

Figure 4: Space-time interaction between the year of first appearance of a hedgerow in the historical archives (a, b), as a proxy for hedgerow age, and the total length of the hedgerow (c, d) on the total number of forest plant species occurring in the understorey of hedgerows not attached (a, c) or attached (b, d) to a woodland. Predictions are based on the best candidate model (see Table 5 for standardized coefficients). All other covariates not displayed on a given plot are set to their median values

How are specialist and generalist species distributed along hedgerows and away from woodlands?

From the initial dataset, we selected only the 29 hedgerows with information on species presence-absence within each of the consecutive 5-m segments along the hedgerow. Similarly, we extracted only the species composition matrix of these 29 hedgerows from the initial species-by-hedgerow matrix.

#
carac %>%
  filter(multi_bande==1) %>%
  select(c(id_haie, 
           annee_med, 
           connection)) %>%
  rename(birth_yr=annee_med, 
         for_con=connection) %>%
  mutate(for_con=factor(for_con)) -> carac_hedgerows_with_bands
#
flo %>%
  filter(multi_bande==1) %>%
  left_join(carac_hedgerows_with_bands, by="id_haie") %>%
  select(c(id_haie, 
           id_bande, 
           num_bande,
           dist_bande,
           birth_yr,
           for_con, 
           starts_with("H_"))) %>%
  rename(dist_start=num_bande,
         dist_for=dist_bande) %>%
  mutate(dist_start=dist_start*5-2.5) %>%
  gather(starts_with("H_"), key="species_id", value="p_a") %>%
  mutate_if(is.character, str_replace_all, pattern="H_", replacement="") %>% 
  mutate_if(is.character, str_replace_all, pattern="_\\.", replacement=" ") %>% 
  mutate_if(is.character, str_remove_all, pattern=" .+") %>% 
  mutate_if(is.character, str_replace_all, pattern="([a-z])\\.([a-z])", replacement="\\1-\\2") %>% 
  mutate_if(is.character, str_replace_all, pattern="_", replacement=" ") %>% 
  arrange(id_haie, id_bande, dist_for, species_id) -> flo_by_band
#

For model calibration, we focused on the list of species (specialists and generalists) that are occurring more than 25 times across all 5-m segements that we surveyed. Here is the code to select those species.

flo_by_band %>%
  filter(p_a==1) %>%
  group_by(species_id) %>% 
  summarise(tot_occ=n()) %>% 
  arrange(desc(tot_occ)) -> occ_by_sp
#
flo_by_band %>%
  left_join(occ_by_sp, by="species_id") -> flo_by_band
#
for_stat <- read.table("species_list.txt", header=TRUE, sep="\t")
for_stat <- as_tibble(for_stat)
#
for_stat %>% 
  select(-c(life_form,
            below_1m,
            above_1m)) %>% 
  rename(species_id=sp_name) %>% 
  mutate(species_id=as.character(species_id)) -> for_stat
#
flo_by_band %>%
  left_join(for_stat, by="species_id") -> flo_by_band
#
flo_by_band %>%
  filter(for_stat %in% c("specialist", "generalist")) %>% 
  filter(tot_occ>25) -> flo_by_band
#

Modelling species distribution

Here is the model we ran to test whether species response curves along hedgerows and away from woodlands are differing between forest specialists and generalists. We used generalised linear mixed-effets models (GLMMs) with a Binomial family as the response variable is binary (i.e. species presence-absence). The list of fixed-effect variables is provided below. We used species name and the hedgerow id as two random intercept terms in our GLMM.

#
best_M_distrib <- glmer(p_a ~ dist_for + 
                              birth_yr +
                              for_stat +
                              for_con + 
                              birth_yr:for_stat +
                              dist_for:for_stat +
                              dist_for:for_con +
                              (1|id_haie) +
                              (1|species_id), 
                        family=binomial,
                        data=flo_by_band)
#
best_M_distrib_scaled <- glmer(p_a ~ gscale(dist_for) + 
                                     gscale(birth_yr) +
                                     for_stat +
                                     for_con + 
                                     gscale(birth_yr):for_stat +
                                     gscale(dist_for):for_stat +
                                     gscale(dist_for):for_con +
                                     (1|id_haie) +
                                     (1|species_id), 
                               family=binomial,
                               data=flo_by_band)
#
table_distrib <- round(summary(best_M_distrib_scaled)$coeff, digits=5)
table_distrib <- as.data.frame(table_distrib)
names(table_distrib) <- c("Coefficient", "Standard error", "Z-value", "P-value")
row.names(table_distrib)[1] <- "Intercept"
row.names(table_distrib)[2] <- "Distance"
row.names(table_distrib)[3] <- "Year"
row.names(table_distrib)[4] <- "Specialists"
row.names(table_distrib)[5] <- "Attached"
row.names(table_distrib)[6] <- "Year:Specialists"
row.names(table_distrib)[7] <- "Distance:Specialists"
row.names(table_distrib)[8] <- "Distance:Attached"
#
Table 6: Best candidate model of the main determinants of the distribution of herbaceous forest species along hedgerows. All continuous variables were z-transformed using the gscale function from the jtools package in R so that coefficient estimates can be interpreted as effect sizes. We used a generalized linear mixed-effects model with a logit link (family = Binomial). The intercept gives the effect size with all other continuous variables set to zero and the forest status factor variable set to the generalists level and the connectivity factor variable set to the non-connected level
Coefficient Standard error Z-value P-value
Intercept -2.60071 0.28871 -9.00815 0.00000
Distance 0.67105 0.24165 2.77698 0.00549
Year -1.43337 0.22218 -6.45145 0.00000
Specialists -0.53494 0.40198 -1.33076 0.18327
Attached 0.57935 0.28568 2.02798 0.04256
Year:Specialists 0.34539 0.13728 2.51599 0.01187
Distance:Specialists -0.77148 0.17628 -4.37653 0.00001
Distance:Attached -2.21396 0.73202 -3.02446 0.00249

Distribution profile of forest specialists and generalists along hedgerows

Here is the code to predict the effect of distance to the closest woodland on the probability of occurrence of both forest specialists and generalists along a given hedgerow, either attached or not (isolated) to a woodland.

#
ilink <- family(best_M_distrib)$linkinv
#
pred <- as_tibble(expand.grid(dist_for=seq(0, 1800, 1), 
                              birth_yr=seq(1500, 2020, 10), 
                              for_stat=c("generalist", "specialist"), 
                              for_con=c(0, 1)))
#
pred %>%
  mutate(for_stat=factor(for_stat)) %>% 
  mutate(for_con=factor(for_con)) -> pred
#         
pred %>% 
  mutate(fit=predict(best_M_distrib, pred, re.form=NA, type="link", se.fit=TRUE)[[1]],
         se_fit=predict(best_M_distrib, pred, re.form=NA, type="link", se.fit=TRUE)[[2]],
         resp_fit=ilink(fit),
         ci_upper=ilink(fit+(2*se_fit)),
         ci_lower=ilink(fit-(2*se_fit))) -> pred
#
pred %>%
  filter(birth_yr==1700, 
         for_stat=="specialist", 
         for_con==1,
         dist_for<=170) -> pred_old_specialist_con
#
pred %>%
  filter(birth_yr==2000, 
         for_stat=="specialist", 
         for_con==1,
         dist_for<=170) -> pred_new_specialist_con
#
pred %>%
  filter(birth_yr==1700, 
         for_stat=="generalist", 
         for_con==1,
         dist_for<=170) -> pred_old_generalist_con
#
pred %>%
  filter(birth_yr==2000, 
         for_stat=="generalist", 
         for_con==1,
         dist_for<=170) -> pred_new_generalist_con
#
pred %>%
  filter(birth_yr==1700, 
         for_stat=="specialist", 
         for_con==0,
         dist_for<=1800) -> pred_old_specialist_discon
#
pred %>%
  filter(birth_yr==2000, 
         for_stat=="specialist", 
         for_con==0,
         dist_for<=1800) -> pred_new_specialist_discon
#
pred %>%
  filter(birth_yr==1700, 
         for_stat=="generalist", 
         for_con==0,
         dist_for<=1800) -> pred_old_generalist_discon
#
pred %>%
  filter(birth_yr==2000, 
         for_stat=="generalist", 
         for_con==0,
         dist_for<=1800) -> pred_new_generalist_discon
#
flo_by_band %>% 
  filter(for_con==1,
         birth_yr<=median(.$birth_yr)) -> flo_by_band_old_con
#
flo_by_band %>% 
  filter(for_con==1,
         birth_yr>median(.$birth_yr)) -> flo_by_band_new_con
#
flo_by_band %>% 
  filter(for_con==0,
         birth_yr<=median(.$birth_yr)) -> flo_by_band_old_discon
#
flo_by_band %>% 
  filter(for_con==0,
         birth_yr>median(.$birth_yr)) -> flo_by_band_new_discon
#
plot_dist_a <- ggplot(flo_by_band_old_con, 
                      aes(x=dist_for, 
                          y=p_a, 
                          colour=for_stat)) +
               geom_point(size=3, 
                          alpha=0.005) +
               geom_line(data=pred_old_specialist_con, 
                         aes(x=dist_for, 
                             y=resp_fit), 
                         colour="green4",
                         linetype="solid",
                         size=1) +
               geom_ribbon(data=pred_old_specialist_con, 
                           aes(x=dist_for, 
                               y=resp_fit, 
                               ymin=ci_lower, 
                               ymax=ci_upper), 
                           fill="green4", 
                           colour=NA, 
                           alpha=0.25) +
               geom_line(data=pred_old_generalist_con, 
                         aes(x=dist_for, y=resp_fit), 
                         linetype="solid", 
                         colour="purple4", 
                         size=1) +
               geom_ribbon(data=pred_old_generalist_con, 
                           aes(x=dist_for, 
                               y=resp_fit, 
                               ymin=ci_lower, 
                               ymax=ci_upper), 
                           fill="purple4", 
                           colour=NA, 
                           alpha=0.25) +
               scale_colour_manual(values=c("purple4", "green4"), 
                                   name="Species\ntype", 
                                   labels=c("Generalists", "Specialists")) +
               scale_fill_manual(values=c("purple4", "green4"), 
                                 guide=FALSE) + 
               guides(color=guide_legend(override.aes=list(size=8, 
                                                           shape=19, 
                                                           alpha=0.25))) +
               labs(x="Distance to woodland (m)", 
                    y="Probability of occurrence",
                    tag="(a)") +
               ggtitle("Attached and ancient") +
               fig
#
plot_dist_b <- ggplot(flo_by_band_new_con, 
                      aes(x=dist_for, 
                          y=p_a, 
                          colour=for_stat)) +
               geom_point(size=3, 
                          alpha=0.005) +
               geom_line(data=pred_new_specialist_con, 
                         aes(x=dist_for, 
                             y=resp_fit), 
                         colour="green4",
                         linetype="solid",
                         size=1) +
               geom_ribbon(data=pred_new_specialist_con, 
                           aes(x=dist_for, 
                               y=resp_fit, 
                               ymin=ci_lower, 
                               ymax=ci_upper), 
                           fill="green4", 
                           colour=NA, 
                           alpha=0.25) +
               geom_line(data=pred_new_generalist_con, 
                         aes(x=dist_for, y=resp_fit), 
                         linetype="solid", 
                         colour="purple4", 
                         size=1) +
               geom_ribbon(data=pred_new_generalist_con, 
                           aes(x=dist_for, 
                               y=resp_fit, 
                               ymin=ci_lower, 
                               ymax=ci_upper), 
                           fill="purple4", 
                           colour=NA, 
                           alpha=0.25) +
               scale_colour_manual(values=c("purple4", "green4"), 
                                   name="Forest herbs", 
                                   labels=c("Generalists", "Specialists")) +
               scale_fill_manual(values=c("purple4", "green4"), 
                                 guide=FALSE) + 
               guides(color=guide_legend(override.aes=list(size=3, 
                                                           shape=19, 
                                                           alpha=0.25))) +
               labs(x="Distance to woodland (m)", 
                    y="Probability of occurrence",
                    tag="(b)") +
               ggtitle("Attached and recent") +
               fig
#
plot_dist_c <- ggplot(flo_by_band_old_discon, 
                      aes(x=dist_for, 
                          y=p_a, 
                          colour=for_stat)) +
               geom_point(size=3, 
                          alpha=0.005) +
               geom_line(data=pred_old_generalist_discon, 
                         aes(x=dist_for, 
                             y=resp_fit),
                         colour="purple4",
                         linetype="solid",
                         size=1) +
               geom_ribbon(data=pred_old_generalist_discon, 
                           aes(x=dist_for, 
                               y=resp_fit, 
                               ymin=ci_lower, 
                               ymax=ci_upper), 
                           fill="purple4", 
                           colour=NA, 
                           alpha=0.25) +
               geom_line(data=pred_old_specialist_discon, 
                         aes(x=dist_for, 
                             y=resp_fit), 
                         linetype="solid", 
                         colour="green4", 
                         size=1) +
               geom_ribbon(data=pred_old_specialist_discon, 
                           aes(x=dist_for, 
                               y=resp_fit, 
                               ymin=ci_lower, 
                               ymax=ci_upper), 
                           fill="green4", 
                           colour=NA, 
                           alpha=0.25) +
               scale_colour_manual(values=c("purple4", "green4"), 
                                   name="Species affinity\nto closed forest", 
                                   labels=c("Generalists", "Specialists")) +
               scale_fill_manual(values=c("purple4", "green4"), 
                                 guide=FALSE) + 
               guides(color=guide_legend(override.aes=list(size=8, 
                                                           shape=19, 
                                                           alpha=0.25))) +
               labs(x="Distance to woodland (m)", 
                    y="Probability of occurrence",
                    tag="(c)") +
               ggtitle("Isolated and ancient") +
               fig
#
plot_dist_d <- ggplot(flo_by_band_new_discon, 
                      aes(x=dist_for, 
                          y=p_a, 
                          colour=for_stat)) +
               geom_point(size=3, 
                          alpha=0.005) +
               geom_line(data=pred_new_generalist_discon, 
                         aes(x=dist_for, 
                             y=resp_fit),
                         colour="purple4",
                         linetype="solid",
                         size=1) +
               geom_ribbon(data=pred_new_generalist_discon, 
                           aes(x=dist_for, 
                               y=resp_fit, 
                               ymin=ci_lower, 
                               ymax=ci_upper), 
                           fill="purple4", 
                           colour=NA, 
                           alpha=0.25) +
               geom_line(data=pred_new_specialist_discon, 
                         aes(x=dist_for, 
                             y=resp_fit), 
                         linetype="solid", 
                         colour="green4", 
                         size=1) +
               geom_ribbon(data=pred_new_specialist_discon, 
                           aes(x=dist_for, 
                               y=resp_fit, 
                               ymin=ci_lower, 
                               ymax=ci_upper), 
                           fill="green4", 
                           colour=NA, 
                           alpha=0.25) +
               scale_colour_manual(values=c("purple4", "green4"), 
                                   name="Species affinity\nto closed forest", 
                                   labels=c("Generalists", "Specialists")) +
               scale_fill_manual(values=c("purple4", "green4"), 
                                 guide=FALSE) + 
               guides(color=guide_legend(override.aes=list(size=8, 
                                                           shape=19, 
                                                           alpha=0.25))) +
               labs(x="Distance to woodland (m)", 
                    y="Probability of occurrence",
                    tag="(d)") +
               ggtitle("Isolated and recent") +
               fig
#
plot_dist_a <- plot_dist_a +
               theme(legend.position="none")
#
plot_dist_b <- plot_dist_b +
               theme(legend.position=c(1, 0.9))
#
plot_dist_c <- plot_dist_c +
               theme(legend.position="none")
#
plot_dist_d <- plot_dist_d +
               theme(legend.position="none")
#
grid.arrange(plot_dist_a,
             plot_dist_b, 
             plot_dist_c, 
             plot_dist_d, 
             nrow=2, 
             ncol=2, 
             widths=c(8, 8), 
             heights=c(8, 8))

Figure 5: Effect of distance to woodland (m) on the distribution of both forest specialists and generalists along hedgerows that are either attached (a, b) or not (isolated) (c, d) to a woodland. For illustrative purposes, we set the year of first appearance of the hedgerow in the historical archives to 1700 (a, c) and 2000 (b, d) for ancient and recent hedgerows, respectively.

Distribution profile of two forest specialist species

Here is the code to run species-specific GLMMs.

#
flo_by_band %>%
  group_by(species_id) %>% 
  nest() -> flo_by_band_nested
#
model_sp <- function(df) {
  glmer(p_a ~ dist_for + 
              birth_yr +
              for_con +
              dist_for:for_con +
              (1|id_haie), 
        family=binomial,
        data=df)
}
#

Predictions for Stellaria holostea

#
sp_i <- flo_by_band_nested$data[[25]]
#
ilink <- family(model_sp(sp_i))$linkinv
#
pred <- as_tibble(expand.grid(dist_for=seq(0, 1800, 1), birth_yr=seq(1700, 2020, 10), for_con=c(0, 1)))
#
pred %>%
  mutate(for_con=factor(for_con)) -> pred
#         
pred %>% 
  mutate(fit=predict(model_sp(sp_i), pred, re.form=NA, type="link", se.fit=TRUE)[[1]],
         se_fit=predict(model_sp(sp_i), pred, re.form=NA, type="link", se.fit=TRUE)[[2]],
         resp_fit=ilink(fit),
         ci_upper=ilink(fit+(2*se_fit)),
         ci_lower=ilink(fit-(2*se_fit))) -> pred
#
pred %>%
  filter(birth_yr==1700, 
         for_con==1, 
         dist_for<=170) %>% 
         mutate(hedge_age=ifelse(birth_yr<=median(sp_i$birth_yr), "Ancient", "Recent")) %>% 
         mutate(hedge_age=factor(hedge_age)) -> old_con
#
pred %>%
  filter(birth_yr==2000, 
         for_con==1,
         dist_for<=170) %>% 
         mutate(hedge_age=ifelse(birth_yr<=median(sp_i$birth_yr), "Ancient", "Recent")) %>% 
         mutate(hedge_age=factor(hedge_age))-> new_con
#
sp_i %>% 
  filter(for_con==1) %>% 
  mutate(hedge_age=ifelse(birth_yr<=median(.$birth_yr), "Ancient", "Recent")) %>% 
  mutate(hedge_age=factor(hedge_age)) -> sp_i
#
plot_ste_hol <- ggplot(sp_i, 
                       aes(x=dist_for, 
                           y=p_a,
                           colour=hedge_age,
                           group=hedge_age)) + 
                geom_point(size=3, 
                           alpha=0.25) +
                geom_line(data=old_con, 
                          aes(x=dist_for, 
                              y=resp_fit), 
                          colour="green4", 
                          size=0.5) +
                geom_ribbon(data=old_con, 
                            aes(x=dist_for, 
                                y=resp_fit, 
                                ymin=ci_lower, 
                                ymax=ci_upper), 
                            fill="green4",
                            colour=NA,
                            alpha=0.25) +
                geom_line(data=new_con, 
                          aes(x=dist_for, 
                              y=resp_fit), 
                          colour="purple4", 
                          size=0.5) +
                geom_ribbon(data=new_con, 
                            aes(x=dist_for, 
                                y=resp_fit, 
                                ymin=ci_lower, 
                                ymax=ci_upper), 
                            fill="purple4",
                            colour=NA,
                            alpha=0.25) +
                scale_colour_manual(values=c("purple4", "green4"), 
                                    name="Hedgerow\nage", 
                                    labels=c("Recent", "Ancient")) +
                scale_fill_manual(values=c("purple4", "green4"), 
                                  guide=FALSE) +
                labs(x="Distance to the \nattached woodland (m)", 
                     y="Stellaria holostea",
                     tag="(b)") +
                fig
#

Predictions for Viola reichenbachiana

#
sp_i <- flo_by_band_nested$data[[30]]
#
ilink <- family(model_sp(sp_i))$linkinv
#
pred <- as_tibble(expand.grid(dist_for=seq(0, 1800, 1), birth_yr=seq(1700, 2020, 10), for_con=c(0, 1)))
#
pred %>%
  mutate(for_con=factor(for_con)) -> pred
#         
pred %>% 
  mutate(fit=predict(model_sp(sp_i), pred, re.form=NA, type="link", se.fit=TRUE)[[1]],
         se_fit=predict(model_sp(sp_i), pred, re.form=NA, type="link", se.fit=TRUE)[[2]],
         resp_fit=ilink(fit),
         ci_upper=ilink(fit+(2*se_fit)),
         ci_lower=ilink(fit-(2*se_fit))) -> pred
#
pred %>%
  filter(birth_yr==1700, 
         for_con==1, 
         dist_for<=170) %>% 
         mutate(hedge_age=ifelse(birth_yr<=median(sp_i$birth_yr), "Ancient", "Recent")) %>% 
         mutate(hedge_age=factor(hedge_age))-> old_con
#
pred %>%
  filter(birth_yr==2000, 
         for_con==1,
         dist_for<=170) %>% 
         mutate(hedge_age=ifelse(birth_yr<=median(sp_i$birth_yr), "Ancient", "Recent")) %>% 
         mutate(hedge_age=factor(hedge_age))-> new_con
#
sp_i %>% 
  filter(for_con==1) %>% 
  mutate(hedge_age=ifelse(birth_yr<=median(.$birth_yr), "Ancient", "Recent")) %>% 
  mutate(hedge_age=factor(hedge_age)) -> sp_i
#
plot_vio_rei <- ggplot(sp_i, 
                       aes(x=dist_for, 
                           y=p_a,
                           colour=hedge_age,
                           group=hedge_age)) + 
                geom_point(size=3, 
                           alpha=0.25) +
                geom_line(data=old_con, 
                          aes(x=dist_for, 
                              y=resp_fit), 
                          colour="green4",
                          size=0.5) +
                geom_ribbon(data=old_con, 
                            aes(x=dist_for, 
                                y=resp_fit, 
                                ymin=ci_lower, 
                                ymax=ci_upper), 
                            fill="green4",
                            colour=NA,
                            alpha=0.25) +
                geom_line(data=new_con, 
                          aes(x=dist_for, 
                              y=resp_fit), 
                          colour="purple4", 
                          size=0.5) +
                geom_ribbon(data=new_con, 
                            aes(x=dist_for, 
                                y=resp_fit, 
                                ymin=ci_lower, 
                                ymax=ci_upper), 
                            fill="purple4",
                            colour=NA,
                            alpha=0.25) +
                scale_colour_manual(values=c("purple4", "green4"), 
                                    name="Hedgerow\nage", 
                                    labels=c("Recent", "Ancient")) +
                scale_fill_manual(values=c("purple4", "green4"), 
                                  guide=FALSE) +
                labs(x="Distance to the \nattached woodland (m)", 
                     y="Viola reichenbachiana",
                     tag="(a)") +
                fig
#

Plotting both species on the same figure

#
plot_ste_hol <- plot_ste_hol +
                theme(legend.position="none",
                      axis.title.y=element_text(face="bold.italic"))
#
plot_vio_rei <- plot_vio_rei +
                theme(axis.title.y=element_text(face="bold.italic"))             
#
grid.arrange(plot_vio_rei, 
             plot_ste_hol, 
             nrow=1, 
             ncol=2, 
             widths=c(8, 8), 
             heights=c(8))

Figure 6: Effect of distance to woodland (m) on the probability of occurrence of (a) Viola reichenbachiana and (b) Stellaria holostea along both recent (year of first appearance of the hedgerow in the historical archives set to 2000) and ancient (year of first appearance of the hedgerow in the historical archives set to 1700) hedgerows that are attached to a woodland