Biodi e si y Da a Jou nal 13: e173529
doi: 10.3897/BDJ.13.e173529
Da a Pape
Coas al g assland ege a ion eco ds om São
Miguel Island (Azo es) and he sou h-wes e n
coas o mainland Po ugal
Hugo Rena o M.G. Calado , An ónio O. Soa es , Ruben Heleno , Paulo A. V. Bo ges
‡ Uni e si y o he Azo es, cE3c – Cen e o Ecology, E olu ion and En i onmen al Changes/Azo ean Biodi e si y G oup,
CHANGE – Global Change and Sus ainabili y Ins i u e, Facul y o Science and Technology, Rua da Mãe de Deus,
9500-321 Pon a Delgada, São Miguel, Azo es, Po ugal
§ Cen e o Func ional Ecology, Associa e Labo a o y TERRA, Depa men o Li e Sciences, Uni e si y o Coimb a, Calçada
Ma im de F ei as, 3000-456, Coimb a, Po ugal
| Uni e si y o Azo es, CE3C—Cen e o Ecology, E olu ion and En i onmen al Changes, Azo ean Biodi e si y G oup,
CHANGE —Global Change and Sus ainabili y Ins i u e, School o Ag icul u al and En i onmen al Sciences, Rua Capi ão
João d’Á ila, Pico da U ze, 9700-042, Ang a do He oísmo, Azo es, Po ugal
¶ IUCN SSC Moni o ing Specialis G oup, Ang a do He oísmo, Azo es, Po ugal
Co esponding au ho : Hugo Rena o M.G. Calado (hugo. [email protected] )
Academic edi o : Rui Elias
Recei ed: 30 Sep 2025 | Accep ed: 27 Oc 2025 | Published: 24 No 2025
Ci a ion: Calado HMG, Soa es AO, Heleno R, Bo ges PAV (2025) Coas al g assland ege a ion eco ds om
São Miguel Island (Azo es) and he sou h-wes e n coas o mainland Po ugal. Biodi e si y Da a Jou nal 13:
e173529. h ps://doi.o g/10.3897/BDJ.13.e173529
Abs ac
Backg ound
The p esen wo k p o ides an in en o y o he plan species eco ded in wo dis inc
coas al g assland ege a ions: he Azo es A chipelago (São Miguel Island) and he
sou h-wes e n coas o mainland Po ugal (Sesimb a and Sines Regions – Se úbal
Dis ic ). Si es we e selec ed in bo h egions o ha e a simila gene al subs a e ( ocky),
la i ude and ele a ion. Thi y-one si es we e selec ed in he coas al g asslands: hi een
we e loca ed on São Miguel Island and eigh een on he mainland, dis ibu ed ac oss
Sesimb a (12) and Sines (6). All si es we e isi ed once du ing he sp ing o 2022. In
each si e, 30 pho os we e aken a 5-me e in e als, o a o al o 930 pho os.
‡ ‡ § |,¶
© Calado H e al. This is an open access a icle dis ibu ed unde he e ms o he C ea i e Commons A ibu ion License (CC BY
4.0), which pe mi s un es ic ed use, dis ibu ion, and ep oduc ion in any medium, p o ided he o iginal au ho and sou ce a e
c edi ed.
The Sesimb a and Sines Regions we e chosen because hey ep esen well-p ese ed
examples o coas al g assland ege a ion on he sou h-wes e n coas o mainland
Po ugal, sha ing simila en i onmen al cha ac e is ics wi h he coas al g asslands o São
Miguel Island, such as ocky subs a e, A lan ic exposu e and compa able la i ude. This
design allows a meaning ul compa ison be ween insula and con inen al ege a ion
unde simila abio ic condi ions, p o iding a s anda dised amewo k o documen ing
coas al plan di e si y ac oss con as ing geog aphic con ex s.
New in o ma ion
Mos eco ds consolida e p e iously documen ed occu ences om na ional eposi o ies,
bu he e hey a e e- alida ed h ough s anda dised, plo -based, geo e e enced su eys
ha ha monise axonomy and me ada a. C ucially, his elease cons i u es he i s
coo dina ed, s anda dised in en o y o coas al g assland ascula plan s spanning he
Azo es and he sou h-wes e n mainland o Po ugal, enabling di ec c oss- egional
compa isons. In addi ion o se e al wi hin-plo dis ibu ional upda es, we documen
mul iple in oduced and in asi e axa, mos no ably Ca pob o us edulis and Oxalis pes-
cap ae, ha a e known o eshape coas al communi ies. As expec ed o A lan ic coas al
g asslands, he lo a is domina ed by As e aceae, Poaceae, Fabaceae and
B assicaceae, p o iding a baseline o long- e m moni o ing and managemen ac oss
bo h a chipelagic and con inen al sho es.
Keywo ds
sampling e en , occu ence, plan s, Azo es, mainland Po ugal, coas al g asslands,
pho og aphic sampling
In oduc ion
G asslands, including coas al a ian s, ha e been widely s udied in e ms o plan and
aunal composi ion (Jones and Donnelly 2004, Fehe e al. 2021). These ecosys ems a e
o pa icula impo ance because hey se e as habi a s o nume ous o ganisms ha
p o ide c ucial ecosys em se ices o humans, such as nu ien cycling, ca bon
seques a ion and pollina ion (Pe e s e al. 2016). Ne e heless, s udies ocusing
speci ically on coas al g assland ege a ion emain compa a i ely sca ce, lea ing much
o be explo ed (Calado e al. 2024).
Vege a ion co e plays a key ole in he s uc u ing o ecosys ems, wi h ce ain o ganisms
being closely linked o he ype o ege a ion p esen , wi hou which hey canno es ablish
hemsel es. Examples include he mona ch bu e ly (Danaus plexippus (Linnaeus,
1758)) and plan s o he genus Asclepias spp. (G eens ein e al. 2022), as well as he
“Lundy cabbage lea bee le” (Psylliodes lu idipennis, Ch ysomelidae), which depends on
he plan Coincya w igh ii, endemic o Lundy Island, England (Comp on e al. 2007). The
2Calado H e al
decline o hese plan popula ions would ce ainly lead o a educ ion o he espec i e
a h opod communi ies.
An h opogenic ac i i ies and clima e change signi ican ly a ec he s uc u e o na u al
ecosys ems (Wick e al. 2016, Lis e and Ga cia 2018, Ba dge e al. 2021, Dai e al.
2022). Oceanic islands a e pa icula ly sensi i e o such p essu es, as hei es ic ed
a ea and isola ion make hem ulne able o ade, ou ism and biological in asions
(Delgado and Rie a 2020, Bo ges e al. 2022, Boiei o e al. 2024, Calado e al. 2025b).
Unde s anding he composi ion o coas al ege a ion is, he e o e, essen ial o assess
how such ac o s in luence ecosys em s uc u e and biodi e si y.
Coas al g assland ege a ion ep esen s a aluable componen o A lan ic ecosys ems,
combining Eu o-A lan ic, Medi e anean and Maca onesian lo is ic elemen s.
Unde s anding i s composi ion ac oss island and mainland con ex s p o ides insigh s
in o biogeog aphical pa e ns, habi a speci ici y and po en ial d i e s o species u no e .
This da a pape aims o documen and make publicly a ailable a s anda dised in en o y
o plan species eco ded in coas al g asslands o São Miguel Island (Azo es) and he
Sesimb a and Sines Regions (Se úbal Dis ic , sou h-wes e n coas o mainland
Po ugal). The da ase cons i u es a baseline o u u e ecological and biogeog aphical
s udies, biodi e si y moni o ing and conse a ion planning.
Gene al desc ip ion
Pu pose:The p ima y objec i e o his s udy is o p o ide a comp ehensi e in en o y o
plan s in he coas al g asslands ac oss São Miguel Island (Azo es A chipelago),
Sesimb a and Sines Regions (Se úbal Dis ic in sou h-wes e n coas o mainland
Po ugal).
Addi ional in o ma ion:The da ase includes in o ma ion on he di e si y and
composi ion o plan s communi ies, eco ded h ough he p ojec 's moni o ing su eys.
P ojec desc ip ion
Ti le:Pheno ypic Plas ici y o Pes and Biological Con ol Agen s: Con as ing Mainland
and Insula Coas al Ecosys ems
Pe sonnel:Paulo A.V. Bo ges, Hugo Calado, Ruben Heleno, An ónio O. Soa es, Lu des
Bo ges.
S udy a ea desc ip ion:The s udy was conduc ed in he coas al g asslands o São
Miguel Island (Azo es A chipelago, No h A lan ic) and sou h-wes e n coas o mainland
Po ugal (Se úbal Dis ic – Sesimb a and Sines Regions) (Fig. 1).
The Azo es A chipelago is loca ed in he cen e o he No h A lan ic, app oxima ely 1,600
km om mainland Po ugal, ex ending abou 600 km be ween San a Ma ia and Co o
Coas al g assland ege a ion eco ds om São Miguel Island (Azo es) and ... 3
(37°–40° N; 25°–31° W). The A chipelago has egula and abundan ain all, an oceanic
clima e wi h ela i ely s able empe a u es, high humidi y h oughou he yea and
pe sis en winds, especially du ing au umn and win e (San os e al. 2004; Bo ges e al.
2019; IPMA 2020).
Mainland Po ugal is loca ed in sou h-wes e n Eu ope (33°–43° N; 9°–6° W), wi h a o al
land a ea o app oxima ely 89,015 km². The Se úbal Dis ic lies on he sou h-wes e n
coas o mainland Po ugal, sou h o Lisbon and encompasses he egions o Sesimb a
and Sines. The mainland has a Medi e anean clima e, cha ac e ised by wa m, d y
summe s and cool, we win e s (Ca alho e al. 2014; de Lima e al. 2015).
Design desc ip ion:Thi y-one plo s we e selec ed, wi h simila gene al subs a e
( ocky), la i ude and ele a ion, in coas al g asslands in bo h ecosys ems: The Azo es
A chipelago and Po ugal mainland.
Funding:H.R.M.G.C. was unded by he Regional FRCT Ph.D. G an M3.1.a/F/012/2021:
Pheno ypic Plas ici y o Pes and Biological Con ol Agen s: Con as ing Mainland and
Insula Coas al Ecosys ems. A.O.S. and P.A.V.B. we e also unded by he p ojec s
Plu iannual Funding FCT-UIDB/00329/2020-2024 - DOI 10.54499/UIDB/00329/2020
(Thema ic Line 1 – in eg a ed ecological assessmen o en i onmen al change on
biodi e si y), Azo es DRCT Plu iannual Funding (M1.1.A/FUNC.UI&D/010/2021-2024)
and PAVB by he p ojec AZORESBIOPORTAL – PORBIOTA (ACORES-01-0145
FEDER-000072). and by he Regional Di ec o a e o Science, Inno a ion and
Figu e 1.
Sampling a eas wi h he plo s indica ed: a Azo es A chipelago and Po ugal mainland; b São
Miguel’s Island; c Se úbal Dis ic ; (i) Sesimb a; (ii) Sines (sou ce: Calado e al. (2024)).
4Calado H e al
De elopmen [Regional Go e nmen o he Azo es] h ough he PROSCIENTIA Incen i e
Sys em M1.1.A/FUNC.UI&D/021/2025 [UI&D/GBA/2025].
Sampling me hods
Desc ip ion:The s udy was conduc ed in he coas al g asslands o São Miguel Island
(Azo es A chipelago, No h A lan ic) and sou h-wes e n coas o mainland Po ugal
(Se úbal Dis ic – Sesimb a and Sines Regions).
Sampling desc ip ion:All 31 si es we e isi ed once du ing he sp ing o 2022 (Ma ch–
May), esul ing in 31 sampling e en s. A each si e, a 45-minu e su ey was ca ied ou o
eco d ege a ion composi ion: 30 minu es we e dedica ed o pho og aphic sampling and
15 minu es o di ec isual obse a ion and p elimina y species iden i ica ion. Each si e
co e ed an a ea o app oxima ely 2500 m², de ined o me hodological consis ency a he
han by minimum a ea calcula ion, o ensu e adequa e spa ial co e age and logis ical
easibili y ac oss all sampling loca ions.
Thi y pho og aphs we e aken pe si e, a 5-me e in e als and 1.2 me es abo e g ound
le el, o alling 930 pho og aphs. Pho og aphs we e aken using a Sony HD Mo ie 720p
came a. No ege a ion co e -abundance scale (e.g. B aun–Blanque ) was applied, as
he s udy ocused exclusi ely on documen ing species occu ences a he han
quan i a i e ege a ion analysis.
Quali y con ol:All pho og aphs we e la e examined indi idually on a compu e o
e i y and con i m species iden i ica ions (Fig. 2). Pho og aphs we e hen o ganised in o
a digi al a chi e o acili a e axonomic alida ion.
When necessa y, ield obse a ions and pho og aphs we e c oss-checked agains online
esou ces, such as he Azo ean Biodi e si y Po al and he Flo a-On da abase, o ensu e
axonomic consis ency.
S ep desc ip ion:In he labo a o y, all pho og aphs we e o ganised and used o
axonomic iden i ica ion. Plan axa we e iden i ied o he species le el whene e
possible, using he Azo ean Biodi e si y Po al and he Flo a-On da abase as e e ences.
When ep oduc i e s uc u es we e absen , mo phologically dis inc plan en i ies we e
classi ied as mo phospecies, ollowing he same ope a ional c i e ia applied in p e ious
a h opod s udies (Calado e al. 2024). Each mo phospecies ep esen s a unique
mo phological uni and is ea ed as an Ope a ional Taxonomic Uni (OTU).
Pho og aphic da a we e ob ained using a Sony HD Mo ie 720p came a, wi h 30
pho og aphs aken pe si e a 5-me e in e als and 1.2 me es abo e g ound le el,
o alling 930 images. This equipmen p o ided su icien image esolu ion o accu a e
pos - ield iden i ica ion o plan axa and mo phospecies.
Coas al g assland ege a ion eco ds om São Miguel Island (Azo es) and ... 5
Geog aphic co e age
Desc ip ion:Gene al spa ial co e age: The s udy was conduc ed in he coas al
g asslands o São Miguel Island (Azo es A chipelago, No h A lan ic) and mainland
Po ugal (Se úbal Dis ic – Sesimb a and Sines Regions).
Coo dina es:37°43'7''N and 38°27'43''N La i ude; 25°51'12''W and 8°47'36''W Longi ude.
Taxonomic co e age
Desc ip ion: Taxonomic anks
Kingdom: Plan ae
Phylum: Magnoliophy a, Pinophy a, P e idophy a.
Class: Genopsida, Liliopsida, Magnoliopsida, Pinopsida, Polypodiopsida.
O de : Alisma ales, Apiales, Aspa agales, As e ales, Bo aginales, B assicales,
Ca yophyllales, Dipsacales, Ephed ales, E icales, Fabales, Fagales, Gen ianales,
Ge aniales, Lamiales, Malpighiales, Mal ales, My ales, Oxalidales, Pinales, Poales,
Polypodiales, P o eales, Ranunculales, Rosales, Sapindales, Saxi agales, Solanales,
Vi ales, Zingibe ales.
Figu e 2.
Examples o g ound pho os om di e en plo s in he Azo es (A) and mainland Po ugal (B),
showing some o he plan species and mo phospecies: 1 – Dimo pho heca u icosa; 2 –
T i olium sp.; 3 – Ca pob o us edulis; 4 – Oxalis pes-cap ae.
6Calado H e al
Family: Acan haceae, Aga aceae, Aizoaceae, Ama an haceae, Ama yllidaceae,
Anaca diaceae, Apiaceae, Apocynaceae, A aceae, Aspa agaceae, Asphodelaceae,
Aspleniaceae, As e aceae, Bo aginaceae, B assicaceae, Cac aceae, Campanulaceae,
Ca yophyllaceae, C assulaceae, Cup essaceae, Denns aed iaceae, D yop e idaceae,
Ephed aceae, E icaceae, Eupho biaceae, Fabaceae, Fagaceae, Ge aniaceae,
Juncaceae, Lamiaceae, Linaceae, Mal aceae, My aceae, Oleaceae, Onag aceae,
O chidaceae, O obanchaceae, Oxalidaceae, Papa e aceae, Pinaceae, Pi ospo aceae,
Plan aginaceae, Plumbaginaceae, Poaceae, Polygonaceae, P imulaceae, P o eaceae,
Rhamnaceae, Rosaceae, Rubiaceae, Sc ophula iaceae, Solanaceae, Tama icaceae,
Thymelaeaceae, T opaeolaceae, U icaceae, Vale ianaceae, Ve benaceae, Vi aceae,
Zingibe aceae.
Tempo al co e age
No es:3 Ma ch 2022 – 17 May 2022.
Collec ion da a
Collec ion name:Rena o_PhD
Collec ion iden i ie :PHEPLA
Specimen p ese a ion me hod:Pho o Reco d
Usage licence
Usage licence:C ea i e Commons Public Domain Wai e (CC-Ze o)
Da a esou ces
Da a package i le:Coas al G assland Vege a ion Reco ds om São Miguel Island
(Azo es) and Mainland Po ugal
Resou ce link: h ps://ip .gbi .p /ip / esou ce? =coas al_plan s
Al e na i e iden i ie s: h ps://www.gbi .o g/da ase /2265c6e2-4ab7-4a 2-93a -ba37
7a7d4e43
Numbe o da a se s:2
Da a se name:E en Table
Cha ac e se :UTF-8
Download URL: h p://ip .gbi .p /ip / esou ce? =coas al_plan s
Coas al g assland ege a ion eco ds om São Miguel Island (Azo es) and ... 7
Da a o ma :Da win Co e A chi e
Da a o ma e sion:1.2
Desc ip ion: The da ase was published in he Global Biodi e si y In o ma ion
Facili y pla o m, GBIF (Calado e al. 2025a). The ollowing da a able includes all he
eco ds o which a axonomic iden i ica ion o he species was possible. The da ase
submi ed o GBIF is s uc u ed as a sample occu ence da ase ha has been
published as a Da win Co e A chi e (DwCA), which is a s anda dised o ma o
sha ing biodi e si y da a as a se o one o mo e da a ables. The co e da a ile
con ains 31 eco ds (e en ID). This GBIF IPT (In eg a ed Publishing Toolki , Ve sion
2.5.6) a chi es he da a and, hus, se es as he da a eposi o y. The da a and
esou ce me ada a a e a ailable o download in he Po uguese GBIF Po al IPT.
Column label Column desc ip ion
e en ID Iden i ie o he e en s, unique o he da ase .
loca ionID Iden i ie o he loca ions, unique o he da ase .
coun y The name o he coun y o majo adminis a i e uni in which he Loca ion occu s
(Po ugal).
coun yCode The s anda d code o he coun y in which he Loca ion occu s (PT).
s a eP o ince The name o he nex smalle adminis a i e egion han coun y (s a e, p o ince,
can on, depa men , egion e c.) in which he Loca ion occu s.
coun y The ull, unabb e ia ed name o he nex smalle adminis a i e egion han
s a eP o ince (coun y, shi e, depa men e c.) in which he Loca ion occu s.
municipali y The ull, unabb e ia ed name o he nex smalle adminis a i e egion han coun y
(ci y, municipali y e c.) in which he Loca ion occu s.
locali y The speci ic desc ip ion o he place.
e ba imLocali y The o iginal ex ual desc ip ion o he place.
loca ionRema ks Commen s o no es abou he Loca ion.
habi a The habi a o an E en (coas al g asslands).
minimumEle a ionInMe es The lowe limi o he ange o ele a ion (al i ude, usually abo e sea le el), in
me es.
decimalLa i ude App oxima e cen e poin decimal la i ude o he ield si e in GPS coo dina es.
decimalLongi ude App oxima e cen e poin decimal longi ude o he ield si e in GPS coo dina es.
geode icDa um S anda d Global Posi ioning Sys em coo dina e e e ence o he loca ion o he
sample collec ion poin s.
coo dina eUnce ain yinMe es Unce ain alue o coo dina e me ics.
coo dina eP ecision Value in decimal deg ees o a p ecision o six decimal places.
8Calado H e al
geo e e enceSou ces Na iga ion sys em used o eco d he loca ion o sample collec ions.
samplingP o ocol The sampling p o ocol used o cap u e he species (45 minu es a an a ea o 2500
m ; 30 minu es aking pho os, 15 minu es iden i iing plan s).
sampleSizeValue A nume ic alue o a measu emen o he size ( ime du a ion, leng h, a ea o
olume) o a sample in a sampling E en .
sampleSizeUni The uni o measu emen o he size ( ime du a ion, leng h, a ea o olume) o a
sample in a sampling E en .
samplingE o The amoun o e o expended du ing an E en (1 pe son).
yea Yea he sample was collec ed (2022).
mon h The in ege mon h in which he E en occu ed.
day The in ege day o he mon h on which he E en occu ed.
e en Da e The da e- ime o in e al du ing which an E en occu ed.
Da a se name:Occu ence Table
Cha ac e se :UTF-8
Download URL: h p://ip .gbi .p /ip / esou ce? =coas al_plan s
Da a o ma :Da win Co e A chi e
Da a o ma e sion:1.2
Desc ip ion: The da ase was published in he Global Biodi e si y In o ma ion
Facili y pla o m, GBIF (Calado e al. 2025a). The ollowing da a able includes all he
eco ds o which a axonomic iden i ica ion o he species was possible. The da ase
submi ed o GBIF is s uc u ed as an occu ence able ha has been published as a
Da win Co e A chi e (DwCA), which is a s anda dised o ma o sha ing biodi e si y
da a as a se o one o mo e da a ables. The co e da a ile con ains 742 eco ds
(occu enceID). This GBIF IPT (In eg a ed Publishing Toolki , Ve sion 2.5.6) a chi es
he da a and, hus, se es as he da a eposi o y. The da a and esou ce me ada a a e
a ailable o download in he Po uguese GBIF Po al IPT.
Column label Column desc ip ion
e en ID Iden i ie o he e en s, unique o he da ase .
ype Type o he eco d, as de ined by he Dublin Co e S anda d.
licence Re e ence o he licence unde which he eco d is published.
ins i u ionID The iden i y o he ins i u ion publishing he da a.
collec ionID The iden i y o he collec ion publishing he da a.
ins i u ionCode The code o he ins i u ion publishing he da a (UAc).
2
Coas al g assland ege a ion eco ds om São Miguel Island (Azo es) and ... 9
