Julho de 2005 ISSN 0874-5250

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Assoóação Lusitana de Fitossociologia (ALFA)
~éd ét:~tlnn ln+r-- rnrt1-;"'..., .... l .o rl .o Ph-U±AC'A-C!. LAJAn1 e (GIl)
 Quercetea , revista da Associação Lusitana de Fitossociologia (ALFA), tem como objectivo dar a
conhecer trabalhos originais de investigação no campo da botânica, designadamente sobre vegetação Preface
e flora. A sua publicação é periódica, pelo menos bienal.

This publication was prepared to support the excursions planned by occasion of the 48th International
Comissão Editorial: Mário Fernandes Lousã, Ma Dalila Espirito Santo e Jorge Henrique Ca pelo. Association 01 Vegetation Science (IAVS) Symposium held in Lisbon July 24 th -29 th 2005.

These excursions were organized by botanists 01 the Lusitanian Association 01 Phytosociology

(ALFA). Two kinds 01 excursions were planned: one day excursions, prepared around Lisbon to be
Comissão Redactorial : Ma Dalila Espirito Santo, Tiago Monteiro Henriques, Sandra Mesquita & Pedro
made during the Symposium; one week excursions, prepared to Madeira (the pre-symposium
Arsén io.
excursion) and to the Azores (the post-symposium excursion). These excursions were planned
considering others made by IAVS through the Iberian Peninsula and Macaronesia. The last one was a
Revisores : Alfredo Asensi Marfil (Málaga), Angel Penas (Leão), Blanca Diez Garretas (Málaga), Carlos 16 day excursion in July 1999, including a lield guide published by Rivas-Martínez et alo t
Aguiar (Bragança), Carlos Neto (Lisboa), Carlos Pinto Gomes (Évora), Francisco Barreto Caldas (Porto),
Each article 01 this volume corresponds to an excursion and tries to provide sufficient inlormation
Jesús Izco Sevillano (Santiago de Compostela), Javier Loidi (Bilbau), João Honrado (Porto), Jorge
Henriqu e Capelo (Lisboa), José Carlos Costa (Lisboa) , Manuel Costa (Valência), Ma Dalila Espírito about the Ilora and plant communities lound during the trip and its stops. The itineraries 01 the
Santo (Lisboa), Eduardo Dias (Angra do Heroísmo), Márío Fernandes Lousã (Lisboa) , Miguel Ladero geobotanical excursions were chosen in order to visit the most representative ecosystems 01 the
Alvarez (Salamanca) , Miguel Sequei ra (Funchal), Roberto Jardim (Funchal), Salvador Rivas-Martínez outskirts 01 Lisbon , the island 01 Madeira as well as the islands 01 Pico, S. Jorge and Terceira
(Madrid), Tomás Dias González (Oviedo). (archipelago of the Azores), regarding flora, vegetation and landscape. The excursions consist of stops
with short lield incursions, being also appreciated some aspects of the landscape Irom the bus, for the
excursions made at the outskirts of Lisbon. ln the stops, a more detailed study of taxa and the mosaics
Secretaria de Redacção, Serviço de Subscrições, Tesouraria: of communities, that we intend to be representative of the regional phytocenosis, will be held. As for the
Departamento de Protecção de Plantas e de Fitoecologia excursions on the islands, the sam e criteria was adopted in some days, but full day walks are also
planned. Some itineraries must be done only with the company of expertise guides, because the access
Instituto Superior de Agronomia
is difficult and some paths are dangerous to be made alone.
Tapada da Ajuda
This volume tries to be useful for the participants in the 48th IAVS Symposium and for those who
1349-018 Lisboa visit these regions, wishing to have summarized a considerable amount of new information, contributing
Fax 21 3635031 to the advance of vegetation science in Portugal.

Tel. 21 363531 66
E-mail jccosta@isa.utl.pt Mª Dalila Espírito Santo

ISSN: 087452250
Depósito Legal: 229670/05
Capa: Sandra Mesquita
Impresso: Gráfica 2000
Editado no Instituto Superior de Agronomia
1 RIVAS-MARTíNEZ, S., J. LOlol ARREGUI, M. COSTA TALENS, T. E. DíAZ GONZÁLEZ & A. PENAS MERINO (eds.) (1999) -
Publicado em 15 de Julho de 2005. Iter Ibericum AD . MIM, !tinera Geobotanica 13: 5-347

3
CONTENTS

The Landscape and Plant Communities of Serra da Arrábida

José Carlos Costa, Jorge Capelo, Pedro Arsénio &Tiago Monteiro
Henriques ..................... ................................................................. ............................................ 7

Natural Succession in Southwestern Portuguese Limy Soils: After Fire Dynamics

(Arrábida Natural Park) and Primary Succession in Sand Dunes (Tip of Peninsula of
Troia)
Otília Correia & Manuel João Pinto ........................................................................................ 27

Landscape Ecology of the Sado River Estuary (Portugal). (Dunes, Fresh and Salt
marshes)
Carlos Neto, Mª Eugénia Moreira & Rute Caraça ................................................................. .43

Sintra Vegetation and Landscape

Sandra Mesquita, Pedro Arsénio, Mário Lousã, Tiago Monteiro Henriques & .
José Carlos Costa ................................................................................................................. ..65

Vegetation and Landscape of Serra de Montejunto

Maria Dalila Espírito Santo, Ilídio Moreira & Patricia Rodríguez González .......................... .83

The vegetation of Madeira Island (Portugal). A brief overview and excursion guide
Jorge Capelo, Miguel Sequeira, Roberto Jardim, Sandra Mesquita &
José Carlos Costa ......................................................... ........................................................ 105

The Azores Centrallslands Vegetation and Flora. Field Guide

Eduardo Dias, Cândida Mendes, Cecília Melo, Dinis Pereira & Rui Elias .......... ..................123
 The Landscape and Plant Communities 01 Serra da Arrábi da

The Landscape and Plant Communities of Serra da Arrábida

José Carlos Costa' , Jorge Capelo**, Pedro Arsénio***, Tiago Monteiro Henriques'

The lield trip takes place in the

biogeographic territories 01 the
Sadensean-Dividing Portuguese
Subprovince (Coastal Lusitan-
Andalusian Province, Mediterranean
Region), throughout the upper
thermomediterranean thermotype and
subhumid ombrotype.

The trip will start in Lisbon (Fig. 1),

which is located in the Olissiponean
Superdistricl, within the Divid ing
Portuguese Sector. During the trip, it is
possible to observe on the North bank
01 lhe Tagus River, on vertisols
originated both Irom acidic and basic
bedrocks, a forest commun ily
dominated by the oleaster-tree (Viburno
tini-O/eetum sy/vestris) . The perenn ial
grassland community Cariei depressae-
Hyparrhenietum sinaieae is a
conspicuous subseral stage 01 this
Fig. 1 - Itinerary of the excursion woodland, predeceased by a kermes
oak (Quereus eoeeifera) scrub
communily (Melieo arreetae-Quereetum eoeeiferae asparagetosum a/bidJ). These communities (except
the later) occur also in Serra da Arrábida .

• Departamento de Protecção de Plantas e de Fitoecologia. Instituto Superior de Agronomia. Tapada da Ajuda 1349-
017 Lisboa. Portugal. jccosta @isa.utl .. pt

,. Departamento de Ecologia. Estação Florestal Nacional. Quinta do Marquês. 2780-159 Oeiras. Portugal.
jorge .capelo@efn.com.pt

... Secção Autónoma de Arquitectura Paisagista. Instituto Superior de Agronomia. Tapada da Ajuda 1349-017
Lisboa. Portugal. arseniop@ isa.utl.. pt

7
Costa, J.C, & aI.
The Landscape and Plant Communities 01 Serra da Arrábida

Crossing the Tagus River, we arrive at the Sadensean Superdistrict, which is inciuded in the
Ribatagan-Sadensean Sector, Two distinct vegetati on series can be lound on this te rritory, both headed
by cork oak (Quercus suber) wood lands: Asparago aphy/li-Querco suberis sigmetum, on sandstone
derived soils, and Oleo sylvestris-Querco suberis sigmetum, on slightly hidromorphic sandy soils, Many
01 these lorests were thinned and translormed into "montados" parklands or into pinewoods 01 Pinus
pinea or Pinus pinaster,

ln this territory, the potential and seral stages 01 the Asparago aphy/li-Querco suberis sigmetum are:
Asparago aphy/li-Quercetum suberis (mature lorest), Buplero fruticosae-Arbutetum unedonis (thicket or
mantle high-shrub hedge), Erico-Quercetum lusitanicae ulicetosum welwitschianii (Iow-shrub
community), Avenulo sulcatae-Stipetum giganteae (perennial grassland) and Erico umbe/latae-Ulicetum
welwitschiani (heath scrubland) .

ln turn, the Oleo sylvestris-Querco suberis sigmetum is made up by the lollowing potential and seral
stages: Oleo sylvestris-Quercetum suberis (mature lorest), Junipero navicularis-Quercetum lusitanicae
(shrubby community) or Asparago aphy/li-Myrtetum communis (shrubby community in
edaphohigrophilous situation), Stipa gigantea community (perennial grasslands), Thymo capite/lati-
Stauracanthetum genistoides (gorse scrubland on deep paleodune sand), Erico umbe/latae-Ulicetum
welwitschiani (heath scrubland on compact substrata) and Corynephoro macranthero-Arenarietum
algarbiense (annual community). The lirst stop will be made in one 01 the local pinewood 01 "Apostiça"
(Fig. 2), in order to look at some 01 the seral stages included in this sigmasyntaxon, well under the pine
canopy. The originallity and rarity 01 some 01 them are noteworthy, namely the shrub community 01 the
endemic juniper (Juniperus navicularis) and dwarl-oak (Quercus lusitanica) - Junipero navicularis-
Quercetum lusitanicae - as well as the Thymo capite/lati-Stauracanthetum genistoides thorny
scrubland.

Serra da Arrábida is a national protected area, classilied under the status 01 'Natural Park'. It
integrates the national NATURA 2000 site list, both as a special area of conservation (Habitats
Directive) and as a special protection area (Birds Directive).

. ',"
Geologically speaking, this sea-Iacing small mountain (501 meters above m.s.m.) is a smalll imestone
outcrop, mostly 01 Jurassic origino Its dominant soils are derived Irom hard calcium carbonate rocks,
some with dolomitical character. ln terms 01 the bioclimate typology (Rivas-Martinez et ai., 2004) it is
located on the thermomediterranean, upper sub-humid belt and biogeographically stands as the
... ,' , :' Arrabidensean Superdistrict (Ribatagan-Sadensean Sector). It shares a large proportion 01 its II0ra and
vegetation with the larger calcareous unit - Dividing Portuguese Sector - lound north 01 the Tagus River
(Costa et ai., 1998).

Its natural vegetation , generally well preserved , has a high natural value due to the lact that the
majority 01 the taxa share a paleomediterranean and/or paleotropical origino This lacto r, together with a
relatively high annual precipitation rate, no hail, and soils originated Irom dolomitical limestone,
determine a vegetation 01 great richness and originality.
.·~t · ..
- "-" -''- '-'- ' -:..:::..:. .

8 9
Costa, J.C. & aI.
The Landscape and Plant Communilies 01 Serra da Arrábida
One such example can be found on lhe dry, rocky slopes facing lhe sea or on ravine gravei deposits:
lhe relicl nanophanerophylic communily dominaled by Euphorbia pedroi: Convolvulo fernandesii-
Euphorbietum pedrai. It forms a xeromorphic permanent phytocoenosis in hiperoceanic, lopographically
xeric and Ihermomediterranean bioclimale. Th is community will be visited on lhe second stop aI lhe
"Califórnia" cliffs over East Sesimbra (Fig. 3).

CD

...

~~"
Fig. 4 - Morro de Jaspe-Selúbal: 1 Ouercus rotundifolia; 2 Ouerco cocciferae-Juniperetum turbinatae;
3 Phlomido Iychnitidis-Brachypodietum phoenicoidis; 4 Iberido micracarpae-Stipetum offneri; 5 Phlomido
purpureae-Cistetum albidi
,j//'JY
/7~ ,:-
~ ")..
T~I H
/ "0'

Fig. 3 - Califórnia-Sesimbra: 1 Ouerco cocciferae-Juniperetum turbinatae; 2 Phlomido purpureae- Anolher stop will be made on "Mata do Solitário" (Fig. 5), a well-known botanical location. From Ihat
Cistetum albidi; 3 Convolvulo fernandesii-Euphorbietum pedroi; 4 Cariei depressae-Hyparrhenietum site iI can be observed lhe climatophilous woodland localed in the well-drained dolomilical slopes, wich
sinaicae; 5 Phlomido Iychnitidis-Brachypodietum phoenicoidis; 6 Sede tum micrantho-sediformis is formed by micro-mesophanerophytes and dominated by lhe tree-quermes oak - Ouercus coccifera
subsp. rivasmartinezii - reaching 14-15 meters in heighl. This small Iree dominates the mature slage of
lhe local vegetalion series, lhe Vibumo tini-Ouercetum rivasmartinesii. The referred woodland has as
Over lhe Serra's summil, an edaphoxerophilous permanenl high-scrub juniper communily is Ihickel or mantle fo resl, a community dominaled by lhe strawberry Iree - Bupleuro fruticosae-Arbutetum
inlerpreled as lhe climax: Querco cocciferae-Juniperetum turbinatae. The slrongly xeric characler of Ihis unedonis - bul iI is also common lo find, on lhe clearings, lhe hemicriplophylic communily Leucanthemo
vegelalion is leslified by lhe co-dominance of Olea europeae var. sylvestris, Asparagus albus and sylvatiei-Cheir%phetum sempervirentis, dominated by Cheirolophus sempervirens. The next seral
ecologically similar taxa in some of Ihese biolopes. Normally, primary posilions of lhe Ouerco- slage is the already referred community co-dominaled by Ouercus coccifera and Juniperus turbina ta
Juniperetum turbinatae have a perennial grass communily as a subslilulion slage: Iberido micracarpae- (Ouerco cocciferae-Juniperetum turbinatae). The communily Phlomido purpureae-Cistetum albidi can
Stipetum offneri. This community, which can be observed in lhe sleepesl and more eroded slopes, is also be found as a second substilution slage within this series . It's a low-scrub communily found in deep
well adapled to fire. The second slage is eilher Salvio sclareoidis-Ulicetum densi thymetosum sylvestris clay-rich soils Ihal suffered erosion of ils top horizons.
or Phlomido purpureae-Cistetum albidi, dominaled by Cistus albidus. Similarly, at Serra da Arrábida lhe
At lhe botlom of the larger valleys, where winter water runs and some higrophilous compensation
Ouercus rotundifolia is always a scrub (2-3 m) never reaching a Iree size and Ihus never eslablishing as
persists during summertime, and at deep clay-rich soils usually derived from limy marl, lhe woodland is
lall woodland. The pulalive explanalion for this facl lies also in lhe high calcium/magnesium soil rale,
dominaled by Ouercus faginea subsp. broteroi - Arisaro clusii-Ouercetum broteroi. A community of
becoming loxic lo lhe holm oak. This kind of vegelalion can be observed aI lhe Ihird slop "Morro de
Rubus u/mifo/ius is ils prickly bramble Ihickel (Lonicero hispanicae-Rubetum ulmifoliae). The olhe r
Jaspe" (Fig. 4).
subslitulion slages of these series are lhe sam e of lhe Viburno tini-Querco rivasmartinezii sigmetum.
 CostarJ.C. & aI.
The Landscape and Planl Communilies 01 Serra da Arrábida

The communily Phlomido Iychnitidis-Brachypodietum phoenicoidis (dominated by the perennial lall

grass Brachypodium phoenicoides) , an importanl Orchidaceae habilal, is ralher ubiquilous on lhe
Serra's loolhills. Often seen on deeper soils, ii also Irequenlly appears in mosaic wilh lhe communily
Salvio sclareoidis-Ulicetum densi thymetosum sylvestris (dominaled by U/ex densus and Thymus zygis
subsp. sylvestris), which occupies lhe rockier and mosl eroded posilions.

Olher equally inleresling communilies occupy some smaller palches 01 Ihis landscape: The Sileno
longiciliae-Antirrhinetum linkiani is a dwarf perennial chasmo-comophyle communily lound on limeslone
walls and on rock crevices 01 Serra da Arrábida. On Ihose locations ii can also be lound lhe chasmolilic
community Narciso calcicolae-Asplenietum rutae-murariae. The succulenl planls Sedum sediforme and
Sedum album varo micranthum are common on Ilal rocks and lorm lhe Sede tum micrantho-sediformis
community.

Finally, a relerence lo lhe community Helianthemo stoechadifoliae-Limonietum virgatae, Ihal takes

place on sea cliffs splashed by marine sall spray.

Synthetic tables and relevees of the most representative communities are presented below:

Synthetic lable 01 Vibumo tini·Oleetum sylvestris J.C. Costa, Capelo & Lousã 1994 (Ouerco-
Oleion, Ouercetalia ilicis, Ouercetea ilieis) , lrom Costa et ai. 1994, 24 relevees: Characteristics: Olea
eurapaea varo sylvestris V, Rhamnus alaternus V, Smilax aspera varo altissima V, Rubia peregrina varo
longitolia V, Vinca difformis V, Arum italicum S. I. V, Arisarum vulgare varo clusii V, Tamus communis V,
Viburnum tinus IV, Pistaeia lentiscus IV, Asparagus albus IV, Ruscus aculeatus IV, Asparagus aphy//us
IV, Ceratonia siliqua III, Ouercus coceifera III, Rosa sempervirens III, Acanthus mo//is II, Rhamnus
oleoides III, Phi//yrea latifolia III, Oaphne gnidium III, Arbutus unedo II, Lonicera implexa II, Osyris alba
II, Corani//a glauca II, Hedera maderensis subsp. iberica II, Lonicera etrusca II, Fraxinus angustifolia II,
Ulmus minor II, Myrtus communis I, Carex distachya I, Anemone palmata I, Iris foetidissima I, Ouercus
suber +, Ouercus rotunditolia +, Juniperus turbina ta +, Jasminum fruticans +, Teucrium fruticans +,
Phlomis purpurea +, Laurus nobilis r, Erica arborea r, Phi//yrea media r, Paeonia broterai r, Bupleurum
fruticosum r, Bupleurum rigidum subsp. paniculatum r, Carex ha//erana r, Euphorbia charaeias r;
companions: Bryonia dioica V, Rubus ulmifolius IV, Smirnum olusatrum IV, Geranium purpureum IV,
w
Piptatherum miliaceum IV, Urtica membranacea IV, Urginea maritima IV, Lonicera peryclimenum subsp.
hispanica III, Parietaria judaica III, Prunus spinosa subsp. insititioides II, Carataegus monogyna subsp.
brevispina II, Ruta chalepensis II, Oactylis hispanica II, Galium aparine II, Brachypodium phoenicoides
II, Orobanche ramosa subsp. nana II, Salvia sclareoides II, Asphodelus lusitanicus II, Calamintha
baetica I, Asparagus aspargoides I, Salpichroa origanifolia I, Opuntia stricta I, Silene dioica I, Ulex
jussiaei I, Cistus monspeliensis I, Cistus albidus I, Cistus salviifolius +, Viola riviniana +, Phagnalon
saxati/e +, Rosmarinus officinalis +, Allium roseum +, Polypodium australe +, Rosa canina r,
Brachypodium sylvaticum r, Silene longicilia r, Barlia robertiana r, Lavandula multifida r, Aristolochia

12 13
Costa, J.C . & aI. The Landscape and Plant Communities 01 Serra da Arrábida

paucinervis r, Biscute//a lusitanica r, Astragalus lusitanicus r, Allium neapolitanum r, Narcissus ulmifolius, + Ulex australis subsp. welwitschianus, + Halimim halimifolium, + Halimium calycinum, +
bulbucodium varo obesus r. Lavandula sampaiona subsp. lusitanica, + Lavandula luisieri, + Thymus vi//osus, + Cistus salvifolius, +
Cistus monspeliensis.

Synthetic table 01 Asparago aphy/li-Quercetum suberis J.C. Costa, Capelo, Lousã & Espírito
Santo t 996 (Ouerco-Oleion, Ouercetalia ilieis, Ouercetea ilicis), Irom Costa et aI. 2002, 30 relevees: Synthetic table 01 Vibumo tini-Quercetum rivasmartinezii Rivas-Martínez, Lousã, T.E. Días,
Characteristics: Ouercus suber V, Smilax aspera varo altissima V, Rubia peregrina varo longifolia V, Fernandez González & J.C. Costa 1990 corro Capelo & J.C. Costa 2001 (Ouerco-Oleion, Ouercetalia
Asparagus aphy//us V, Ouercus faginea subsp. broteroi IV, Rhamnus alaternus IV, Daphne gnidium IV, ilicis, Ouercetea ilicis) , Irom Capelo & Almeida 1993, 10 relevees: Characteristics: Ouercus coceifera
Hedera maderensis subsp. iberica IV, Teueium scorodonia IV, Arbutus unedo III, Olea europea var. subsp. rivasmartinesii V, Viburnum tinus V, Pistacia lentiscus V, Arbutus unedo V, Phi//yrea latifolia V,
sylvestris III, Rosa sempervirens III, Pistacia lentiscus III, Ouercus coccifera III, Myrtus communis III , Smilax aspera varo altissima V, Ruscus aculeatus V, Arisarum vulgare var. clusii V, Phi//yrea media III ,
Osyris alba III, Ruscus aculeatus III, Euphorbia charaeias III, Ouercus lusitanica III, Lonicera etrusca III , Asplenium onopteris IV, Rubia peregrina var. longifolia IV, Olea europeae var. sylvestris III, Ouercus
Arisarum vulgare var. clusii III, Deschampsia stricta III, Sei//a monophy//os III, Phi//yrea latifolia II, coccifera subsp cocciofera III, Bupleurum fruticosum III, Deschampsia stricta III, Hyacintoides hispanica
Viburnum tinus II, Erica arborea II, Phi//yrea angustifolia II , Carex distachya II, Asplenium onopteris II, III, Myrtus communis III, Coroni//a glauca II, Erica arborea II, Vinca difformis II, Lonicera implexa II ,
Pulicaria odora II, Vinca difformis II, Iris foetidissima II, Epipactis tremolsii II, Cephalantera longifolia II , Phi//yrea angustifolia II, Ouercus faginea subsp. broteroi I, Osyris lanceolata I, Rhamnus oleoides I,
Laurus nobilis I, Lonicera implexa I, Biarum galiani I, Luzula forsteri subsp. baetica I, Asparagus Rhamnus alaternus I, Jasminum fruticans I, Daphne gnidium I, Asparagus aphy//us I, Peaonia broteroi I,
acutifolius I, Rhamnus oleoides +, Ouercus x airensis +, Genista tournefortii +, Hyacintoides hispanica Carex distachya I, Genaria diphy//a I; companions: Rubus ulmifolius IV, Tamus communis IV, Ceterach
+, Anemone palmata +, Sanguisorba hybrida +, Melica minuta subsp. arrecta +, Coroni//a glauca r, offieinarum III, Holometecium serieium II, Teucrium scorodonia II, Lonicera periclymenum subsp.
Paeonia broteroi r, Carex ha//erana r; companions : Rubus ulmifolius V, Tamus communis IV, Cistus hispanica I, Geranium purpure um I, Aristolochia paucinervis I.
salviifolius IV, Ulex jussiaei IV, Crataegus monogynea subsp. brevispina III , Prunus spinosa subsp.
insititioides III, Lonicera peryclimenum subsp. hispanica III, Lavandula luisieri III, Brachypodium
phoenicoides III, Cheirolophus sempervirens III, Calamintha baetica III, Origanum virens III, Urginea Synthetic table 01 Arisaro c/usii-Quercetum broteroi Br.-BI., P. Silva & Rozei ra 1956 (Ouerco-
maritima III, Geranium purpureum III, Aristolochia paueinervis III, Lavandula luisieri II, Pteridium O/eion, Ouerceta/ia i/ieis, Ouercetea i/ieis), Irom Capelo et alo 2002, 7 relevees: Characteristics:
aquilinum II, Erica scoparia II, Erica cinerea II, Lithodora prostrata II, Cistus crispus II , Genista Ouercus faginea subsp . broteroi V, Viburnum tinus V, Smi/ax aspera varo altissima V, Ruscus acu/eatus
triacanthus II, Cistus psilosepalus II, Clinopodium vulgare subsp. arundanum II, Piptaterum mileaceum V, Arisarum vu/gare var c/usii V, Arbutus unedo IV, Rubia peregrina var. /ongifo/ia IV, Asp/enium
II, Asphodelus lusitanicus II, Dactylis glomerata subsp. lusitanica II, Thapsia vi//osa II, Arum italium II, onopteris IV, Carex distachya IV, Phi//yrea la tifo/ia III, Phi//yrea media III, Erica arborea III, Ouercus
Ouercus pyrenaica I, Castanea sativa I, Fraxinus angustifolia I, Ca//una vulgaris I, Tamus communis I, coccifera III, Vinca difformis III , Lonicera etrusca III, Genista tournefortii III, Teucrium scorodonia III, /ris
Ulex airensis I, Rosa canina I, Bryonia dioica I, Prune//a vulgaris subsp. estremadurensis I, foetidissima III, Hyacintoides hispanica III, Se/agine//a denticu/ata III , Laurus nobi/is II, Myrtus communis
Pseudarrhenatherum longifolium I, Geum sylvaticum I, Agrimonia eupatoria I, Origanum virens I, II , Coroni//a g/auca II, Lonicera imp/exa II, Phi//yrea angustifo/ia II, Jasminum fruticans II, Paeonia
Lathyrus clymenum I, Clinopodium vulgare I, Polypodium australe I, Cytisus striatus +, Brachypodium broteroi II, Euphorbia charaeias II , Bup/erum rigidum subsp. panicu/atum II, O/ea europeae var.
sylvaticum +, Rosmarinus officinalis +, Hypericum perforatum +, Silene longicilia +, Frangula alnus r, sy/vestris I, Acer monspessu/anum I, Rhamnus a/aternus I, Deschampsia stricta I, Asparagus aphy//us I,
Salix atrocinerea r, Erica lusitanica r, Orobanche hedera r, Stachys germanica subsp. lusitanica r, Anemone pa/mata I, Asparagus acutifo/ius I, Cepha/antera /ongifo/ia I, Me/ica minuta subsp. arrecta I,
Digitalis purpurea r, Ranunculus ficaria r, Carduus broteroi r, Smirnium olusatrum r, Arrhenatherum Pu/icaria odora I; companions: Rubus u/mifo/ius V, Tamus communis IV, Geranium purpureum IV,
album subsp. baeticum r, Agrostis curtisii r, Cistus ladaniferi +, Tuberaria lignosa r, Simethis matiazii r, Ca/amintha baetica III, Brachypodium sy/vaticum III, Cistus sa/vifo/ius III, Cheir%phus sempervirens II ,
Astragalus lusitanicus r, Cistus monspeliensis r, Inula conyza r, Orobanche latisquama r. Origanum virens II, Salvia sc/areoides II, Po/ypodium cambricum var. serru/atum II, Brachypodium
phoenicoides I, Lonicera peryclimenum subsp. hispanica I, Arist%chia paucinervis I, Euphorbia
amygda/oides I, Agrimonia eupatoria I, Narcissus ca/cico/a I, Arum italicum I, Be//is sy/vestris I, Carex
Relevee 01 Oleo sylvestris-Quercetum suberis Rivas Goday, F. Galiano & Rivas-Martínez in divu/sa I, Carex flacca varo serru/ata I, Piptaterum mi/eaceum I, Asp/enium trichomanes var.
Rivas-Martínez 1987 (Ouerco-O/eion, Ouerceta/ia i/icis, Ouercetea i/icis) , Cova da Morena, 200 m2, N, quadriva/ens I.
100 m, Irom Capelo & Almeida 1993: Characteristics: 4 Ouercus suber, 2 O/ea sy/vestris, 2 Asparagus
aphy//us, 1 Arbutus unedo, + Ouercus coceifera, + Pistaeia /entiscus, + Juniperus navicu/aris, + Smi/ax
aspera var. altissima, + Rubia peregrina varo longifo/ia, + Ruscus acu/eatus; companions: + Rubus
Costa, 'J.C . & aI. The Landscape and Plan! Communities of Serra da Arrábida

Synthetic table 01 Ouerco cocciferae-Juniperetum turbinatae (Rivas-Martínez 1975) Rivas - Relevée 01 Convo/vu/o fernandes ii-Euphorbietum pedroi G. Pedro ex Capelo 2003 (Asparago a/bi-
Martínez, Lousã, T.E . Díaz, Fernández-González & J.C. Costa 1990 (Asparago a/bi-Rhamnion o/eoidis, Rhamnion o/eoidis, Pistacio /entisci-Rhamneta/ia a/atemi, Quercetea i/ieis): Califórnia (Sesimbra), 60
Pistacio /entisci-Rhamneta/ia a/atemi, Quercetea i/ieis), Iram Capelo & Almeida 1993, 27 relevees: m2, S, 120 m, Characteristics: 4 Euphorbia pedroi, 1 O/ea europeae varo sy/vestris, 1 Rhamnus
Characteristics: Juniperus turbinata V, Rhamnus o/eoidis V, Quercus coccifera IV, Pistacia /entiscus o/eoides, 1 Phi//yrea angustifo/ia, 1 Pistacia /entiscus, + Juniperus turbinata + Asparagus a/bus, +
IV, Lonicera imp/exa IV, Smi/ax aspera varoaspera IV, Phi//yrea angustifo/ia III, Rhamnus a/aternus III , Asparagus aphy//us, + Arisarum vu/gare varo c/usii; companions: 1 Rosmarinus officina/is, + Cistus
O/ea europea varo sy/vestris III, Rubia peregrina varo /ongifo/ia III, Oaphne gnidium III, Asparagus monspe/iensis, + Cistus a/bidus, + Brachypodium retusum, + Hyparrhenia hirta.
aphy//us III, Arisarum vu/gare varoclusii III, Anemone pa/mata III, Arbutus unedo II, Jasminum fruticans
II , Osyris alba II, Ruscus acu/ea tus II, Bup/erum rigidum subsp. panicu/atum II, Carex distachya II ,
Coroni//a g/auca II, Asparagus a/bus II, Myrtus communis II, Ceratonia si/iqua II, Phi//yrea /atifolia I, Synthetic !able of Erico-Ouercetum lusitanicae Rothmaler ex 8r. -8i. , P. Silva & Rozeira 1964
Phi//yrea media I, Genista toumefortii I, Ph/omis purpurea I, Euphorbia charaeias I, Lonicera etrusca I, (Quercion fruticosae, Pistacio /entisei-Rhamneta/ia a/atemi, Quercetea i/icis) , Irom Costa et alo2002, 28
Me/ica minuta subsp. arrecta I, Sei//a monophy//us I, Gennaria diphy/la I, Quercus rotundifo/ia +, Osyris relevees: Characteristics : Quercus /usitanica V, Myrtus communis IV, Asparagus aphy//us IV, Serratu/a
/anceo/ata +, Asparagus acutifo/ius +, Vibumum tinus +, Bup/erum fruticosum +, Rosa sempervirens +, monardii IV, Oaphne gnidium III, Sei//a monophy//us III, Phi//yrea angustifolia III, Quercus suber III ,
Hyacintoides hispanica +, Paeonia broteroi r, Asp/enium onopteris r; companions: Brachypodium Rubia peregrina varo /ongifo/ia III, Rhamnus a/atemus II, Euphorbia transtagana II , Arbutus unedo II ,
phenicoides III, Rosmarinus officina//is II, Cistus a/bidus II, Cistus monspe/iensis II, Oacty/is hispanica II , Anemone pa/mata II, Pulicaria odora II, Pistacia /entiscus I, O/ea europea var sy/vestris I, Quercus
Antirrhynum linkianum II, Astraga/us /usitanicus II, Origanum virens II, Cistus crispus II, Cistus sa/vifo/ius coccifera I, Osyris alba I, Carex oediposty/a I, Carex distachya +, Centaurea africana +, Erica arborea +,
II , Urginea maritima II, Carex ha//erana II, Asphode/us aestivus II, Pulicaria odora II, Lavandu/a /uisieri Se/agine//a denticu/ata +, Oanthonia decumbens +; companions: Erica scoparia V, Cistus sa/viifo/ius
II ,Ca/amintha baetica I, Stahe/ina dubia I, Ruta cha/epensis I, U/ex densus I, Erica scoparia I, IV, Lavandu/a /uisieri IV, U/ex jussiaei III, Lithodora prostrata subsp. /usitanica III , Erica umbe//ata III ,
Asphode/us /usitanicus I, Be//is sy/vestris I, Sa/via scareoides I, Sanguisorba spachiana I, Centaurium Ca//una vu/garis III, Asphode/us /usitanicus III, Pterospartum tridentatum subsp . tridentatum III , Genista
grandiflorum I, Carex flacca I, Anthy//is vu/neraria subsp. maura I, Oaucus maximus I, Convo/vu/us triacanthos III, Brachypodium phoenicoides II, Cistus crispus II, Urginea maritima II, Thapsia vi//osa II,
altheoides I, Erygium di/a ta tum I, Rubus u/mifo/is +, Stipa offeneri +, Thymus mastichina +, Lathyrus Avenu/a su/cata II, Stipa gigantea II, U/ex airensis II, Simethis maltiazzi II, U/ex austra/is subsp.
sy/vestris +, Tu/ipa austra/is +, Ophris fusca +, Anaga//is mone//i var. /inifolia +, Andrya/a integrifo/ia +, we/witschianus II, Oacty/is g/omerata subsp. /usitanica II, Ho/cus /anatus II, Agrostis curtisii I, Pteridium
/ris subbiflora +, Narcissus bu/bucodium subsp. obesus +, Ph/omis /ychinitis +, Tamus communis r, aqui/inum I, Thymus vi//osus I, Cistus psi/osepa/us I, Arrhenatherum a/bum I, Ha/imium ocymoides I,
Ceterach officinarum r, Narcissus ca/cico/a r. Tuberaria /ignosa I, Cistus monspe/iensis I, Erica austra/is +, Stachys officina/is subsp. a/geriensis +,
U/ex minor r, Lonicera peryclimenum subsp. hispanica r, Rubus u/mifo/ius r, Crataegus monogyna
subsp. brevispina r, A/lium pruinatum r, Hyparrhenia hirta r, Friti/aria /usitaica subsp. stenophy//a r,
Synthetic table 01 Asparago aphy/li-Myrtetum communis Rivas-Martínez, Cantó, Fernández- Oipecadi serotini r, Sesamoides canescens r.
González & Sanchez-Mata ex JC. Costa, Lousã & Espírito Santo 1996 (Asparago a/bi-Rhamnion
o/eoidis, Pistacio /entisci-Rhamneta/ia a/a temi, Quercetea i/icis) , Irom Costa et alo2004, 12 relevees:
Characteristics: Myrtus communis V, Asparagus aphy//us V, Pistacia /entiscus V, Quercus coccifera IV, Synthetic table 01 Junipero navicu/aris-Ouercetum lusitanicae (Rothmaler 1954) Rivas-Martínez,
Phi//yrea angustifo/ia IV, Quercus suber IV, Smi/ax aspera var. aspera IV, Oaphne gnidium III, Rubia Lousã, T.E. Díaz, Fernández-González & J.C. Costa 1990 (Quercion fruticosae, Pistacio /entisei-
peregrina var /ongifo/ia III, O/ea europaea var sy/vestris III, Quercus /usitanica II, Rhamnus o/eoides I, Rhamneta/ia a/atemi, Quercetea i/icis), Irom Costa et ai. 1994, 8 relevees: Characteristics: Quercus
Rhamnus a/atemus I, Osyris alba I, Asparagus acutifo/ius +, Pyrus bourgeaena +, Arbutus unedo +, /usitanica V, Juniperus navicu/aris V, Serratu/a a/ca/ae V, Corema a/bum IV, Oaphne gnidium IV,
Tamus communis +, Crataegus monogynea subsp. brevispina +; companions: Cistus sa/viifo/ius V, Phi//yrea angustifo/ia III, Euphorbia transtagana III, Sei//a monophy//us II, Anemone pa/mata III ,
Rubus u/mifo/ius III, U/ex austra/is subsp. we/witschianus III, Brchypodium phoenicoides III, Cistus Asparagus aphy//us II, Quercus coccifera II, Arbutus unedo I, Pistaeia /entiscus I, Osyris alba I, Ruscus
psi/osepa/us III, Ho/cus /anatus II, Cistus crispus II, Erica scoparia II, Cistus /adanifer II, Cynodon acu/eatus I, Quercus suber I; companions : Stipa gigantea V, Ha/imium ca/ycinum IV, Ha/imium
dacty/on II , Foenicu/um vu/gare subsp. piperitum II, Oitrichia viscosa subsp . viscosa II, U/ex jussiaei I, halimifo/ium IV, U/ex austra/is subsp. we/witschianus IV, Erica umbe/ata IV, Ca//una vu/garis IV, Cistus
Tori/is neg/eta I, H%schoenus romanus subsp. australis I, Lavandu/a /uisieri I, Hyparrhenia sinaica +, sa/viifo/ius IV, Lithodora prostrata subsp. /usitanica IV, Tuberaria gultata IV, Thymus capite//atus III ,
Cistus monspe/iensis +, Arundo donax +, Oaucus crinitus +, Oacty/is g/omerata subsp. /usitanica +, Stauracanthus genistoides III, Lavandu/a sampaiona subsp. /usitanica III, Lavandu/a /uisieri III, Urginea
Sco/ymus macu/ata +, Mantisa/ca sa/mantica +, Rumex crispus +, Juncus acutiflorus subsp. rugosus +, maritima III , Thapsia vi//osa III, Genista triacanthus III , Linaria spartea III, C/adonia sp. III , /beris /inifolia
Ha/imium ca/yeinum +, Ha/imium halimifo/ium +, Tuberaria lignosa +, Ca//una vu/garis +. subsp. we/witschii II, Cistus crispus II , Pterospartum tridentatum subsp. tridentatum II , Tuberaria /ignosa
II , Andrya/a integrifo/ia II, Helichrysum picardii varo virescens I, Orchis mascu/a I, Cytinus machrantherus

16 17
Costa, J.C. & aI. The Landscape and Plant Communities 01 Serra da Arrábida

I, Magydarispanicifolia I, Asphodelus aestivus I, Arrhenatherum album I, Pteridium aquilinum I, Relevée 01 Phlomido purpureo-Cistetum albidae Rivas-Martínez, Lousã, T.E. Díaz, Femández-
Elaeoselinum gummiferum I. González & J.C. Costa 1990 (Ulici argentei-Cistion ladaniferi, Lavanduletalia stoechadis, Cisto-
Lavanduletea): Santana (Sesimbra), 20m 2, S, 210m, Charac!eris!ics: 3 Cistus albidus, 2 Cistus
monspeliensis, 1 Cistus salviifolius, 1 Lavandula luisieri, 1 Thymus mastichina, + Phlomis purpurea, +
Synthetic table 01 Bupleuro fruticosae-Arbutetum unedonis Capelo, J.C. Costa' & Rivas-Martínez Cistus crispus, + Astragalus lusitanicus; Companions: 2 Rosmarinus officinalis, 1 Thymbra capitata, 1
2002 (Arbuto unedonis-Laurion nobilis, Pista cio lentisci-Rhamnetalia alatemi, Quercetea ilicis), Iram Brachypodium phoenicoides, 1 Oaphne gnidium, + Hyparrhenia sinaica, + Carlina corymbosa, +
Capelo et ai. 2002, 9 relevees : Charac!eris!ics: Arbutus unedo V, Erica arborea V, Vibumum tinus V, Arisarum vulgare, + Oactylis hispanica, + Reichardia picroides, + Asparagus aphy//us, + Urginea
Coroni//a glauca V, Smilax aspera varo aspera V, Phi//yrea angustfolia V, Pistacia lentiscus V, maritima, + Sipa gigantea.
Asparagus aphy//us V, Buplerum fruticosum IV, Rubia peregrina var. longifolia IV, Myttus communis IV,
Quercus coccifera subsp. coccifera IV, Phi//yrea latifolia IV, Oaphne gnidium IV, Rosa sempervirens IV,
Rhamnus alatemus IV, Vica difformis IV, Olea europea varo sylvestris III, Lonicera implexa III, Ruscus Synthetic table 01 Thymo capitellati-Stauracanthetum genistoides (Rothmaler 1954) Rivas-
aculeatus III, Arisarum vulgare var. clusii III, Osyris alba III, Euphorbia characias III, Quercus faginea Martínez, T.E. Díaz & Fernández-González (Coremion albi, Stauracantho genistoidis-Halimietalia
subsp. broteroi II, Rhamnus oleoides II, Lonicera etrusca II, Laurus nobilis II, Anemone palmata II , commutati, Cisto-Lavanduletea) , Iram Neto, 2002, 29 releves: Charac!eris!ics: Stauracanthus
Buplerum rigidum subsp. paniculatum II, Jasminum fruticans II, Genista toumefottii II, Juniperus genistoide V, Thymus capite//atus V, Halimium calycinum V, Halimium halimifolium V, Lavandula
turbinata I; companions: Rubus ulmifolius IV, Teuc'rium scorodonia IV, Brachypodium phoenicoide IV, sampaiona subsp. lusitanica V, Ulex australis subsp. welwitschianus V, Helichrysum picardi var
Crataegus monogyna subsp. brevispina IV, Tamus communis III, Erica scoparia III, Cistus virescens IV, Armeria royana II, Lithodora prostra ta subsp. lusitanica II, Corema album II , Malcolmia
monspeliensis III, Cistus salviifolius III, Calamintha baetica III, Cheirolophus sempervirens III, Pulicaria lacera subsp. gracilima II, Oianthus broteri subsp. hioxianus II, Cistus salviifolius I, Iberis linitolia subsp.
odora III , Geranium purpureum III , Prunus spinosa subsp. insititiodes II, Lonicera hispanica II , welwitschii I, Euphorbia boetica +, Cistus libanotis r, Armeria pinifolia r; companions: Cladonia
Lavandula luisieri II, Picris spinitera I, Astragalus lusitanicus I, Cistus crispus I, Oactylis glomerata potentosa III, Cladonia mediterranica III, Arrhenatherum album III, Conyza bonariensis III, Pinus
subsp. hispanica I, Cistus albidus I, Bryonia dioica I, Iris foetidissima I, Salvia scareoides I, Geum pinaster II, Corynophorus canescens varo maritimus II, Carlina corymbosa II, Santolina impressa II ,
sylvaticum I. Sedum sediforme II, Euphorbia pottlantica II, Oaphne gnidium II, Pinus pinea II, Eleasolinum
gummiterum I, Oiltrichia viscosa subsp. viscosa I, Juniperus navicularis I, Pimpine//a vi//osa I,
Asparagus aphy//us +, Quercus suber +, Cytisus grandiflorus +, Rosmarinus officinalis +, Anemone
Synthetic table 01 Erico umbelletae-U/icetum welwitschianií Capelo, J.C. Costa, Neto & Lousã in palmata +, Scrophularia frutescens +, Ca//una vulgaris r, Thapsia vi//osa r, Sesamoides canescens r,
J.C. Costa, Capelo, Neto, Espírito Santo & Lousã 1997 (Ericion umbe//atae, U/icetalia minoris, Ca//uno- Fritilaria lusitaica subsp. stenophy//a r, Retama monosperma r, Genista triacanthos r, Cistus
Ulicetea) , Iram Costa et ai. 1997, 12 relevees: Charac!eris!ics: Ulex australis subsp. welwitschianus V, psilosepalus r.
Ca//una vulgaris V, Genista triacanthos V, Erica umbe//ata IV, Erica australis subsp. australis IV,
Tuberaria lignosa IV, Erica scoparia IV, Pterospattum tridentatum subsp. tridentatum III , Agrostis cuttisii
III , Thymus vi//osus II, Simethis mattiazzi II, Cistus psilosepalus II, Erica erigena +; companions: Cistus Synthetic table 01 Salvio sclareoidis-Ulicetum densi thymetosum sylvestris Rivas-Martínez,
salviifolius V, Halimium halimifolium V, Lavandula luisieri III, Cistus ladaniter III, Cistus crispus III , Lousã, T.E. Díaz, Fernández-González & J.C. Costa ex Capelo, J.C. Costa, Lousã & Neto 1992
Thymus capite//atrus III, Rosmarinus officinalis III, Oaphne gnidium III, Halimium calycinum III, Lithodora (Serratulo estremadurensis-Thymenion sylvestris, Saturejo-Thymbrion capitatae, Rosmarinetalia
prostra ta subsp. lusitanica III, Pulicaria odora III, Briza maxima II, Quercus lusitanica II, Asphodelus officina//is, Rosmarinetea officinalis), Iram Capelo et ai. 1993, 10 relevees: Charac!eris!ics: U/ex
lusitanicus II, Cladonia mediterranea II, Arrhenatherum album II, Cladonia potentosa II, Juniperus densus V, Thymus zygis subsp. sylvestris V, Salvia sclareoides V, Rosmarinus officinalis, Sideritis
navicularis II, Arbutus unedo II, Stipa gigantea II, Asparagus aphy//us II, Corema album II, Myttus hirsuta varo hittula IV, Stahelina dubia IV, Serratula baetica subsp. lusitanca III, Avenula occidentalis III ,
communis I, Phi//yrea angustitolia ' 1, Stauracanthus genistoides I, Margotia gummifera I, Serratula Phagnalon rupestre III, Fumana thymifolia II, Cistus albidus II, Thymbra capitata II , Micromeria graeca
monardii I, Pistacia lentiscus I, Anemone palmata I, Lavandula sampaiona subsp. lusitanica I, Osyris subsp. micrantha II, Anthy/lis vulneraria subsp. maura II , Serratula estremadurensis I, Battsia aspera I,
alba I, Carex ha//erana I, Oittrichia viscosa +, Piptaterum miliaceum +, Centaurea aspera +, Pimpine//a Iberis procumbens subsp. microcarpa I, Helianthemum origanitolium I, Va//eriana tuberosa I, Cistus x
vi//osa +, Holcus lanatus +, Carpobrotus edulis +, Rubus ulmifolius +, Gladioulus italicus +, Pteridium pulverulentus I, Nepeta mu/tibracteata I; Companions: Brachypodium phoenicoides V, Oaphne gnidium
aquilinum +, Lonicera peryc/imenum subsp. hispanica +, Helichrysum picardi varo virescens +, V, Lavandula luisieri IV, Carex ha//erana IV, Eryngium dilatatum IV, Centaurium erythraea subsp.
Centaurium erythreae +, Erygium dilatatum +. grandiflorum IV, Cistus salviifolius IV, Bupleum rigidum subsp. paniculatum IV, Asparagus aphy//us IV,
Pistacia alentiscus IV, Smi/ax aspera var. aspera IV, Cistus monspe/iensis III, Oacty/is hispanica III ,
 The Landscape and Plant Communities of Serra da Arrábida
Costa, J.C. & aI.
Cistus albidus, t Cistus salviifolius, t Nepeta tuberosa, t Scabiosa atropurpurea varo villosa, +
Quercus coccifera III, Rubia peregrina varo longifolia III, Juniperus turbina ta II I, Rhamnus oleoides III ,
Centaurium erythraea subsp. grandiflorum.
Euphorbia portlantica III, Helichrysum stoechas II, Olea europeae varosylvestris II, Genista tournefortii
II , Phlomis purpurea II, Phillyrea angustifolia II, Lonicera implexa II, Oaucus crinitus II, Hyparrhenia
sinaica II , Reichardia intermedia II, Cistus crispus II, Astragalus lusitanicus II , Convolvulus altheoides II ,
Relevée 01 Iberido mieroearpae-Stipetum offnerií Rivas-Martínez, Lousã, T.E. Díaz, Fernández-
Anagallis monelli varo linifolia, Carlina corymbosa II, t Narcisus bulbocodium subsp. obesus II ,
González & J.C. Costa 1990 (Thero-Brachypodium retusum, Lygeo-Stipetalia, Lygeo-Stipetea): Serra
Sanguisorba minor subsp . spachiana II, Reichardia picrioides II , Phlomis Iychnitis II, Urginea maritima II ,
do Risco, 4 m2, SW, 250 m, Iram Rivas-Martínez et aI. , 1990: Characteristics: 3 Stipa oftnerii, 1
Pulicaria odora II, Stipa gigantea II, Pallenis spinosa II, Brachypodium distachyon II, Tulipa australis I,
Ornithogalum concinnum, 1 Sedum sediforme, 1 Phagnalon saxatile, 1 Gladiolus reuteri, 1
Rhamnus alaternus I, Plantago serraria I, Atractylis gummifera I, Scabiosa atropurpurea I, Cynara
Brachypodium retusum, t Iberis procumbens subsp. microcarpa, t Oactylis hispanica, t Narcissus
cardunculus I, Allium paniculatum I, Santolina rosmarinifolia I, Serapias vomeracea I, Schoenus
calcicola; Companions: 1 Arisarum vulgare varo clusii, 1 Hyacintoides hispanica, t Asphodelus
nigricans I, Cheirolophus sempervirens I, Sanguisorba ancestroides I, Erica scoparia I, Melica minuta
aestivus, t Astragalus lusitanicus, t Rosmarinus ofticinalis, t Fumana laevipes, t Gennaria diphylla, t
subsp. arrecta I, Bituminaria bituminosa I, Phagnalum saxatile I.
Sedum album varo micranthum, t Valantia hispida, t Calendula suftruticosa varoalgarbiensis, t Urginea
maritima, t Galium verrucosum.

Synthetic table 01 Cariei depressae-Hyparrhenietum sinaicae Br. -BI ., P. Silva & Rozeira 1956
corro J. C. Costa, Capelo, Lousã & Esp írito Santo 2002 (Hyparrhenion sinaicae, Hyparrhenetalia hirtae,
Synthetic table 01 Avenulo suleatae-Stipetum giganteae Capelo, J.C. Costa, Lousã & Espírito
Lygeo-Stipetea), Iram Costa et aI. 1994, 20 relevees : Characteristics: Hyparrhenia sinaica V,
Santo in. J.C. Costa, Capelo, Lousã & Esp írito Santo 2002 (Agrostio castellanae-Stipion gigantaea,
Phagnalon saxatile V, Oactylis hispanica V, Reichardia picroides, Convolvulus altheoides V, Asphodelus
Agrostietalia castellanae, Stipo giganteae-Agrostietea castellanae) , Irom Costa et ai. 1992, 9 relevees:
lusitanicus IV, Bituminaria bituminosa III, Erygium dila ta tum III, Thapsia vilosa II, Oaucus crinitus II ,
Characteristics: Stipa gigantea V, Agrostis castellana V, Brachypodium phoenicoides V, Avenula
Salvia sclareoides II, Brachypodium phoenicoides II, Gynadriris sisyrichium II, Micromeria graeca II ,
sulcata subsp. sulca ta IV, Stachys ofticinalis subsp. algeriensis IV, Arrhenatherum album III, Oactylis
Plantago serraria II, Anacamptis pyramidalis I, Lathyrus ocrus I, Carex depressa t,
hispanica III, Oactylis glomerata subsp. lusitanica III, Asphodelus lusitanicus III, Thapsia villosa II I,
Pseudarrhenateurum pallens t; companions: Foeniculum vulgare subsp. pipeperitum V, Urginea
Avenula sulcata subsp. gaditana I, Festuca durandoi I, Holcus annus I; companions : Holcus lanatus V,
maritima V, Oxalis pes-caprae V, Piptaterum mileaceum IV, Reichardia intermedia IV, Asphodelus
Erica scoparia III, Agrostis curtisii III, Thymus villosus II I, Lavandula luisieri III, Carlina corymbosa III ,
fistulosus IV, Salvia verbenaca III, Asparagus albus III, Verbascum sinuatum III , Cynara humilis III ,
Ulex jussiaei III, Quercus lusitanica II I, Anthyllis gerardi II , Asparagus aphyllus II, Brila maxima II ,
Atractylis gummifera III, Lobularia maritima II, Ononis natrix subsp . hispanica II , Paronychia argentea II ,
Lagurus ovatus II, Ulex minor II, Brachypodium sylvaticum II, Euphorbia characias II, Plantago afra I,
Asparagus aphyllus II, Scorpiurus muricatus II, Sedum album II, Brachypodium distachyon II ,
Plantago coronopus I,
Sanguisorba spachiana I, Echium tuberculatum I, Ruta calepensis I, Thymbra capitata I, Vicia sativa I,
Anthyllis vulneraria subsp. maura I, Oelphinum plantagineum I, Campanula rampunculus I, Vicia lutea t,
Silene vulgaris t, Narcissus bulbucodium subsp. obesus.
Synthetic table 01 Corynephoro maeranthero-Arenarietum algarbiense P. Silva & Teles ex Rivas-
Martínez & Izco 2002 (Anthyllido hamosae-Malcolmion lacerae, Malcolmietalia, Tuberarietea guttata e),
Iram Neto, 2002, 28 relevees: Characteristics: Corynephorus macrantherus V, Rumex
Relevée 01 Phlomido Iyehnitidis-Braehypodietum phoenieoidis Br.-BI., P. Silva & Rozeira 1956
bucephalophurus subsp . hispanicus V, Erodium aethiopicum subp. pilosum V, Tuberaria guttata V,
(Brachypodion phoenicoidis, Brachypodietalia phoenicoidis, Festuco-Brometea erectl) , EI Carmen, 50
Linaria spartea V, Hypochaeris glabra V, Logfia gallica V, Silenes cabriflora IV, Polycarpon alsinifolium
m2, NW, 220 m: Characteristcs: 3 Brachypodium phoenicoides, 2 Oactylis hispanica, 2 Eryngium
IV, Astrolinum linum-stellatum III, Micropyrum tenellum II I, Loeflingea baetica varo micrantha III ,
dila ta tum, 1 Mantisalca salmantica, 1 Plantago serraria, 1 Salvia sc/reoides, 1 Asphodelus lusitanicus, 1
Leucojum tricophyllum II, Silene littorea II, Coronilla repanda subsp. repanda II, Aira caryophyllea II ,
Allium roseum, 1 Oaucus crinitus, 1 Anthyllis vulneraria subsp. maura, t Ophrys lutea, t Phlomis
Bril a maxima II, Anthyllis hamosa II, Malcolmia lacera subsp. gracilima II , Ononis baetica II , Silene
Iychnitis, t Lathyrus amphicarpus, t Aceras anthropophorum, t Anacamptis pyramidalis, t Convolvulus
colorata II, Mibora minima II, Jasione montana II, Pimpinella villosa II, Euphorbia exigua II, Arenaria
altheoidis, t Urginea maritima, t Fritilaria lusitanica, t Narcisus bulbocodium subsp. obesus, t Phleum
algarbiense I, Vulpia membranacea I, Ononis broteriana I, Ornithopus isthmocarpus I, Tolpis barbata I,
bertolonii; companions: 1 Cynodon dactylon, 1 Plantago lanceolata, 1 Bupleurum rigidum subsp. Ornithopus pinnatus I, Evax pygmaea subsp. ramosissima t , Campanula lusitanica subsp. matritensis
paniculatum, 1 Rosmarinus ofticinalis, t Carex hallerana, t Campanula rampunculus, t Pulicaria odora, t , lonopsidium acaule t , Lotus castellanus t , Paronychia cymosa t, Arenaria conimbricensis r, Agrostis
t Asparagus aphyllus, t Cynara humilis, t Phalaris caerulescens, t Agrotis stlolonifera, t Anagallis
tenerrima r, Loeflingia laveresiana r, Logfia gallica r, Lathyrus angulatus r; companion: Secio gallicus
arvensis varo linitolia, t Sanguisorba minor subsp. spachiana, t Thymus lygis subsp. sylvestris, t
Costa, J.C. & aI.
The Landscape and Plant Communities 01 Serra da Arrábida

IV, Spergularia purpurea IV, Vulpia alopecurus III, Spergula arvensis III, Juncus capitatus III , Bromus Spergularia australis II, Carlina corymbosa subsp. major II, Armeria pungens subsp. major I,
rigidus III, Avena longiglumis II, Euphorbia portlantica II, Sesamoides canescens II, Centranthus Helichrysum decumbens I; companions : Euphorbia portlantica V, Helianthemum marifolium V,
caleitrapa II, Corynophorus canescens varomaritimus II, Silene ga/lica II, Lupinus angustifolius subsp. Calendula suffruticosa subsp. algarbiensis IV, Sedum sediforme V, Catapodium marinum V, Lobularia
reticulatus II, Papaver setigerum II, Dipecadi serotinum I, Carduus meonanthus I, Raphanus maritima III, Cruciane/la maritima III, Lotus creticus III, Parapholis incurva III, Rosmarinus officinalis III ,
raphanistrum I, Chamaemelum mixtum I, Brassica barrelieri subsp. oxyrrhina I, Arrhenatherum album +, Echium tubeculatum III, Astericus maritimus III, Atriplex halimus II , Brachypodium phoenicoides II ,
Avena barbata +, Anaga/lis arvensis +, Ste/laria media +, Anarrhinum belidifolium +, Anemone palmata Eleaesolinum gummiferum II , Beta maritima II , Asparagus aphy/lus II, Salsola vermiculata II, Pa/lenis
+, Centaurea sphaerocephala +, Silene nicaensis +, Sci/la monophy/lus +, Papa ver rhoeas +, spinosa II, Hordeum marinum II, Erygium dilatatum II, Cistus salviifolius II, Daphe gnidium var. maritima,
Paronychia argentea +, Rumex angiocarpus +, Radiola linoides r, Galium minutulum r. II , Asperula seabra II, Juniperus turbina ta II, Asperula seabra II, Carpobrotus edulis I, Anaga/lis monelli
varo microphy/la I, Thymus zygis subsp. sylvestris I, Mesembryanthemum nodiflorum I, Mucozonia
hispida I, Thapsia vi/losa I.
Relevee 01 Leucanthemo sylvatici·Cheirolophetum sempervirentis J.C. Costa, Ladero, T.E Díaz,
Lousã, Espírito Santo, Vasconcelos, Monteiro & Amor 1993, at Mata do Solitário, 8 m2, SE, 210m:
(Stachydo lusitanicae·Cheirolophenion sempervirentis, Origanion virentis, Melampyro·Holcetalia, References
Trifolio·Geranietea) Characteristics: 4 Cheirolophus sempervirens, 2 Picris spinifera, 1 Teucrim AG UIAR, C., COSTA, J.C., CAPELO, J., AMADO, A., HON RADO, J., EspíRITO SANTO, M.D. &
scorodonia subsp. scorodonia 1 Origanum virens, 1 Clinopodium vulgare subsp. arundanum, 1 Vinca LOUSÃ, M. (2003) - Aditamentos à vegetação de Portu gal continental. Silva Lusit. 11 (1): 101·11 1.
difformis, 1 Calamintha baetica, 1 Sedum forsteranum, + Campanula rampunculus, + Stachys BRAUN·BLANQUET, J., PI NTO DA SILVA, A.R. & ROZEIRA, A. (1956) · Résultats de deux excursions
germanica subsp. lusitanica, Companions: 2 Brachypodium phoenicoides, 1 Geranium purpureum, + géobotanique à travers le Portugal septentrional & moyen II. Chenaies à leuilles caduques
Salvia sclareoides, + Arisarum vulgare var. c1usii, + Bituminaria bituminosa. (Ouereion occidentale) et chenaies à leuilles persistentes (Ouereion faginae) au Portugal. Agron.
Lusit. 18 (3): 167·234
BRAUN·BLANQUET, J., PINTO DA SILVA, A.R. & ROZEIRA, A. (1964)· Résultats de deux excursions
Relevée 01 Si/eno longicilae·Antirrhinietum Iinkianum Ladero, Valle, M.T. Santos, Amor, Espírito géobotanique à trave rs le Portugal septentrional & moyen III . Landes à Cistes et Ericacées (Cisto·
Santo, Lousã & J. C. Costa 1991 (Calendulo lusiotanicae-Antirrhinion linkiani, Phagnalo saxatilis- Lavanduletea et Ca/luno·Ulicetea). Agron. Lusit. 23 (4): 229·313.
Rumicetalia indurati, Phagnalo-Rumicetea induratae), Serra Arrábida near Conve nto, 2 m2, NE, 320 m, CAPELO, J. (1996) • VIII Nota à sintaxonomia das orlas herbáceas Ilorestais do SW da Península
Characteristics: 2 Antirrhinum linkianum, 1 Biscute/la lusitanica, 1 Calendula suffruticosa subsp.
Ibérica. Silva Lusit. 4 (1) : 123·1 25.
lusitanica, 1 Phagnalon saxatile, 1 Sanguisorba multiculmis, 1 Melica minuta + Silene longieilia, + Arabis CAPELO, J. (2003) - Syntaxonomical disposal 01 the Euphorbia pedroi Molero & Rovira communities, a
sadina, + Sedum album varo rnicranthurn; Companions: 1 Biturninaria bituminosa, 1 Hyparrhenia syn·endemism 01 Serra da Arrábida (Portugal) sea·cliffs - Convolvulo fernandesii·Euphorbieturn
sinaica, + Reichardia picrioides, + Sedum sediforme.
pedroi. Silva Lusit. 11 (1): 123·124.
CAPELO, J. & ALMEIDA, A. F. (1993) . Dados sobre a paisagem vegetal do Parque Natural da Serra
da Arrábida: proposta de uma tipologia li tos sociológica. Silva Lusit. 1(2): 217·236.
Relevée 01 Narciso calcicolae·Asplenietum ruta murariae Espírito Santo, Ladero & Lousã 1995 CAPELO , J. & COSTA, J.C. (2002) . Notícia acerca dos carrascais arbóreos da Serra da Arrábida. Silva
(Asplenion petrachae, Asplenetalia petrachae, Asplenietea trichomanis) Serra da Arrábida near
Lusit. 9 (2): 269-271.
antennas, 2m 2, SW, 485 m, Characteristics: 2 Narcissus calcicola, 1 Melica minuta, 1 Asplenium CAPELO, J., COSTA, J.C., EspíRI TO SANTO, M.D. & LOUSÃ, M (1993) - As comunidades
petrachae, + Phagnalon rupestre, + Asplenium ceterach, + Asplenium trichomanes, + Sanguisorba camefíticas dos calcários do Centro-Oeste Português (Serratulo estremadurensis-Thymenion
ancistroides; companions: + Arabis sadina, + Calendula suffruticosa subsp. lusitanica. sylvestris, suball. nova). ln Guia Geobotânico das XIII Jornadas de Fitossociologia; 99-11 8. I. S.
Agron omia. Lisboa.
CAPELO, J., COSTA, J.C., LOUSÃ, M, MESQU ITA, S. (2002) - A aliança Ouercion fruticosae
Synthetic table 01 Helianthemo stoechadifoliae·Limonietum virgatae J. C. Costa, Lousã & Capelo in Rothmaler 1954 em. Rivas·Martínez, Lousã, Díaz, Fernández-Gonzalez & J.C. Costa 1990.
Aguiar, J.C. Costa, Capelo, Amado, Honrado, Espírito Santo & Lousã 2003 (Crithmo-Daucion halophili, Ouercetea 3: 99-110.
Crithmo-Limonietalia, Crithmo-Limonietea), Irom Costa et ai. 1998, 9 relevees: Characteristics: CASTROVIEJO, S. et ai. (ed) (1986·2003) - Flora Iberica. 1, 2, 3, 4, 5, 6, 7, 8, 19, 14. Real Jardín BoI.
Limoniurn virgatum V, Helianthemum apeninum subsp. stoechadifolia V, Daucus halophilus V, Dactylis Mad rid. Madrid.
marina V, Crithmum maritimum V, Plantago coronopus subsp. occidentalis IV, Frankenia laevis IV,
Costa, J.C. & aI. The Landscape and Plant Communities of Serra da Arrábida

COSTA, J.C., AGUIAR, C., CAPELO, J., LOUSÃ, M & NETO, C. (1998) - Biogeografia de Portugal GÉHU, J.M . & S. RIVAS-MARTíNEZ (1981) - Notions fondamentales de phytosociologie. ln
Continental. Quercetea O: 3-56. Syntaxonomie. 5-33. Ed. J.Cramer. Vadu z.
COSTA, J.C., CAPELO, J. EspíRITO SANTO, M.D. & LOUSÃ, M. (2001) - Corrección nomenclatural LADERO, M., VALLE, C.J., SANTOS, M.T. , AMOR, A., EspíRITO-SANTO, M.D. , LOUSÃ, M. &
de los sintaxones basados en Hyparrhenia hirta dei sector Divisorio portugués. Lazaroa 22: 135- COSTA, J.C. (1991) - Sobre la végetación y flora rupicola de las intercalaciones calcare as de los
136. sectores Divisorio portugés y Bei rense litoral. Candollea 46 (1): 53-59.
COSTA, J. C. , CAPELO, J. EspíRITO SANTO, M.D. & LOUSÃ, M. (2002) - Aditamentos à vegetação LOUSÃ, M., COSTA, J.C. , CAPELO, J., PINTO GOMES, C. & C. NETO, C. (1999) - The vegetation of
do Sector Divisório Português. Silva Lusit. 10 (1): 119-128. the stretch between Faro and Évora. ln Rivas-Martínez, S., J. Loidi, M. Costa, T.E. Díaz & A. Penas
COSTA, J.C., CAPELO, J. & LOUSÃ, M. (1994) - Os bosques de zambujeiro (Olea europaea L. varo (ed) - Iter Ibericum A.o. MIM. (Excursus geobotanicus per Hispaniarn et Lusitaniam, ante XLII
sylvestris Miller): vegetação potencial dos vertissolos das áreas termomediterrânicas da Syposium Sociatatis Internationalis Scientiae Vegetationis Bilbao mense lulio celebrandu dicti
Estremadura portuguesa. Anais do Inst. Sup. Agron. 44 (2): 497-513. Anm). Itinera Geobot. 13: 149-168.
COSTA, J.C. , CAPELO, J., LOUSÃ, M. & AGUIAR, C. (1994) - Communautées de Juniperus au NETO, C. (1994) - Notas sobre a flora e a vegetação do Cabo EspicheI. Finesterra. 55/56: 201-214.
Portugal. Colloques Phytosoc. 22: 499-526. NETO, C. (2002) - A flora e a Vegetação do Superdistrito Sadense (Portugal). Guineana 8: 1-269.
COSTA, J.C., CAPELO, J., LOUSÃ, M. & EspíRITO SANTO, M.D. (1996) - Asparago aphy/li-Querceto Universidad dei País Vasco.
suberis sigmetum - a new coark-oak woodlands vegetation series of central-west Portugal. A case- RIVAS-MARTíNEZ, S. (1976) - Sinfitosociologia, una nueva metodologia para el estudio dei paisaje
study of an integrated approach to the forest syntaxonomy. Livro de Resumos do I Congreso de la vegetal. Anales Inst. Bot. Cavanilles 30: 69-87.
Federación Internacional de Fitosociologia: 66. Oviedo. RIVAS-MARTíNEZ, S., DíAZ, T.E., FERNANDEZ-GONZÁLEZ, F., IZCO, J., LOIDI, J., LOUSÃ, M. &
COSTA, J.C., CAPELO, J., LOUSÃ, M. & EspíRITO SANTO, M.D. (1997) - Sintaxonomia da vegetação PENAS, A. (2002) - Vascular Plant communities of Spain and Portugal. Addenda to Sintaxonomical
halocasmofítica das arribas marítimas portuguesas (Crithmo-Staticetea Br.-BI. 1947) Itinera checklist of 2001 (part I, II). Itinera Geobot. 15 (1 , 2): 5-922.
Geobot. 11: 227 -247. RIVAS-MARTíNEZ, S., FERNÁNDEZ-GONZÁLEZ, F. & LOIDI, J. (1999) - Checklist of plant
COSTA J. C., CAPELO, J., LOUSÃ, M. & EspíRITO SANTO, M.D. (2002)- Os sobreirais do Divisório communities of Iberian Peninsula, Balearic and Canary islands to suballiance levei. Itinera Geobot.
Português: Asparago aphy/li-Quercetum suberis. Quercetea 3: 81-98. 13: 353-451.
COSTA, J.C., CAPELO, J., LOUSÃ, M. & EspíRITO SANTO, M.D. (2004) - As comunidades de RIVAS-MARTíNEZ, S., FERNANDEZ-GONZÁLEZ, F. , LOIDI, J., LOUSÃ, M. & PENAS, A. (2001)
Asparago albi-Rhamnion oleoidis Rivas Goday ex Rivas-Martínez 1975 do Divisório Português. Sintaxonomical checklist of vascular commun ities of Spain and Portugal to association levei. Itinera
Quercetea 4: 31-43. Geobot. 14: 5-341
COSTA, J.C., CAPELO, J., NETO, C. , EspíRITO SANTO, M.D. & LOUSÃ, M. (1997) - Notas RIVAS-MARTíNEZ, S., LOIDI , J., COSTA, M. , DíAZ, T.E . & PENAS, A. (ed .) (1999) - Iter Ibericum A.o.
fitosociológicas sobre os tojais do Centro e Sul de Portugal. Silva Lusit. 5 (2): 275-282. MIM. (Excursus geobotanicus per Hispaniam et Lusitaniam, ante XLII Syposium Sociatatis
COSTA, J.C., LADERO, M., DíAZ, TE , LOUSÃ, M., EspíRITO SANTO, M.D., VASCONCELOS, T., Internationalis Scientiae Vegetationis Bilbao mense lulio celebrandu dicti Anm). Itinera Geobot. 13:
MONTEIRO, A. & AMOR, A. (1993) - Guia Geobotânico da Excursão das XIII Jornadas de 5-347.
Fitossociologia: 1- 98. I. S. Ag ronomia. Lisboa. RIVAS-MARTíNEZ, S., LOUSÃ, M., DíAZ T.E., FERNANDEZ-GONZÁLEZ, F. &. COSTA, J.C. (1990)-
COSTA, J.C., LOPES , M.C, CAPELO, J. & LOUSÃ, M. (2001) - Sintaxonomia das comunidades de La vegetación dei sur de Portugal (Sado, Alentejo y Algarve) Itinera Geobot. 3: 5- 126.
Prunus lusitanica L. subsp. lusitanica no ocidente da Península Ibérica. Silva Lusit. 8 (2): 253-263. RIVAS-MARTíNEZ, S., A. PENAS & T.E. DíAZ (2004) - Biogeographic and bioclimatic maps of Europe.
COSTA; J.C., LOUSÃ, M. & EspíRITO SANTO, M.D. (1997) - Vegetação do Parque Natural da Ria Serviços Cartográficos da Universidad de Léon.
Formosa. Studia Bot. 15: 69-157. SAMPAIO, G. (1947) - Flora Portuguesa. Ed 2. Imprensa Moderna. Porto.
EHRENDDORFER, F. (1994) - Geobotânica. ln Strasbourg et aI. Tratado de Botánica: 871-978. TUTIN et ai. (1964-1980) - Flora Europaea (I-V). Cambridge University Press.
Ediciones Omega. Barcelona. VALDÉS, B., S. TALAVERA & E. GALlANO (1987) - Flora vascular de Andalucía occidental. Vol. 1-3.
EspíRITO SANTO, M.D ., LADERO, M. & LOUSÃ, M. (1996) - Comunidades rupícolas do Parque Ketres. ed. Barcelona.
Natural das Serras de Aire e Candeei ros . Studia Bot. 14: 13-22.
FRANCO, J.A. (1971-1984) - Nova Flora de Portugal (Continente e Açores). voll e II. Lisboa.
FRANCO, J.A. & M.L. ROCHA AFONSO (1994 - 1998 -2003) - Nova Flora de Portugal (Continente e
Açores). vollll , fasc. 1,2, 3. Escolar Editora. Lisboa.
Quercetea 7: 27-41, 2005
ALFA, Lisboa. Portugal

Natural Succession in Southwestern Portuguese Limy Soils: After Fire

Dynamics (Arrábida Natural Park) and Primary Succession in Sand
Dunes (Tip of Peninsula of Troia)

Otília Correia' & Manuel João Pinto*

South of Lisbon , crossing the bridge over the Tagus River, we reach the "Setúbal Peninsula". It is a
large area (500 km2) that extends from the Tagus River to the Arrábida Chain and bordered by the
Atlantic Ocean on its western
boundary (Fig. 1). Low altitudes
(50-60 m), smooth relief and
sandy landscapes characterise
this region (1). Its proximity to the
city of Lisbon is responsible for an
intense human occupation and
degradation of natural landscape,
preserved only in restricted areas
expanding southwards.

The "Serra da Arrábida" Natural

Park, is located in Arrábida Chain
(2) an elongated mountain, located
on the southern border of the
Setúbal Peninsula, about 40 km
south of Lisbon (from 38º 27' to
38º 30' N latitude and 8º 55' to 9º
02' W longitude). The Península
de Tróia region corresponds to the
estuarine area of the Sado river
(3) (Fig.1).
_ ===_
1r) o 10
-_-l
<O Km

Figure 1. Geographical situation and localities to be

visited

• Faculdade de Ciê ncias da Universidade de Lisboa. Campo Grande. 1749-016. Lisboa. Portugal.
odogato@fc.ul.pt
Correia, O. & Pinto, M.
Natural Sueeession in Southwestern Portuguese Limy Soils

PART 1- Atter tire dynamics (Arrábida Natural Park) ln opposition, the Chain's low lands are predominantly 01 marly clay nature or aeeumulation, leading to
Otília Correia
a very different type 01 soils. The low erosion rates allow the development 01 thick brown soils, with
Universidade de Lisboa, Faculdade de Ciências, Centro de Ecologia e Biologia Vegetal. Campo organic matter accumu lation. Therelore, wate r and nutrients retention is improved, providing conditions
Grande, 1749-016 Lisboa. odgato@ fc.ulpt lor the growth 01 abundant vegetation. We see leached brown soil s, 01 caleic and neutral to alkaline
nature (pH=6.5 to 7.5), corresponding to Calcie Chromic CAMBISOLS.

Vegetation and climate

I. INTRODUCTION
Climatic data are only available Irom meteorological stations at Sesimbra and Setúbal. The climate is
Geomorphology and soils
clearly Mediterranean, with the dry season occurring in the summer and lasting about 4 months (Fig.2).
The Arrábida Chain (2) is over 30 km long and 5 km broad, extending Irom Espichei Cape, on the
The Emberger climagram enables the classilication 01 Mediterranean bioclimates 01 different sites in
West, to Palmela castle, on the Easl. Clearly detached Irom the surrounding areas, it is a
Arrábida (according with the data obtained by Albuquerque, 1964), as subhumid and humid bioclimates
geomorphologic unit, with the ocean on the South and West (O m) and the sandy plain (previously
with warm and hot winters, except lor Formosinho (500m elevation, 1330mm precipitation, Fig .2).
described) on the North and East (under 100m). It reaches its maximum height at Formosinho (501 m)
and there are several tops over 300m. It corresponds to a well-delimited area 01 Mesozoie (200 to 100 SetúbGl (35m) 16' C 746,8 nun A
(30) 120
million years old) limestone roeks, uplilted Irom the more recent surrounding terrains. The shape 01 this
E-W elongated chain is the result 01 tectonic movements active during the alpine compression, about 5 100
million years ago, originating a narrow lolded and upl ilted structure resulting in several anticlinal and
sinclinal systems. The Formosinho/Sol itário lormation is the more important one, with an ENE-WSW 40 80

~
orientation. 35

~y-1?
30
U
Its complex geomorphology resulted in a strong reliel with steep slopes (nearly hall 01 the area has ~~ 25
I

slopes between 50 and 70%). Due to the strong reliel, true soils proliles are almost absent lor most 01 20 ........ . 40

. \8<
. . . . .....
. .. . . . . . .
the Serra. 15

10 20
The large majority 01 rocks that build up the "Serra da Arrábida", are limestone 01 dillerent kinds, with
variable chalk and clay contents. Locally, some sands and clays occupy an important area. Broadly, it is
J F M A M J J A S o N D
possible to correlate lithology and re liel, stating that high places and steep slopes correspond to hard IVlontllS
limestone, and low regions with smooth slopes correspond to detrital deposits. As a consequence, B
Q --~~
dillerent kinds 01 soil are expected to develop in these two opposite situations, as we will see. Fonnosinho
240
The crystalline limestone, that lithologieally and morphologically delines the Arrábida Chain, resis ts
strongly to physical erosion. The main process 01 landscape evolution is, therelore, by ehemical 200
Pr~-AtÚ.IJ.tU:o

dissolution 01 calcium carbonate, thus developing karstic leatures. The result is an insoluble residue 01 C o r1i.c-.
tU •
oxidized clays with strong red colours - the "terra rossa ". 160 Carrrten

The high places oceupied by the limestone and the associated steep slopes lavou r an intense erosion 120 H1J.IlLiol
~ • Portirilio
01 residual soil, inhibiting the development 01 thick soils, suitable lor vegetation colonization. The
capacity 01 water retention by these soil s is very low, and the rainwater rapidly inliltrates through the 80

dissolution cracks 01 limestone, or runs down the slopes into the valleys or sea. Therelore, we see red
-3 Cold CooI 3 Tempente 7 Hot
"rocky" soils with incipient brown A horizons, alkaline due to their high caleium content (pH=7.5 to 8.5). o 11 T-'C
m
These characteristics correspond to the classilication as Calcic Rodochromic LUVISOLS . Figure 2. A - Ombrothermic diagram lor Setúbal. B - Climatogram showing the Emberger pluviometric
quotient (Q) and the mean minimum temperature 01 the coldest month (m) lor the Mediterranean
climate: location 01 several the Arrábida sites.
 -
Correia, O. & Pinto, M.
Natural Succession in Southwestern Portuguese Limy Soils

As a consequence, diflerent microclimates and soils generate conditions lor the development 01 angustifolia and Rosmarinus officinalis. On the northern slopes 01 Arrábida, a climax lormation with
dlflerent types 01 vegetatlon. The natural vegetation belongs to querei communities where lorests 01 o.
Quercus faginea, coccifera and Phillyrea latifolia is presen!.
Quercu~ faginea ssp. broteroi and Quercus coccifera can be lound in the valle~s and mid-slope
depresslons .. The area covered by these lorests is much reduced due to the strong human impact for Climax lormations can also be lound in the valleys with deeper soil proli les, especially on their north
many centunes (slnce the Arab occupation) and to the Irequent occurrence 01 lire. Kermes oak lacing slopes (Mata Coberta, Mata do Solitário) . These lorests are dominated by Quercus faginea, and
(Quercus coccifera) lormations considered as degradations 01 the lorest, cover most 01 the area in include Acer monspessulanum, Phillyrea latifolia, o.coccifera, Arbutus unedo, Pistacia lentiscus,
patches with a maximum height 011 .5 m. Viburnun tinus, Smilax aspera, etc. At the base 01 these slopes nearly pure lormations 01 Acer can be
lound.
The continuous and strong action olli re and grazing also caused some regression in the vegetation,
resultlng ln the Irequent presence 01 dwari shrubs, namely Irom Cistaceae and Labiatae species. Usually growing as shrubs, some 01 these species (O. coceifera, Phillyrea latifolia, Pistacia lentiscus)
develop here as several meters high (12-15m) trees. For this reason, Chodat (1909) was the lirst to
The vegetation prolile along a North~South transect shows diflerent physiognomic vegetation types, consider that Arrábida is one 01 the last places in the world where remains 01 a tertiary lorest can still be
ranglng Irom ploneer ruplcolous lormatlons to the residual patches 01 the climax deciduous lorest (Fig 10und.On the remaining sites, sh rub lormations are present - maquis and garrigue - wilh Quercus
~. .
coccifera, Phillyrea angustifolia, Pistacia lentiscus, Rosrnarinus officinalis, Arbutus une do, Erica
The habitats and vegetation types encountered along this North/South transect can assume a ârborea, Lavandula luisieri and several Cistus species . Juniperus phoenicea is ollen present, especially
seemingly unlimited number 01 variations depending on local conditions and human land use histories on south-Iacing slopes.
malnly the time since lire. The combination 01 great many habitats (topography, elevation, slope, etc) The mosl important impacts wilhin the area are wildlires and quarry activities: the largest one is at
wlth a large number 01 posslble pathways and stages 01 degradation and regeneration gives to this Outão, in the eastern pari 01 the Serra, with ils concrele lactory (SECIL). ln the last lew years marble
landscape a particular mosalc 01 habitats.
quarries have developed in lhe western pari 01 lhe Serra, not lar Irom Sesimbra.

Strategy types and regeneration after fire

Mxed sclerophyll
deciduous Mxed sc lerophyll cpen
'MXldland ln ali Mediterranean-climate regions evergreen trees and shrubs are the predominant lile lorms
N~ presen!. Broadly speaking, in the Mediterranean basin, plant traits regarding drought resistance can be
B C 50
.... s O
synthesized in a lunctional classilication : (i) drought semi-deciduous and (ii) evergreen sclerophylls
o m species, considered to be drought toleran!. These species difler in several morphological and
physiological adaptalions to the water and nutrient constraints 01 the Mediterranean ecosystems. ln
evergreen sclerophylls, leaves persist through periods 01 unlavourable weather and soil moisture, and
drought deciduous species or semi-deciduous species, which are considered drought avoiders,
500 m drop leaves during the unlavourable dry period. During this period , the semi-deciduous shrubs retain
o a small tull 01 apical leaves, which may difler morphologically and physiologically Irom winter/spring
Scree and . Ulhosoi ls
Fissu
~~~~7~m I-Umic rich lerra LilhosoiJs terra
Brown
hurnic rie
Lithosoils
Humc rie Ulhosoils terra
leaves, and which can rapidly expand when soil moisture becomes available.
soi ls terra ross

Hurni d Sub-hulllid HUlllid Sub-hulllicl

Generally, a given plant community may be dominaled by any one 01 the above types. The resulting
community is generally related to lhe availability 01 moisture in the habita!. The moistest sites are
dominated by evergreen trees, lollowed by evergreen shrubs and then drought deciduous shrubs on
increasingly drier sites.
Figure 3. Vegetation prolile along a North-South transect (adapted Iram Catarino el al.1982
Ali these lile lorms show a range 01 activity patterns which are seasonally related to their rooting
depth. Evergreen sh rubs are generally more deeply rooted than deciduous species , and tend to occupy
more mesic sites wilhin regions 01 Medilerranean climale. Droughl deciduous shrubs in Medilerranean
On the southern slopes graws open woodland Irom the Oleo-Ceratonion. The dominant species are syslems are characlerized by lower leal mass per area (g/cm 2), higher leal nilrogen conlenls, lower leal
Olea europaea var. sylvestris, Ceratonia siliqua, Juniperus phoenicea, Pistacia lentiscus, Phi//yrea area index and shallower rools Ihan evergreen sclerophylls, bul a larger rootlshool ralio. From a
physiological poinl 01 view, Ihey exhibit maximum rales 01 assimilalion and leal conduclance roughly
Correia, o. & Pinto, M. Natural Succession in Southwestern Portuguese Limy Soils

doubling those of evergreen, but less ability to regulate water loss through stomatal closure. This results like Smílax aspera, Rubus ulmifolius and Lonicera implexa behave as resprouters. The initial increase in
in higher productivity in drought deciduous species during favourable periods for growth and cover is mainly due to resprouters.
metabol ism, whereas evergreens maintain a decreasing but positive carbon balance through the ln the first spring alter fire there is usually a flush of herbaceous plants which arise from soil-stored
drought. seeds, dormant bulbs or other underground parts. ln Spring, these areas were covered by numerous
Plant species in Mediterranean-type ecosystems regenerate alter fire either by seed germination or by geophytes with underground regenerative and/or reserve organs (bulbs, rhizomes, and tubers), which
resprouting fram below-g round organs . These regeneration strategies are closely tied with physiological avoid fire to a certain extent, like Narcissus bulbocodium (ANNEX V, Directive 92/43 CE E, 1992), AI/ium
and morphological attributes. roseum, Hyacinthoides hispanica, Anemone palma ta, Arisarum vulgare, Asphodelus spp. and some
orchidaceae and gramineae. The first two species sprout and flower in great abundance and form a
Fires are one of the major disturbances shaping Mediterranean plant communities. Wildfires in carpet of colloured flowers alter fire. We can also find some annual species - anthracophytes, which
Portugal are responsible for the largest portion of forest destruction in the country. The Mediterranean colonize bare areas alter fire; these are olten annual nitrophytes.
type climate, with long, warm and dry summers, and the accumulation of flammable vegetation, also
favour the natural wildfires. The numerous adaptations to fire shown by plants dominating these The obligate seeders were killed, and since they propagate only by seeds and fire promotes their
ecosystems suggest that fire has been a selective force for a long time. germination, a large number of small seedlings were present. These species belong to Cistaceae such
as Cistus albidus, C. monspeliensis, C. crispus and C. salvifolius and Labiateae like Rosmarinus
The Mediterranean ecosystems are well adapted to recurrent fires , for two reasons. First, many officinal/is, Lavandula luisieri, Thymus mastichina, Phlomis purpurea and others.
shrubby species resprout fram underground organs like stumps, and others from rhizomes, and also
seeds are produced at an early age. These seeds can remain dormant in the litter and in the soil for ln these areas, one of the dominant species before fire, Juniperus phoenicea, is highly susceptible to
very long periods. Second, the species have developed characteristics favouring their flammability, and fire, since his propagation is inhibited by it. This species does not resprout and the seeds are destroyed
they depend on the occurrence of fire for perpetuation and for optimal growth. by fire (Clemente et ai. 1996, Clemente 2002). ln this site, we can also find some endemic and
protected species, like Arabis sadina and Iberis procumbens subsp.microcarpa (ANNEX II, Directive
The woody shrubs have been grouped into two categories, according to their mechanism of 92/43 CEE, 1992).
regeneration alter fire: (i) obligate resprouters and (ii) obligate seeders. The evergreen
sclerophyllous shrubs were classified as obligate resprouters, while the deciduous and semi-deciduous 2. Mata Coberta (Mixed Scleropphyll Woodland/ Mediterranean Forest)
species were included in the obligate seeders group since they do not resprout after fire and regenerate The vegetation shows higher structural complexity and biomass on the valley with deep soil profiles.
only through seed germination. The species included in each group show different demographic
This woodland is highly diverse in its architecture, appearance and woody species composition (C-
patterns over the post-fire succession: population density of resprouters is generally similar before and
Fig.3). We can reach this woodland following a narrow 1-2 Km road surrounded by adense
alter fire, whereas that of obligate seeders may suffer large fluctuations depending on the fire regime. sclerophyllous maquis. ln this path of low and high maquis vegetation, the sclerophyllous and semi-
ln this field trip we will look at some of the major community types encountered in Arrábida due to deciduous species present are the same observed in the first site. ln addition, other subshrubs like
different abiotic factors and time since fire. We will go from a community recently burned by a wildfi re (in Coronil/a valentine, Cheirolophus sempervirens, Bupleurum fruticosum. Phagnalon saxatile,
July 2004) to adense woodland (forest) in protected north valleys- pre-cl imax vegetation, more than Helichrysum stoechas, and Osyris quadripartita are found. The road gently descends towards Mata
100 years without fire. Coberta. The elevation is around 340m and the slope ranges from 0-5%. Here, in the protected valley
under mesic and more humid conditions, and in the absence of anthropogenic factors, we find a mixed
We can consider a spatial gradient of recovered vegetation patches (recovered time since last fire) sclerophyllous forest dominated by deciduous and sclerophyllous trees - Mata Coberta. This mixed
overlaid by an aridity gradient from drie r places to mesic ones. sclerophyllous Mediterranean forest, according to its structure, is assumed to be in a pre-climax state
1. Mixed sclerophyll scrub with a recent fire (2004) (Catar~no et ai. 1982). This forest has the sam e composition as high maquis, but better developed
canoples with clear distinction of trunk and canopy, representing older stages of high maquis
The dense cover of maquis vegetation (corresponding to letters D and E - Fig.3) was destroyed by an formations. This condition has been achieved due to the absence of fire for several decades in these
intense fire in July 2004. Regeneration of the plant community started rapidly alter fire. areas. This forest has a very complex structure, with trees, shrubs, lianas and herbs. Mediterranean
ln these stands we can observe the evergreen sclerophyllous species that regrow strongly alter fire, woodland is highly diverse in the growth forms, morphology, physiology and phenology. ln these
regenerating vegetatively from underground buds • obligate resprouters, like Quercus coccifera, woodlands, we can find a mixture of trees and shrubs from deciduous species.
Phil/yrea latifolia, Phil/yrea angustifolia, Arbutus unedo, Myrtus communis, PistaGia lentiscus, Erica
arborea, Olea europaea, Rhamnus Iycioides subsp . o/eoides and Oaphne gnidium. Also some lianas
Correia, O. & Pinto, M.
Natural Succession in Southwestern Portuguese Limy Soils
Several sclerophyllous species like Quercus coccifera, Phillyrea latifolia, Arbutus unedo, Pistacia
lentiscus and others can attain tree dimensions, which is apparently not common in other Mediterranean
Part 11- biological traits in natural succession along sand dune
countries. Quercus faginea subsp. broteroi is the dominant tree. Acer monspessulanum can be found in gradient in the Península of Tróia.
this forest where relief and aspect provide enough humidity and low radiation. The demographic studies Manuel J. Pinto
performed in this forest showed interesting differences in the population structures of the main species. Botanic Garden. National Museum of Natural History. University of Lisbon. R. da Escola Politécnica,
While Phil/yrea latifolia, Quercus coccifera and other sclerophylls present much more equilibrated
54. 1250-102 Lisbon. mjpinto @fc.ulpt
populations, with higher numbers of young oak plants, presents an old population without regeneration
(Catarino et ai. 1982), and Acer monspessulanum presents an invading population with many seedlings
and saplings and few mature individuais. ln this dense forest, sh rubs of obligate seeders are not Introduetion
present.

ln the understory we can find some perennial herbs, like Paeonia broteroi, Hyacinthoides hispanica,
Iris foetidissima, Arum italicum, Arisarum vulgare and Ruscus aculeatus (classified for legal protection The Peninsula of Tróia is a sandy territory,
under ANNEX V, Directive 92/43 CE E, 1992). Some lianas like Smilax aspera and Rubus ulmifolius are approximately 16 km long per 1 km wide,
also very abundant. stretching between the estuary of Sado's river
and the Atlantic oceano It was progressively
Certain species regionally rare today, are found in this protected site; this is the case of Laurus nobilis, formed along different geologic periods. Several
which is highly susceptible to fire .
core areas age more than 2000 years and
3. Near EI Carmen site (Mixed Seleropphyll serub-14 years sinee fire) possibly were part of ancient scattered bodies of
a complex system of barrier islands. Other parts
Following the sam e road , we leave the "Mata" and arrive to a high dense maquis. Elevation ranges
of the peninsula were formed recently. A debate
from 250-300m and the slope from 16 to 25%. ln Sepiember 1991, by the end of a relatively dry season,
about the effect of the 1755 tsunami has focused
a wildfire burned about 200 ha of maquis and garrigue. This fire was severe and consumed ali the
the origin of an important dune ridge, that
aboveground biomass and surface organic material to the edge of the Mata Coberta (Clemente et aI. longitudinally cross the northern part of the
1996). Vegetation recovery was rapid and attained 100%. ln this stand , co-occurring species from the territory, differentiating two morphological
two groups - obligate resprouters and obligate seeders - were observed with high cover.
contrasting subareas. These areas are different
According to most definitions, Mediterranean communities would be considered resilient to fires. regarding sand ridges orientation and the
Following such a perturbation, community structure is altered only briefly and, under most burning associated wind regimes that in specific geologic
regimes, the species composition at a given time alter fire does not change appreciably from one fire periods were responsible for their respective
cycle to the next. The evergreen sclerophyllous shrub component re-establishes itself rapidly alter fire, formation. Tróia had evolved to a heterogeneous
by resprouting from belowground vegetative parts and/or by recruitment of seedlings from soil-stored continuous territory linked to the main land,
seeds. As the relative abundances of shrub species return to pre-fire leveis, the species which recruit offering different migratory opportunities for
1 km
large numbers of seedlings are thinned out. animais and plants.

The different mode of reestablishment produces very different demographic patterns alter fire. Figure 1. Visit area in the Northern part of
Obligate seeders survive only as a dormant seed pool in the soil. Recruitment is restricted to the first
the Peninsula of Tróia, showing main field During the last forty years, an important part of
year alter fire, producing a large even-aged cohort, which suffers extreme mortality. ln contrast, obligate trip itinerary. the territory near the northern tip of Adoxe was
resprouters show little demographic change alter fire and regain their original cover more rapidly than
formed succeeding shoreline migration.
obligate seeders (Clemente et aI. 1996, Clemente 2002). ln this site we can observe that the population Shoreline retreat follow the variation in the magnitude and position of outer sand banks deposits, known
of obligate seeders, although very dense and with a high cover, begins to experience some mortality. by Banco do Cambalhão, accumulated outside the harbour and consolidating a renewed main
dlscharge channel. As a direct consequence of this phenomenon , the peninsula is now migrating and
enl . I .
arglng ast towards westwards. Dunng the summer 01 2004 was already possible to cross the recent
Correia, O. & Pinto, M. Natural Succession in Southwestern Portuguese Limy Soils

outer sand bank, from the pre-existent tip of Adoxe to the nearest lighthouse 2 km farther West, walking The aim of this field trip is to observe and study local and subregional patlerns of combination of
by foot, at low tide. biological traits that irrespective to taxonomic properties that characterise plant communities, are most
related to residual component due to species turnover along pH gradients.
Following the shoreline migration towards Northwest and West, a chronological sequence of sand
dune ridges parai lei to the shore has been emerged. Youngest dunes are located towards both
Table 1. Four biological traits and correspondent categories
shorelines, forming a larger sector ai ong the oceanic side. Older dune bodies occur in the central part
The abundance of four
upon the main longitudinal geographic axe, developing transversely towards Northeasl. As a global
• height of the crown centroid [h] : biological traits due to
result of this chronological decrease in sand deposition and thus exposure to soil organic complexation
a) 0-10 b)10-30 c)30-70 d) 70-130 e) 130-210 f»130 cm plant aggregation and
and cation leaching processes, is the remarkable edaphic gradient that transversely crosses the
• crown diameter [e]: community organisation
a) 0-10 b) 10-20 c)20-40 d) 40-80 e) 80-160 f) >160 cm is focused in this visit, as
ilmol /9 %(Olg . Carbon)x1 00 well it is addressed
450 ,~'----------------------------------------------~- 60
500
70
t • life span [Is]:
a) <=3 years b) 3-8 years c) 8-15 years d) >15-24 e) >24 years
causative functional
400 ~'--------------------------~--------------~--~ 50
350 '
300 '
250'
7 ,'I
7
/
1
40
• leaf concentration on the canopy [Ie]:
length proportion ai ong the branches, from distal to proximal
effects of four biological
traits: plant height,

~~~ l,. \~. ~~

crown diameter, life
/ position, that are more densely covered by leaves
span and leaf
1~~ ,~ I I Lo a) 1/4-1/2 b) 1/2-111 c) 1/1-2/1 d) 2/1-4/1 e) >4/1
concentration on
1 3 9 10 11 12 13 14 15 16 17 18 19 20 branches. Vegetation is
Atlantic shore ( > Estuary shore
composed by mosaics of non-thorny dwarf sclerophyllous shrubs rich in Cistaceae species. Foredunes
Northeast are covered by typical dwarf and flexible graminoid vegetation strongly related to coastal and wind
8Juthwest dynamics. •

Altho.ugh the visit area is illustrative of the chronosequence in the sand dune system, data used in the
... Organic carbon l ecologlcal analysis mentioned in this trip guide, come from a wider analysis that gathered 290
vegetation and soil samples scatlered over 5.5 km sand dune Northern sector of the peninsula. Some
sample locations are indicated in the figures 4c and 4d.
Figure 2. Variability ai ong a coast-to-coast transect in the amount of cation Ca+ 2and percentage of
orqanic carbon in soil samples collected at 10/15cm below orqanic horizon
peninsula, consistent with the chronosequence gradient of dune ridges. This gradient is visualised by Combination of biological traits
soil pH (fig. 4a), that starting in central acid and old core areas is progressively increasing towards the
shoreline. It is based on the increase in organic carbon consistently with the gradient of decalcification. How much informative is the selected set of biological traits?
Older dunes have much less calcium carbonate and much larger amount of organic carbon in soil, while .Biological traits were categorised in several modalities presented in table 1. When regressing the
the reverse characterises younger dunes. Floristic gradients follow this main effect in such a way that blologlcal traits data table onto the residual component not explained by species turnover along pH
the relationship found when a global floristic data table of 55 perennial plant species is regressed onto gradlents, the R2 for the first factorial axe achieves 0.21, P =0.0009 (Monte Cario test). This result
the soil pH, R2 for the first factorial axe ranges 0.17, p=O.OOOOO (Monte Cario test). When applying suggest global moderate causative influence, being particularly high concerning leaf concentration on
multivariate factorial analysis and the noise of this floristic data table is c1eaned out by first component branches which accounts for a levei of correlation ratio of r=0.43. Other trait groups accounted for
extraction, which then is regressed onto the soil pH , the amount of variability explained raises to respectively 0.17, 029 and 0.12. Crown diameter is the second most influential trait (r=0.29). .
R2=0.78, p=0.0001. This result states the great importance of soil properties on plant species turnover,
and its influence aid to predict species composition on plant communities at each step of the gradient
(Pinto & Andrade, submitted). The regressive residual component not explained by soil pH is prone to
further investigation.
 Natural Succession in Southwestern Portuguese Limy Soils
Correia, O. & Pinto, M.

Table 2. Correspondance between cluster represented by the formula h-e-Is-Ie. 80th general
When analysing residuais alter studying the influence of soil pH on the floristic table, and then
considering 6 magnitude classes numbers indicated in map d) of figure 5 and formulas depict a rank gradation for each trait.
combinatorial formulas of trait categories This gradation is characterised in the first set, by
of that component, the
contribution for each one of that an increase in crown diameter, life span and the
classes by each trait category is ability to concentrate leaves in the outer part of the
indicated in figure 3. Cluster Main formula of canopy. ln the formula h-e-Is-Ie the gradation is
number combined traits characterised by rank increase in ali categories,
but the crown diameter only is relevant when
4 c1 height and life span are small. These patterns
ln this figure, it is clear that when
6 c2-ls1-lc5 suggest that competitive ability for light resources
the residual magnitude is small ,
8 c3-ls4-lc3 is an important factor at local scale, constraining
also it is small the contribution of
11 h1-c1-ls1-lc5 community structure and organisation. At the
each category whatever the trait
7 h1-c2-ls1-lc5 scale of the gradient, those patterns seems to be
considered. Those areas are
2 h5-ls4-ls5-lc2 implicated in the forming processes of zonati on.
mostly related to primary
h5-ls5-lc2 (smaller Crown diameter loose relevance alter shrub height
succession and also to recovered
sectors that suffered disturbance 3 scores in ali overcome a certain threshold . Crown diameter is
due to rural activities, and initiated categories) particularly important in open communities where
a secondary processes of h5-ls5-lc2 (Iarger nucleation is still incipient. Formula e-Is-Ie is
12
succession alter ploughing, scores) located westwards over soils rich in calcium and
mowing, and pasture grazing smaller amounts of organic carbono The most
abandonment. Floristic residual is important shilt occur when Is4 beco me dominant over Is1 (i.e. ls1 ->ls4) and accordingly the e2->e3 and
larger in areas less disturbed in le5->le3. Thus microsite occupancy activating competitive exclusion seems to be a relevant component
the past or even pristine for at of the nucleation. Leaf concentration on the outer parts of the individual canopy would had a
least the last 80 years. For these considerable influence balancing competitors.
areas, communities integrate When height is important within the structure of the communities, such as those belonging to general
Figure 3. Variation in the levei of explanation produced by four shrubs of large height, large crown formula h-e-Is-Ie, then crown diameter become negligible following the shift h1->h5. This shift occurs in
biological traits in six classes of residual magnitude. Residuais diameter, large life span and younger dunes (yet behind fo redune domain), normally near the first or second main dune ridge. Two
represent the non-explained component of a floristic data table smaller rank of leaf concentration. different life span categories, Is4 and Is5 characterise a particular area disturbed during the fourties by
alter regression onto soil pH at a depth of 10-15cm. When the residual magnitude are severe oceanic overwash. The explanation for the reinforcement of life span in the architecture of plant
quoted medium or small-medium, communities of this sector, probably relies on the selective disturbance agent that altered competitive
the amount of negative contributions is larger than the positive one. This means that the levei of forces for light resources, favouring the continuity of older resistant elements having strong root system,
explanation due to biological traits is low and insufficient fer these classes, therefore suggesting that while constrained the dominance of short-living small size poorly rooted shrubs, more or less dependent
communities within these ranks should be investigated under other set of biological traits . This on the soil seed bank for recruitment. Renewed opportunities arose from this event, which had favoured
interpretation also should be applied when prior relationship with soil pH gradients is focussed. past niche apportionment, contributing for the particular patterns of individual aggregation shown.

The increase in the local abundance of each trait category in the formula h5-ls5-le2 is a relevant
phenomenon. This one characterises certain spots located in younger dunes, sli ghtly decalcified or,
What kind of funetions are suggested by the eombination of the seleeted set of biologieal
located so near the shore, that due to localised progradative erosion, are prone to calcium concentrati on
traits?
effect in the top-soil due to short range sand blown landwards from beach source. ln these areas, a
Two major trait combinations dominate in visit are a (cC figure 4d). The first one relies on crown larger number of shrubs have grown to higher standards, increasing the proportion of elements that
diameter associated to lile span and leal concentration. For simplicity, the former could be indicated by comparatively show those traits . Competition lar light in the context 01 more abundant soil resources
the formu la e-Is-Ie (cl. table 2). The other one relies on a more complex set of combined traits, actually
Correia, O. & Pinto, M. Nalural Succession in Soulhweslern Portuguese Limy Soils

forces slralificalion among shrubs, which aclually is arare fealure in Ihis vegelalion globally dominaled
by synusial organisalion.

Formula h5-ls5-lc2 is dominanl in a vasl area where lhe florislic residual upon lhe soil pH is quoled
above lhe small-medium rank. This vegelalion Iype covers a large conlinuous lerrilory characlerised by
older dunes, where lhe rale of change of soil pH comparalively decreases slower easlwards. Species
lurnover in Ihis particular landscape seclor seems lo cope wilh olher biological Irails. Nevertheless, lhe
dominanl combinalion of sludied Irails governs lhe seleclion of species Ihal malching slruclurally
diverge physiologically.

REFERENCES

ALBUQUERQUE, J. P.M. (1964). Esquema Climático da Península de Setúbal. Agricultura 21 :8-13.

CATARINO, F.M., CORREIA, O.A., CORREIA, AJD. (1982). Structure and Dynamics of Serra da Arrábida
Mediterranean vegetation. Ecologia Mediterranea VIII:203-222. a) ma p of soi! pH at 10115cm c) six classes of residual magnitude, Le.
beneath the organic layer, components of a 290x55 floristic data table Ihat
CHODAT, R. (1909). Excursions botaniques en Espagne et au Portugal. Buli. Soe. GeniiVe (2"dser.) 1:1-132. interpolated for a regional are not explained by linear regression onto
CLEMENTE, A.S., REGO, F.C. CORREIA, O.A. (1996). Demographic patterns and productivity of post-fire scale analysis by universal soil pH. The magnitude 01 lhe ranks considered
kriging. are: s-small, sm-small/medium , m-medium,
regeneration in portuguese mediterranean maquis. Int. JWildland Fire 6, 5-12. ml-medium/large, Harge, vl-very large.

CLEMENTE,A.S. (2002) . Alter Fire Vegetation Dynamics in Serra da Arrábida. PhD Thesis. University of Lisbon.
Pp.196.
PINTO, M. J., ANDRADE, F. (submitted). Plant species aggregation effects on the zonation pattern of a sand dune
system.

b) map of calcium content in soil d) map of most important biological trait

sam pies collected at 10/15cm clusters. Clusters indicated by numbers
beneath the organic layer. were achieved by non-hierarchical clustering
Symbol dimensions are roughly of trait sample contributlon in Analysis of
proportional to calcium concentration Inertia. Only trait categories scoring above
rneasured in f.l moi g" the mean plus one unit of standard devlation
were selected and included in the processo
Thus, each cluster represenls a first
1km importance levei of combinatorial trall
categories within each sample. Those
combinations are indicated in lable 2.

Figure 4 . Maps depicting results from chemical analysis af sail and

bialagical traits
Quercetea 7: 43-64, 2005
ALFA, Lisboa. Pottugal

Landscape Ecology of the Sado River Estuary (Portugal).

(Dunes, Fresh and Salt marshes)

Carlos Neto', M. Eugénia Moreira' & Rute Caraça"

INTRODUCTION

On the vestibular section, the Sado River drains into a lagoonal estuary, from Alcácer do Sal to
Setúbal-Outão and Tróia; here, it reaches the sea through the outlet called Barra do Sado, between the
high cliff of Serra da Arrábida and the tip of the spit of Tróia.
The chanel of Barra de Sado is cut into the deltaic plain of the submarine delta of the Sado. The
instability of the sediments in the channel is considerable, and increased by the dredging wich is
necessary to mantain the estuarine navigation (Setúbal-Setenave harbours), older then the Roman
times.
ln fact the human occupation of the margins of the estuary began during the Paleolitic period.
Roman an Arabian people created the first estuarine industries - the fishing and the salt production,
destinate to exporto Ali around the estuary archaeological remnants from the Roman occupation have
been found, specially ceramics and ruins of "Cetarea".
During the Medieval period the explotation of salt, the fishing and the hunting were the most important
activities around the estuary. To defend the estuarine commerce, a chain of castles and harbours was
built; Outão, Setúbal, Palmela and Alcácer do Sal are evidence of the fac!.
Ove r this century the cork and cereal (wheat) production, transformed the forested landscape of the
estuarine margins. The cultivation of the rice has been another important activity, moreover on the low
fluvial plains and around the estuary on the reclamed mud-flats. The problem of this cultivation is the
dryness of the climate and the consequente sal inization of the soil (of solontchak type) .
The activities increasing in the present days are related to the fish production, in parks on mud-flats
(some of them substituting old salines), and the tou rism on the spit of Tróia.
The study of the flora and the vegetation of the litoral strip, was done from a phytosociological
perspective, cross-referenced with geomorphological, pedological and climatological studies carried out
in order to understand the principal factors governing the distribuition of the floristic communities .

, Departamento de Geografia da F.L.U.L. Alameda da Universidade 1600-2 14 LISBOA - PORTUGAL.

phone: 00351217920000; fax: 00351217960063. carlosneto@fl,ul,ptand m.albergaria@netcabo.pt
**Departam ento de Planeamento Biofísico ,e Paisagístico, Universidade de Évora. Colégio Luís António
Verney, Rua Romão Ramalho nº 59.7000 EVORA - PORTUGAL. rfmc @potugalmail.pt
 Landscape Ecology of the Sado River Estuary
Neto, Carlos & aI.

ln general, the vegetation in the area under study is influenced by a characteristic mediterranean
climate with an antlantic influence and a lower thermomediterranean bioclimate, according to the BIOGEOGRAPHY OF THE REGION AROUND THE SADO
classification of Rivas-Martinez (1996).
The flora and the vegetation of the area have been arranjed in five broad biogeosystems which are , .... --, ' .
," '-','
, 3c
characterized by the particular floristic communities and by the lito-morpho-pedological characteristics: .,,
,,
--- ,,
- beaches and coastal dunes under the influence of saliness ,,
- dunes and inland sandy coverings
- surfaces made up Marateca formation sandstone and conglomerate
- peat-bogs
- marshy areas wihout peat soils
The flora of these units is mostly mediterranean. However, various atlantic floral elements are to be
found as a result of Quaternary climatic fluctuations. The originality of many of the floristics communities
identified, resides in this suoerimposition of the atlantic and mediterranean species in the sam e territory.

As a result of profound, ancient anthropic action, the floristic communities corresponding to the various
simple types, are poorly represented and the substitution stages, damaged to a greater or lesse r
degree, make up the dominant vegetation.

ITINERARY OF THE FIELD TRIP

. 0'
+Itinerary
1st STOP (Castle of Alcácer do Sal)
Alcácer do Sal lies in the district of Setúbal
and is one of the most important towns
located on the margins of the Sado Estuary. It
has been known since very remote times for
the salt industry developed along the estuary,
because of Mediterranean climate of this
region.
Known as Salacia during the Roman times,
it was so important that the Romans built
there, one of the most important castles of
the Iberian Peninsula. ln 715 the Moors
conquered it and named it Alcaçar de Salaria
(Gacer, al-cacer or kazar means fortress) that Picture 2 - Biogeography of the region around the Sado Estuary and the terminal sector of the Tagus River
developed into its present name. The Moors
MEDITERRANEAN REGION; WEST MEDITERRANEAN SUBREGION;
built a new wall using taipa (a mixture of
clayish earth, rolled pebbles, water and I -Mediterranean West Iberian Province; Lusitan-Extremadurean Subprovince
straw) in its construction. ln no other castle in 1. Marianic-Monchiquensean Sector; 1.1. Baixo Alentejano-Monchiquense Subsector; 1.1.a. Serrano-Monchiquense
the Peninsula was this technique used.
W (. ln ;U Km Superdistrict; 1.1.b. Baixo-Alentejano Superdistrict; 1.2. Araceno-Pacense Subsector; 1.2.a. Alto-Alentejano Superdistrict
- = -=
Ruins of Salacia, 100 to 300 Y AG (the II-Coastal Lusitan-Andalusian Province; Sadensean-Divinding Portuguese Subprovince
Picture 1 - 1st Stop - Gastle of Alcácer do Sal; 2nd forum and wall) where found around the
Stop - Murta Dam lake; 3trd Stop - Garrasqueira 1. Divinding Portuguese Sector; 2. Ribatagan-Sadensean Sector; 3.a. Sadensean Superdistrict; 3.b. Arrabidian
Moorish castle, and have been escavated
Fishing harbour of the Sado Estuary; 4th Stop - since 1983. Superdistrict; 3.c. Ribatagan Superdistrict; 3.d. Olissiponean Superdistrict; 4. Algarvian Sector; 4.a. Vicentine Coast Sup.
Tróia Peninsula- Botanic Reserve.
Neto, Carlos & aI. Landscape Ecology of the Sado River Estuary

Until the construction 01 the railway, the harbour 01 Alcacer do Sal played an important role in the Caracteristic species 01 association and higher syntaxonamic units: Salix salvifolia ssp. australis,
shipment 01 goods Irom the Alentejo to Lisbon. Many warehouses, where the products were stored, can Rubus ulmifolius, Salix atrocinerea, Vitis sylvestris, Salix neotricha, Tamus communis, Hedera
still be seen along the riverbanks. canariensis, Bryonia dioica.
Accompanying species: Phragmites australis, Lythrum salicaria, Typha domingensis, Arundo donax;
80th the salt industry and the port related activities have almost disappeared. Agriculture, mainly rice
Samilax aspera varo nigra.
crops, cattle breeding and lorestry by-products (cork, pine wood, eucalyptus, resin and pine seeds) is
still 01 great importance to this region. These activities associated with the great potentialities 01 the 3 - Blackberry thickets (Lonicero hispanicae-Rubetum ulmifoliae)
Sado River Natural Reserve have contributed to a signilicant development 01 tourism.
It is a common community among the riparian vegetation that can be lound along the riverbanks 01
From the castle, one can see landscape dominated by the terminal sector 01 the Sado River. Among the southwest Iberian Peninsula and represents the Iringe or the replacement phase 01 alder-groves,
the Portuguese rivers, the Sado River has the largest hydrographical basin with 7640 km 2 and a elm-groves, poplar trees, willow groves and ash groves. It is very common in the basin 01 the Sado
typically Mediterranean climate characterized by intense Iloods and long dry periods. The growth 01 River as a replacement phase 01 the riparian bushes and it is easily identilied by the predominance 01
vegetation associated to it, is controlled by the hydrographical regimen. Thus, alder-groves are rare and
blackberry bushes (Rubus ulmifolius).
willows (Salix atrocinerea and Salix salvifolia) , ash-groves and blackberry thickets predominate in the
riparian vegetation , mainly in the lower part 01 the Sado River. The riparian vegetation grows on the Characteristic species 01 association and higher syntaxonamic units: Rubus ulmifolius; Lonicera
margins 01 the river as the all uvium soils are occupied mainly by rice lields. hispanica; Rosa canina; Tamus communis; Bryonia dioica; Crataegus monogyna ssp. brevispina;
Fraxinus angustifolia; Vitis sylvestris; Arum italicum; Silene latifolia.
ln the valley slopes, the groves 01 cork-oaks and 01 hoalm-oaks are dominant. A psammophylic cork- Accompanying species: Scirpoides holoschoenus, Smilax aspera; Torilis arvensis; Cyperus longus
oak grove (Oleo-Quercetum suberis) develops in shallower sands. A non psammophylic cork-oak grove ssp. badius; Lythrum salicaria; Arundo donax; Rubia peregrina; Asparagus aphy//us; Pistacia lentiscus,
(Asparago aphy//y-Quercetum suberis) develops in detritical lormations (gravei, sandstones and
Ulexminor.
conglomerates) and an hoalm-oak grave Myrto communis-Quercetum rotundifoliae develops in
schistous lormations. ln the lower sector 01the Sado River near the estuary, the riparian vegetation is
4 - Ash-groves (Ficario ranunculoidis-Fraxinetum angustifoliae)
dominated by the lollowing plant communities:
It constitutes the second line of vegetation from the river inland ward. It is dominated by ash-groves
1 - Reedy thickets (Typho angustifoliae-Phragmitetum australis) and poplar trees, wich are extremely abundant in the basin of the Sado River. It is very common in
thermo-mesomediterranean territories in western Iberia. The destruction 01 these ash groves leads to
Typical plant community 01 the Atlantic-Mediterranean region and very common along the River Sado their replacement by blackberry thickets dominated by Rubus ulmifolius.
and many 01 its tributaries. It prelers eutrophic or mesotrophic biotopes and it usually corresponds to a
Characteristic species of association and and higher syntaxonamic units: Fraxinus angustifolia; Rubus
permanent community that occupies the lirst line 01 vegetation 01 the riparian geosystem of the lower
ulmifolius; Arum italicum; Populus nigra; Ranuculus ficaria; Salix salvifolia ssp. australis, Salix
Sado, partially submerged during most 01 the year.
It can be lound along Iresh water streams and does not tolerate an increase in the salinity leveI. ln
sally or brackish environmenls 01 the Sado Esluary, this communily is replaced by lhe associalion
Scirpetum compacto-litoralis (Scirpion compacto-litoralis, Scirpetalia compacti, Phragmito-
Magnocaricetea) also characlerized by lhe presence of Phragmites australis and Typha domingensis as
characleristic species bul wilh Iwo importanl and differenlial species Bolboschoenus maritimus varo
compactus and Schoenoplectus litoralis.
Characleristics species 01 association and higher synlaxonamic units Schoenoplectus lacustris,
Phragmites australis, Typha domingensis, Lythrum salicaria, Typha latifolia, Apium nodiflorum, Lycopus
europaeus, Iris pseudacorus, Oenanthe crocata.
Accompanying species: Calystegia sepium, Paspalum paspalodes, Nerium oleander.

2 - Willaw graves (Salicetum atrocinereo-australis)

It is an endemic community in Soulhern Portugal and very common along lhe River Sado and many
01 its tribularies. II can usually be lound in lhe lirsl line 01 arboreous vegetation, is florislically very poor
and wilhoul under cannopy lormalions because it grows on lhe margins 01 Medilerranean slreams and
is subjecl to a slrong Ilooding torrential regimen.
 Landscape Ecol ogy of the Sado River Estuary
Neto, Carlos & aI.
atrocinerea, Bryonia dioica, Rosa canina, Crataegus monogyna ssp. brevispina, Ulmus minor, Alnus
glutinosa, Populus alba, Solanum dulcamara.
Accompanying species : Scirpoides holoschoenus, Rumex crispus, Torilis neglecta, Lythrum salicaria,
Cyperus longus ssp. badius, Oenanthe crocata, Mentha suaveolens, Arundo donax, Calystegia sepium,
Sonchus oleraceus, Rubia peregrina, Foeniculum piperitum, Ficus carica, Smilax aspera, Asparagus
aphyllus, Vicia sativa, Avena barbata, Holcus lanatus, Lepidophorum repandrum, Convolvulus arvensis,
polygonum lapatypholium, Parentucellia viscose, Equisetum ramosissima.

5 - Rush thicket (Holoschoeno-Juncetum acutí)

The thermo-mesomediterranean rush thicket 01 southwestern Iberian Peninsula that grows in sandy
and moist soils with the water-table closer to the sUrface, in the winter. This community represents the
replacement phase 01 the ash-groves Ficario ranunculoidis-Fraxinetum angustifoliae.
Characteristic species 01 association and higher syntaxonamic units: Scirpoides holoschoenus,
Phalaris caerulescens, Cyperus longus ssp. badius, Mentha suaveolens, Agrostis stolonifera, Plantago
lanceoiata, Lythrum junceum, Bromus hordeaceus, Lolium perenne, Juncus acutus, Cynodon dactylon,
Polypogon maritimum, Lotus uliginosus.
Accompanying species: Torilis negletá, Dittrichia viscosa ssp. revoluta, Daucus maritimus, Hordeum
leporinum, Lythrum salicaria, Convolvulus arvensis, Paronychia argentea.

2nd STOP (Murta Dam lake)

Here one can observe a flat bottom depression that corresponds to an interdunal corridor with a
water-table close to the surlace without being subject to great Iluctuations throughout the year. The
Marateca lormation lorms the depression bottom oriented in the direction north-south and is limited to
east and to west by two sandy accumulations 01 psammytic regosoils and podzols with allioz (surraipa)
The moist interdunale depression owes its origins to the rain-water that inliltrates in the dunes until
the low permeability levei 01 the Marateca Formation, where an extensive water table is lormed and
then cul by the inlerdune depression. Thus, as no water Ilows on the dune sUrface, it Ilows to the Ireatic
levei that supplies it to the depression even during summer.
This kind 01 depressions can be lound throughout the Sado Estuary, lhe Galé Coast and, the
Vicentine Coast and, due to the permanent humidity it is possible to observe typical Atlantic and Euro-
Siberian plant communities . Due to the lact that such communities are so rare in the Mediterranean part
01 Europe and that they include some endemic and rare plants and thal their geographical distribulion is
limitid, there is enormous interest in preserving this community.
These moi sI depressions com pose "Atlantic islands" in an open Mediterranean environment and the
vegetation is associated with climatic Iluctuations typical 01 the Quaternary period. During the coo I and
moist phases, the Atlantic and Euro-Siberian vegetation migrated southwards . During the expansion
phases 01 the Mediterranean climate, such as the corrent phase, it only remains in these places 01
permanent humidily due to local and particular hydrological circulation.
Associated with these interdune moist depressions, peat soils can be lound in the South 01 Portugal.
These soils are composed 01 organic matter 01 "muck" type, which has the highest rale 01
decomposition among ali types 01 organic matter that distinguish peat-bogs.
ln the Murta Dam lake, small Iloating islands, made 01 a large amount 01 organic matter (piclure 3),
are lhe result 01 a small dam that covered the peat-bog wilh water and peat rise up to the surface due to
its low density. Thus, in lhe Murta Dam lake the peat-bogs Sphagnum auriculatum, the vegetation
 ---
Neto, Carlos & aI. Lanclscape Ecology of lhe Sacio Ri ver ESlu ary
litorel/etea and the hygrophylic rush thicketsic with Cirsium palustre can be observed in similar but not
very deep depressions along the Sado River are represented sparsely here. On the other side, it is
possible to lind important communities 01 emerging macrophytes, 01Iloating macrophytes rooted in the
soil, and mainly palustrian willow groves 01 the Alnetea-glutinosae class (picture 3).
The lollowing plant communities can be lound in the Murta Dam lake:

I- PALUSTRIAN ECOSYSTEMS

1 - Palustrian willow groves (Cariei lusitanieae-Salieetum atroeinereae)

The palustrian willows bel ong to the class Alnetea glutinosae. This class is composed 01 wood and
shrub communities typical 01 holarctic marshy areas. ln the Mediterranean these communities are rare
and considered a relict as they represent traces 01 Ouaternary glaciations that remained as biotopes
with an edalic compensation (edapho-hygrophylic and mi re). ln the peat soils 01 the Superdistrict 01 the
Sado River, the class Alnetea glutinosae is represented by the Alnetalia glutinosae order and the Alnion
glutinosae alliance, associating the marsh bushes with alder groves (Alnus glutinosa) based on a
European distribution. ln the Sado ri ver, the Alnion glutinosae alliance is represented by the endemic
association 01 peat soils 01 the Superdistrict 01 the Sado River Cariei lusitanicae-Salicetum atrocinereae.
Characteristic species 01 association and higher syntaxonamic units: Salix atrocinerea, Carex
paniculata ssp. lusitanica, Thelypteris palustris, Myrica gale, Frangula alnus, Salix salvifolia ssp.
australis, Alnus glutinosa. Differentials species 01 Phragmito-Magnocericetea: Lythrum salicaria, Iris
pseudacorus Phragmites australis, Lycopus europaeus, Cladium mariscus, Typha domingensis, Carex
pseudocyperus, Calystegia sepium, Typha latifolia, Schoenoplectus tabernaemontani, Carex
lamprocarpa, Cyperus longus ssp. badius. Differentials species 01 the Genistion micrantho-anglicae:
Ulex minor, Erica ciliais, Genista anglica ssp. ancistrocarpa.
Accompanying species: Rubus ulmitolius, Eleocharis multicaulis, Cirsium palustre, Lonicea
periclymenum ssp. hispanica, Erica erigena, Mol/inia caerulea, Galium palustre, Juncus effusus,
Schoenus nigricans, Hydrocotyle vulgaris, Potentil/a erecta, Hypericum elodes, Polygonum salicifolium,
Scrophularia sublyrata, Scirpoides holoschoenus, Pteridium aquilinum, Anagallis tenel/a, Hypericum
humitusum, Erica lusitanica, Rhynchospora rugosa, Festuca arundinacea, Holcus lanatus,
Schoenoplectus mucronatus, Carex de missa, Lobelia urens, Cal/una vulgaris, Erica scoparia,
Mantisalca salmantica, Oenanthe lachenalii, Rubia peregrina ssp . longitolia.

2- Marsh community of Myriea gale

Plant cornmunity dominated by Myrica gale and accompanied by some large helophytes such as
Phragmites australis, Lythrum salicaria, Iris pseudacorus, Lycopus europaeus, Cladium mariscus,
Typha domingensis, Carex pseudocyperus, Typha latifolia, Schoenoplectus tabernaemontani and
typical species 01 peat heaths such as Genista anglica ssp. ancistrocarpa. The occurrence 01 the
previously mentioned species is occasional. Very often the community 01 Myrica gale lorms adense
and monospecilic shruby lormation that constitutes a recovery step 01 Alnetea glutinosae (Cariei
lusitanicae-Salicetum atrocinereae).

I
'----
 --
Neto, Carlos & aI.
Lanclscape Ecology of the Sacio River E stuary

3 - Reedy thickets Phragmites australis (Typho angustifoliae-Phragmitetum australis) When the depth of water reduces , the necessary conditions for the development of hygrophyle heath
thickets, hygophyle rush thickets, peat bog with Sphagnum and palustrian willow groves are created.
It is a plant community dominated by slender reeds (Phragmites australis) , lake-scirpus
(Schoenoplectus lacustris) and narrow-Ieal-cattail (Typha angustifolia, Typha latifolia and Typha Characteristic species of association and higher syntaxonamic units: Cladium mariscus, Phragmites
dominguensis) very Irequent in lagoons and dams along the Sado River and the Galé Coast. It is a australis, Lythrum salicaria, Lycopus europaeus, Carex lusitanica.
typical community 01 Iresh water streams and does not tolerate an increasing salinity. It prefers Accompanying species: Cyperus longus ssp. badius, Scirpoides holoschoenus, Galium palustre,
eutraphic ar mesotrophic enviranments with low water energy and peat ar gley sai Is. Lythrum junceum, Juncus acutiflorus ssp. acutiflorus.

ln general reed thickets Typho angustifoliae-Phragmitetum australis represent a permanent phase of 6 - Fresh water communities formed by rooted plants with floating leaves (Myriophy/lo-
the vegetation and do not present a serial logic as it happens in the Murta Dam lake. However, when Nupharetum lutei)
the water is not very deep, the association may constitute a replacement phase 01 the palustrian wil low
graves of Salix atrocinerea. . An association dominated by Nuphar lutea, Polygonum salicifolium, Nymphaea alba and
Ceratophyllum demersum growing in aquatic environments of deep and caim Iresh water. This
Characteristic species of association and higher syntaxonamic units: Phragmites australis, community dominated by hydrophytes is very Irequent in the Supperdistrict 01 the Sado River, in dams
Schoenoplectus lacustris, Typha domingensis, Iris pseudacorus, Lycopus europaeus, Lythrum salicaria, that do not dry in the summer. ln the Murta Dam lake, this association constitutes an almost pure
Bolboschoenus maritimus, Typha latifolia, Alisma plantago-aquatica, Mentha aquatica, Scrophularia community 01 Nymphae alba, only accompanied by Polygonum salicifolium.
auriculata varoglabra ta, Sparganium erectum ssp. neglectum.
Accompanying species: Juncus maritimus, Hypericum humifusum, Calystegia sepium, Galium Characteristic species of association and higher syntaxonamic units: Nymphaea alba, Polygonum
palustre, Rubus ulmifolius, Lotus uliginosus, Salix atrocinerea, Agrostis stalonifera, Tamarix africana, salicifolium, Nuphar lutea, Potamogeton po/ygonifolius.
Scirpoides holoschoenus, Juncus effusus varo effusus, Salix salvifolia ssp. australis. Accompanying species: Myriophyllum altemiflorum, Callitriche stagnalis

4 - Community of high cyperaceous (Caricetum pseudocypero-Iusitanicae) 7 - Panico-Paspaletum vagina ti

It is a tall, dense and herbaceous community, poor in species, typical of peat soils that are Irequently This community, wich is almost monospecilic, is dominated by Panicum repens, grawing on the sandy
submerged in enviranments of low energy waters. margins 01 lakes, ponds and dams that are characterized by great Iluctuations of water levei due to the
ri ce fields.
ln the Murta Dam lake, this community occurs in areas more than 10 cm deep, during most of the
year, and constitutes a degradation step ar fringes of the Salix atrocinerea peat bog forest. Willow It occurs in nitrophylic sandy soils, poor in organic matter. During the winter, when dam lakes are fu ll,
graves Carici lusitanicae-Salicetum atrocinereae grow in small peat floating islands and an association they are covered with water. Panicum repens may be temporarily submerged 01 1m deep. During the
with Caricetum-pseudocypero lusitanicae was observed on the margins of such small islands. The soil summer this community may reveal a significant hydrological delicit.
is always covered with water. Contact with palustrian willow graves justifes its presence among the
accompanying species 01 some typical species 01 the relerred willow graves, such as Myrica gale,
11- PSAMMOPHYTIC ECOSYSTEMS
Thelypteris palustris and Salix atrocinerea.
Characteristic species 01 association and higher syntaxonamic units: Carex paniculata ssp. lusitanica, 8 - Thyme/gorse thickets (Thymo capite/lati-Stauracanthetum genistoidis)
Carex pseudocyperus, Cladium mariscus, Phragmites australis, Lythrum salicaria, Thelypteris palustris, It is lormed by small shrubs dominated by Stauracanthus genistoides and Thymus capitellatus. It is
Schoenoplectus tabemaemontani, !ris pseudacorus. sometimes dense, typical 01 the psammitic regosoils of the interior stabilized dunes 01 the Superdistricts
Accompanying species: Hydrocotyle vulgaris, Myrica gale, Ludwigia palustris, Salix atrocinerea, of the Sado River and the Vicentine Coast (north of Melides). It is a vegetal population wich is rich in
Frangula alnus. psammophytic endemism (Thymus capitellatus, Armeria rouyana, Armeria pinifolia, Malcolmia lacera
ssp. gracilima), dominant throughout the Sado landscape. This domain results from the anthropic
5 - Moist Mediterranean Ibero-Atlantic, Atlantic and Central European marshes Cladium origins of the gorse thickets Stauracanthus genistoides, wich is the first step in the degradation 01 the
mariscus (Cladietum marisci) Juniperus navicularis bushy formations due to anthropical processes (the exploitation of farming and
It constitutes a moister natural Iringe (permanent phase of the vegetation) of Salix atrocinerea forestry). Planting of stone pine (Pinus pinea), maritime-pine (Pinus pinaster) and eucalyptus trees
palustrian bushes or a more advanced degradation step, compared to the palustrian community of (Eucalyptus camaldulensis) leads to the destruction of pre-existing bushes. Deep land ploughing in
Myrica gale. Cladium mariscus forms dense, tall (up to 2,5m) and almost pure, populations, because podzol also destroyed the alioz that prevents deep roots Iram grawing. These proloundly changed
they avoid the shorter species to grow. biotopes are colonized by characteristic species of the association Thymo-Stauracanthetum genistoidis.

The large amount 01 biomass supplies a large quantity 01 dead-Ieaf coverage to the soil, which Characteristic species of association and higher syntaxonamic units: Saturacanthus genistoides,
decomposes very slowly as a result of its characteristics and due to the existing water in the soil Thymus capitelatus, Halimium calycinum, Halimium halimifolium ssp. multiflorum, Lavandula
throughout the year. This phenomenon may origininate very thick peat soils. pedunculata ssp. lusitanica, Ulex australis ssp. welwitschianus, Malcolmia lacera ssp. gracilima,
Oianthus broteri ssp. hinoxianus, Gistus salvifolius, Iberis linifolia ssp . welwitschii, Euphorbia boetica.
 Lancl scape Ecolo gy of the Sacio Ri ver Estuary
Neto, Carlos & aI.

Accompanying species. Conyza bonariensis, Cladonia mediterranica, Cladonia potentosa, Characteristic species of association and higher syntaxonamic units. Erodium aethiopicum ssp.
Arrhenatherum album, Santolina rosmarinifolia varo impressa, Sedum sediforme, Pinus pinaster, pilosum, Linaria spartea, Corynephorus divaricatus ssp . macrantherus, Rumex bucephalophorus ssp.
Corynephorus canescens va ro maritimus, Carlina corymbosa, Euphorbia portlandica, Helichrysum hispanicus, Tuberaria guttata, Hypochaeris glabra, Logfia ga//ica, Silene scabriflora, Polycarpon
picardi varo picardi, Oaphne gnidium, Pinus pinea, Elaeoselinum gummiferum, Oittrichia viscosa ssp. alsinifolium, Asterolinum linum-ste//atum, Micropyrum tene//um, Loeflingia baetica varo micrantha,
revoluta, Pimpine//a vi//osa, Rosmarinus officinalis, Asparagus aphy//us, Anemone palmata, Stipa Leucojum trichophy//um varo trichophy//um, Silene littorea, Aira caryophy//ea, Coroni//a repanda ssp.
gigantea, Centaurea sphaerocephala ssp. sphaerocephala, Ca//una vulgaris, Carpobrotus edulis, repanda, Silene ga//ica, Briza maxima, Anthy//is hamosa, Lupinus angustifolius ssp. reticulatus, Ononis
Genista triacanthos, Fritillaria lusitanica, Juniperus navicularis, Quercus suber, Retama sphaerocarpa, baetica varo baetica, Jasione montana ssp . blepharodon, Euphorbia exigua, Vulpia membranacea,
Cytisus grandiflorus, Thymus carnosus, Scrophularia frutescens, Armeria pungens ssp. pungens, Ononis broterana, Ornithopus sativus ssp. isthmocarpus, Arenaria algarbiensis, Tolpis barbata,
Calendula suffruticosa ssp. algarbiensis, Eucalyptus camaldulensis. Ornithopus pinnatus, Rumex angiocarpus, Evax pygmaea ssp. ramosissima, Lotus subbiflorus ssp.
caste//anus, Paronychia cymosa, lonopsidium acaule, Loeflingia tavaresiana, Arenaria conimbricensis,
9 - Sub-moist psammophytic, thermomediterran ean juniper tree Juniperus navicularis of the Lathyrus angulatus, Agrostis tenerrima.
Sado river - (Daphno gnidii-Juniperetum navicularis) Accompanying species : Senecio ga//icus, Spergularia purpurea, Juncus capitatus, Spergula arvensis,
Vulpia alopecuros, Bromus rigidus, Avena longiglumis, Euphorbia portlandica, Mibora minima,
It is adense and sometimes tall shrub dominated by Juniperus navicularis, endemic in the Sado River Centranthus calcitrapae ssp. calcítrapae, Sesamoides canescens ssp. canescens, Corynephorus
and the Vicentine coast that grows in the inland stabilized dunes where the mature step of the system is canescens var. maritimus, Silene colorata, Malcolmia lacera ssp. gracilima, Papa ver somniferum ssp .
constituted. The dryness and nutrient-poor psammophytic and sometimes podzol soils associated with setigerum, Andryala arenaria ssp. parvipila, Carduus meonanthis, Oipcadi serotinum, Brassica barrelieri
the stabilized dunes that are not subject to the influence of the saline spray, prevent the development of ssp. oxyrrhina, Chamaemelum mixtum, Raphanus raphanistrum ssp. microcarpus, Arrhenatherum
cork-oak groves (Quercus suber); thus the xerophytic savine thickets Juniperus navicularis are the most album, Avena ba rba ta, Anagalis arvensis, Ste//aria media, Anarrhinum be//idifolium, Anemone palma ta,
developed plant formation. Cork-oaks groves Oleo-Quercetum suberis only occur in places in which the Carpobrotus edulis, Centaurea sphaerocephala ssp. sphaerocephala, Cynoglossum creticum, Silene
sand depth is lower, composing a climax sclerophylic and psammophylic bus h of the class Quercetea nicaeensis, Sci//a monophy//os, Papaver rhoeas, Poronychia argentea, Campanula lusitanica ssp .
ilicis. So, in deep and dry sands, the mature step of the system is represented by Juniperus navicularis, matritensis, Elaeoselinum gummiferum, Radiola !inoides, Ga!ium mínutulum.
while in wetter sands (with a higher levei of moisture due to its proximity to the water-table) the mature
step is represented by cork-oak groves Oleo-Quercetum suberis. 3rd STOP (Carrasqueira Fishing harbour of the Sado Estuary)
Characteristic species of association and higher syntaxonamic units: Juniperus navicularis, Oaphne The sucession of the salt marsh communities shown in the picture, nowadays can not be observed in
gnidium, Phi//yrea angustifolia, Corema album, Osyris alba, Anemone palma ta, Osyris quadripartita, Carrasqueira harbour, because of the marsh antropogenic erosion, caused by the enlargement,
Pistacia lentiscus, Rubia peregrina ssp. longifolia, Sci//a monophy//os, Arbutus unedo. deepening, and weidening of the main channels and by trailling of the boats on the high marsh where
Accompanying species: Ca//una vulgaris, U/ex australis ssp. welwitschianus, Rosmarinus offícinalis, they are repared and parked. Also the high marsh is connected to dykes built to reclaim the mud flats,
Halimium halimifolium ssp. multiflorum, Cladonia mediterranica, Lithodora prostrata ssp. lusitanica, transformed in rice-fields. Haliomiono portulacoidis-Sarcocornietum alpinii dominates the marsh, with
Pinus pinaster, Cladonia potentosa, Cistus salvifolius, Erica umbe//ata, Halimium calycinum, Lavandula narrow fringes of Spartinetu m maritimae, along the narrow tide channels, and marsh pools. Cistancho
pedunculata ssp. lusitanica, Erica australis, Cytisus grandiflorus, Genista triacanthos, Chamaespartium phelypaeae -Arthrocnemetum fruticosae are mixed with Suaedetum verae. Along the dykes, in very few
tridentatum, Quercus suber, Cistus ladanifer, Arrhenatherum album, Euphorbia terracina, Friti//aria places, small patches of the transitiion marsh of Polygono equisetiformis-Juncetum marítimi, can be
lusitanica, Thapsia vi//osa, Senecio ga//icus, Pinus pinea, Stauracanthus spectabilis ssp. spectabilis, seen, far from the harbour.
Thymus capite//atus, Eríca scoparia ssp. scoparia, Helichrysum picardi varo virescens, Iberis linifolia ssp.
welwitschíi, Stauracanthus genistoides. The scheme of the figure 5 can be apreciated along the Eastern side of Tróia spit. Glose to the
estuarine beach of the Roman Ruin, the subtidal salt grasses of Zoosteretum no/tii, psammophylic (fine
10 - An nual psammophytic community of the inland dunes Anachorlo-Arenarietum sand), can rise the water levei during the spring low tides. Southern most, dense populations of
algarbiensis of the Sado River and Vicenti ne coast Spartinetum maritimae colonize sandy small spits and bars , and allow the low marsh of horse-grass to
associate the Sarcocornio perennis-Puccinelietum convolutae and to spread on the muddy sllke. The
It is a pioneer therophytic plant community that openly colonizes the glades of shrubs Stauracanthus high marsh has no erosion scarp and is cut in small islands by the tidal creeks . Hal/Omlono
genistoides and Stauracanthus spectabilis (Thymo capite//ati-Stauracanthetum genístoidis and Thymo portulacoidis-Sarcocornietum alpinii is locate along the margins and into the depressions, some places
camphorati-Stauracanthetum spectabilis). It occupies biotopes that are very poor in organic matter and associated to Spartina marítima and the Sarcocornio perennis-Puccinelietum convolutae. On the convex
characterized by strong daily and annual temperature fluctuations . Under these conditions, the sectors of the mudflat, Cistancho phelypaeae-Arthrocnemetum fruticosae, and Inulo cnthmoldls-
competition moved by perennial plants is weak and only the small pioneer therophytes can reach in the Arthrocnemetum glauci and Cistancho phaelypaeae-Suaedetum verae, grow and form dense
spring, the conditions of humidity, temperature and light necessary for their survival. It is aPortuguese
endemic association, distributed throughout the Superdistricts of the Sado River and the Vicentine
Coast. The floristic diversity is enormous and the presence of two important Portuguese endemisms can
be observed (Ionopsidium acaule and Malcalmia lacera ssp. gracilima).
Neto, Carlos & aI. Landscape Ecology of lhe Sado River Estuary

communities. The ecotone slat-marsh-dune is colonized by plants Ihat have their roots into the sand
covered muddy soils, such as Po/ygono equisetiformis-Juncetum maritimi, or are delinitely
psammophillic such as Arthemisio-Armerietum pungentis that extend to lhe upper limit 01 lhe freatic salt-
water leveI.

Ta ble 1- Flonstlcal composilion oI the Carrasqueira sa t mars hes communilies.

1 2 3 4 5 6 7 8 9
l ostera nollii X
Spartina maritima X X X X
Limonium vulaare X X X X
Sarcocomia perennis X X X X
Puccinellia convoluta X X X X
Halimione portulacoides X X X X X X X
Sarcocomia alpini X X X X X
Sarcocomia fruticosa X X X X X X X
Cistanche phelvoaea X X X X X
Triqlochin bulbosa SSP. barrelieri X X X X X
Sperqularia maritima X X
Puccine/lia foucaudii X X X
Arthrocnemum macrostachyum X X X X X
Limonium ferulaceum X X X X X
Inula crithmoides X X X X X X
Artemisia caerulescens X
Limonium daveaui X X
Limonium lanceolatum X
Suaeda vera X X X X X
Polygonum equisetiforme X X X X
Elymus elonaatus X X X
Juncus maritimus X X X X
Juncus acutus X X X
Carex extensa X
Sperqularia salina X X
Centaurium spicatum X
Centaurium tenuiflorum X
Sonchus maritimus X
Oenanthe lachenalli X
Aster trioolium ssp. pannonicus X
Lactuca saliana X
Apium araveolens X
Bolboschoenus maritimus X X X
Phragmites australis X X
Typha dominaensis X
Frankenia laevis X X X X
Salsola vermiculata X
Atriplex halimus X
Beta vulgaris ssp. maritima X
Plantago coronopus ssp. ceratophy/la X X
Salicomia lutescens X
Salicomia oatula X X X X
Suaeda maritima X X X X X X
 paz

Neto, Carlos & aI. Landscape Ecology of lhe Sado Ri ver Estuary

Suaeda sp/edens
Sa/so/a soda
Parapho/is filiformis X X X X
Hordeum marinum X X X ''' ..
Po/ypogon maritimus X X X
Parapho/is incurva
Beta macrocarpa X Q)
>
Cressa cretica X Q;
ln
Spergu/aria bocconei X X X Q)
IX:
Mesembryanthemum nodiflorum X X ", . .~
Atrip/ex prostata X X X c
m
Sonchus tenerrimus X Õ
Oxalis pes-caprae X e?.
m
Hedypnois cretica X ::;
-'. , ' ln
Avena barbata X ., . . .. .!::
c
Cotu/a coronopifo/ia X . : :.. . ...
' Q)
c..
Me/i/otus messanensis X m
Hordeum /eporinum X X . -. :õ
Anacyc/us radiatus X .=
Q)

Piptatherum miliaceum X .s
Pha/aris paradoxa X õ
ln

1- Zosteretum no/ti (CosIa et ai. 1991); 2-Spattinetum maritimae (CosIa et ai. 1997); 3-Sarcocornium
. '. Ec
:::l
perennis-Puccinellietum convolutae(Cosla et ai. 1997); 4- Halimiono pottulacoidis-Sarcornietum alpini E
E
(CosIa et ai. 1997); 5 - Cistancho phelypaeae-At1hrocnemetum fruticosae (CosIa et ai. 1997); 6 - o
o
Polygono equisetiformis-Juncetum maritimi (CosIa et ai. 1997); 7- Inulo crithmoidis-Atthrocnemetum o
glauci (CosIa et ai. 1997); 8- Cistacho phelypaeae-Suadetum verae; 9 - Frankenio laevis-Salsoletum ~
..c
vermiculatae (CosIa et ai. 1997) c.
o
E
E
4th STOP (Tróia Peninsula- Botanic Reserve) m
ln
c.
Q)
One can observe lhe sequence of planl communilies from lhe shoreline lowards lhe inland as a .s
consequence of lhe changes of lhe biolope physical characteristics such as the reduction of the õ
undulalion and lhe saline spray influence, reduclion of lhe wind speed , increase of sand slabilizalion, E
~
grealer paedogenelic developmenl ele. As a resull, lhe coaslal psammophylic communilies grow in Ol
parallel slrips along coaslline organizing a microgeosigmelum in which plan! populalions are no! .!!!
"O
included in a serial logics bu! represenl permanenl sleps of lhe vegelalion. Q)
>
~
"E
Q)
1 - Annual psammophytic halonitrophilous vegetation, western Mediterranean - 5alsolo kali- ln
Q)

Cakiletum aegyptiacae o..

Q)
IX:
II is a halonilrophylic and psammophylic communily dominaled by pioneer Iherophyles Ihal grow CD
prefe rably in lhe upper par! of lhe medi um seclor of lhe beach or in lhe lransilion zone lo lhe high beach Q)
::;
where Ihey are subjecl lo a direcl and permanenl influence of sall bolh in lhe soil and in lhe air, a õ
conslanl and easy movemenl of lhe sandy subslralum, permanenl wind action and somelimes of wave c::
and currenls lidaI. II is an associalion poor in species Ihal grows in beach sands wilh organic delrilus
carri ed by lhe sea du ring high lides and slorms. II reaches ils peak of developmenl during lhe end of lhe
spring and in lhe summer, when lhe condilions of Iighl, waler, lemperalure and sand slabilily are more
favou rable.
Neto, Carlos & aI.
Landscape Ecology of lhe Sado River Estuary

Characteristic species of association and higher syntaxonamic units: Caki/e maritima ssp. maritima, 4 - Chamaephytic, psammophytic and thermophytic vegetation of the coastal penestabilized
Sa/so/a ka/i ssp. ka/i, Euphorbia pep/is dunes of the Sado River Bassin, Vicentine and Algarve Coast - Artmemisio crithmifoliae-
Accompanying species: Po/ygonum maritimum, E/ymus farctus ssp. boreo-atlancticus, Euphorbia Armerietum pungentis
para/ias, Eryngium martimum, Pancratium maritimum, Ca/ystegia so/danel/a The communities of the Ammophi/ion australis alliance that grow in unstable dunes towards the inland
are gradually replaced by the chamaephytic communities of the alliance Crucianel/ion represented by
2 - Perennial halopsammophytic vegetation of Europe Atlantic beaches - Euphorbio paraliae-
the Artemisio crithmifo/iae-Armerietum pungentis association that colonizes the penestabilized dunes
Agropyretum junceiformis
from the Tagus River mouth until Cadiz.
It correspond to an herbaceous plant communily lhat colonizes the hig-beach sand, hardly covering Its composition includes some important endemisms on the southern coast of Portugal (From the
the ground from the coastline particularly in the flat seclors invaded by the wave currents and high tides.
Tagus River until Ponta Umbria, in Spain), such as Thymus camosus and Linaria /amarckii.
II is composed of a halopsammophytic vegelation, cyclically affected by lhe waves (specially during
during storms), and therefore it is subject to a strong sand movement. The percentage of organic matter The greater distance from the sea, the less influence of marine spray and the prevailing winds ,
in the soil is extremely low and the salin ity, both in the soil and in the air, is relatively high. E/ymus allowing for the development of a diversified flo ra that covers the ground. The movement of sandy
farctus ssp. boreo-atlanticus is the dominanl plant and has to lolerale being covered or uncovered by particles is almost inexistent, which associated to a soil rich in nutrients leads to the development of a
the ondulation waves and the wind. Its radicular system plays an important role in the sand stabilization community dominated by chamaephytes.
and its aereal part interferes in the accumulation processus of the beach sand.
Characteristic species of association and higher syntaxonamic units: He/ichrysum ita/icum varo
Characteristic species of association and higher syntaxonamic units.· E/ymus farctus ssp. boreo- picardi, Armeria pungens ssp. pungens, Ammophi/a arena ria ssp. austra/is, Artemisia crithmifo/ia,
atlanticus, Eryngium maritimum, Euphorbia para/ias, Ca/ystegia so/danel/a, Pancratium maritimum, Crucianel/a maritima, Lotus creticus, Ma/co/mia /ittorea, Corynephorus canescens varo mantlmus,
Medicago marina, Po/ygonum maritimum, Crucianel/a maritima, Artemisia crithmifo/ia, Otanthus Pancratium maritimum, Anagal/is monelli varo microphyl/a, Sesel/i tortuosum, Ononis natrix ssp .
maritimus, Ammophi/a arenaria ssp. austra/is, Linaria /amarckii ramosissima, Scrophu/aria frutescens, Eryngium maritimum, Hemiaria maritima, Medicago marina,
Accompanying species: Crithmum maritimum, Caki/e maritima ssp. maritima, Sa/so/a ka/i ssp. ka/i, Euphorbia port/andica, Cyperus capitatus, Euphorbia para/ias, Thymus camosus, Linaria /amarckii.
Carpobrotus edu/is, Reichardia gaditana, Euphorbia pep/is, Cutandia marítima Accompanying species: Sedum sediforme, Pimpinel/a vil/osa, Carpobrotus edu/is, Reichardia
gaditana, Antirrhinum majus ssp . cirrhigerum, Senecio gal/icus, Si/ene /ittorea, Ca/endu/a suffuticosa
3 - Psammophytic community of unstable dunes of Divisório Português, Tagano-Sadense, ssp. a/garbiensis, Ononis broterana, Senecio vu/garis, Lagurus ovatus, Cynog/ossum creticum, Corema
Algarviense, Onubense and Gaditano - Loto cretici-Ammophiletum australis a/bum, Santolina rosmarinifolia varo impressa, Lobu/aria maritima, Caki/e maritima ssp. maritima,
Anchusa calcaria, Arrhenatherum a/bum, Oianthus hinoxianus, Lotus subbiflorus ssp. castel/anus,
This communily is dominaled by Ammophi/a arenaria ssp. austra/is thal densly populates the top of Vu/pia a/opecuros, Stauracanthus genistoides, Crithmum maritimum, Po/ypogon maritimum, Cerastium
small, high-beach dunes (coastal nebkhas) and the movable dunes influenced by the marine spray. It
g/omeratum, Rumex bucepha/ophorus ssp. hispanicus, Car/ina corymbosa.
grows between the populations of Euphorbio-Agropyretum that colonize lhe beach and the populations
of the chamaephyles that belong lo lhe Crucianel/ion alliance. Thus , it inhabits the fixed inland dunes. 5 - Annual psammophytic community of the penestabilized and unstable coastal dunes of the
High density tufts of Ammophi/a are a barrier to the aeolian current that lose energy and, by Sado Estuary and Vicentine Coast - Herniario a/garvicae-Linarieturr. fica/hoanae
consequence, sandy particles deposil causing the rising of the topographic surface and the burial of the
aereal part of the plant. During the stability periods, the plants develop their aereal bodies creating It is a therophytic community comprised of pioneer species that occupy biotopes adjacent to the
radicular syslems that can reach 9m in deep. The sands mobility may also exhume the plant, however chamaephytic communities of the Crunianel/ion maritimae alliance (Artemisio crithmifo/iae-Armerietum
the community plays an important role in the dune stabilization. pungentis) and the crowberry thickets (Rubio /ongifo/iae Coremetum a/bi) that grow on the coastal
penestabilized dunes. It is endemic in Portugal and is distributed throughout the Sado superdistricts and
Characteristic species of association and higher syntaxonamic units. Ammophi/a arenaria ssp. the Vicentine Coast. Among the characteristic species of association and higher syntaxonamic units,
austra/is, Othantus maritimus, Eryngium maritimum, Ca/ystegia so/danel/a, Euphorbia para/ias, two important Portuguese endemisms occur (Linaria fica/hoana and Hemiaria a/garvica). The former
Pancratium maritimum, Lotus creticus, Medicago marina, Hemiaria maritima, Po/ygonum maritimum, (Linaria fica/hoana) characterizes the entire geographycal area of the association distribution (between
Crucianel/a maritima, Armeria pungens ssp. pungens, Artemisia crithmifo/ia, E/ymus farctus ssp. boreo- the Peninsula of Tróia and Sagres). Hemiaria a/garvica can only be found towards the south of Sines
atlanticus, Ma/co/mia /ittorea , Linaria /amarcki, Thymus camosus, /beris procumbens ssp. procumbens. until Sagres.
Accompanying species.· Caki/e maritima ssp. maritima, Carpobrotus edu/is, Cynog/ossum creticum,
Si/ene /ittorea. Characteristic species of association and higher syntaxonamic units: Linaria fica/hoana, Si/ene
/ittorea, Policarpon a/sinifo/ium, Rumex bucepha/ophorus ssp. hispanicus, Erodium aethiopicum ssp.
pi/osum, Ononis broterana, Tuberaria guttata, Hypochaeris glabra, Vu/pia membranacea, Cutandia
maritima.
Accompanying species: Senecio gal/icus, Ma/comia /ittorea, Sedum sediforme, Papa ver somniferum
ssp. setigerum, Euphorbia port/andica, Hemiaria maritima, Anagallis mone/Ii varo microphyl/a, Pimpinel/a
vil/osa, Corynephorus Gqhescens varo maritimus, Cyperus capitatus, Ca/endu/a suffuticosa ssp.
 Neto, Carlos & aI. Landscape Ecology' of the Sado River Estuary

algarbiensis, Cynoglossum creticum, Silene colora ta, Capobrotus edulis, Sesamoides canescens, B - Annual subnitrophylous and ombrophyllous community of the coastal stabilized dunes of
Leontodon taraxacoides ssp. longirostris, Asterolinum linum-stelatum, Cerastium glomeratum, Mibora Gaditano-Onubo-Algarviense of Geranio purpurei-Galietum minutuli
minima, Senecio vulgaris, Avena longiglumis, Centhranthus calcitrapa, Rumex angiocarpus, Reichardia
gaditana, Vulpia alopecuros. It is a therophytic, psammophytic, ombrophytic and subnitrophytic association inhabitus under the
savine (Juniperus turbina ta), in the coastal stabilized dunes.
6 - Psammophytic and heliophytic crowberry thickets (Corema album) typical of the littoral It is composed 01 small ombrophytic therophytes, requiring sub-nitrophyles and registers a great
penestabilized dunes of the Atlantic coasts of the Iberian Peninsula - Rubio longifoliae- vitality in the savine coastal areas 01 the Tróia Peninsula. The nitrilication 01 the biotope is increased by
Coremetum albi the anthropic origin, due to the garbage left by visitors and to organic dejects 01 animais.
It is a termophylic and heliophyte community dominated by Corema album that very olten constitutes Characteristic species 01 association and higher syntaxonamic units: Geranium purpureum, Urtica
almost pure, high density populations. It can be lound on the coastal penestabilized dunes representing membranacea, Galium minutulum, Myosotis ramosissima ssp. ramosissima, Centranthus calcitrapae
towards the south 01 Aveiro, the Iringe or lirst replacement step 01 savina wood (Juniperus turbinata) 01 ssp. calcitrapae, Stellaria media, Cardamine hirsuta, Geranium molle, Mercurialis annua, .Cerastium
the alliance Juniperion turbinatae (Osyrio quadripartitae-Juniperetum turbinatae). glomeratum, Galium verrucosum, Arabidopsis thaliana, Oxalis pes-caprae, Myosotls persoonfl.
Characteristic species 01 association and higher syntaxonamic units: Corema album, Antirrhinum Accompanying species: lonopsidium acaule, Erodium aethiopicum ssp. pilosum, Rumex
majus ssp. cirrhigerum, Rubia peregrina ssp. longifolia, Asparagus aphyllus, Scilla monophyllos bucephalophorus ssp. hispanicus, Scrophularia sublyrata, Silene latifolia ssp. latifolia, Aetheorhiza
Accompanying species: Sedum sediforme, Capobrotus edulis, Cistus salvifolius, Halimium calycinum, bulbosa ssp. bulbosa, Umbilicus rupestris.
Senecio gallicus, Lavandula pedunculata ssp. lusitanica, Helichrysum italicum ssp. picardii, Lithodora
prostra ta ssp. lusitanica, Calendula suffruticosa ssp. algarbiensis, Cladonia mediterranica, Ulex australis
ssp. welwitschianus, Halimium halimifolium ssp. multiflorum, Carlina corymbosa, Arrhenatherum album, BIBLlOGRAPHY
Armeria pungens ssp. pungens, Euphorbia portlandica, Cladonia potentosa, Silene littorea, Thymus
carnosus, Stauracanthus spectabilis ssp. spectabilis, Pinus pinaster, Erodiurn aethiopicurn ssp. pilas um, BRAUN-BLANOUET, J.; ROZEIRA, A. & PINTO DA SILVA, A. R. (1972) - "Résultats de trois
Rumex bucephalophorus ssp. gallicus, Corynephorus canescens varo maritimus, Reichardia gaditana, excursions géobotaniques à travers le Portugal septentrional et moyen, IV. Equisse sur la
Ammophila arenaria ssp. australis, Ononis natrix ssp. natrix. vegetation dunale», Agronomia Lus~., 33(1-4): 217 -234.
BRAUN-BLANOUET, J.; ROZEIRA, A. & SILVA, A. R. P. DA (1964) - "Résultats de trois excursions
7 - The savine thicket of the Coastal, thermo-mediterranean dry-submoist dune typical of the géobotanique à travers le Portugal septentrional et moyen - III Landes à cistes et ericacées (Cisto-
provinces of Gaditano-Onubo-Algarviense and Divisório Português - Osyrio quadripartitae- Lavanduletea et Calluno- Ulicetea»>, Agronomia Lusitanica 23(4) .
Juniperetum turbinatae CASTROVIEJO, S. & ai. (1986-1997) - Flora Iberica. (Planas Vasculares da Península Ibérica e Islas
Baleares). Madrid, Real Jardim Botánico, Volumes I a V.
It is adense and tall shrubby lormation that can be lound in the coastal stabilized dunes with a grey COSTA, J.C., EspíRITO-SANTO, D. & LOUSÃ, M. (1994) - "The Vegetation 01 Dunes 01 Southwest
sandy soil (non podzolic) still subject to the spray marine inlluence. ln such dunes the poor soils and the Portugal ". Silva Lusitana, Ano II, Vol. 2, nº1, pp. 51-68
direct inlluence 01 the maritime winds do not allow the development 01 Ouercetalia i/icis. Juniperus COSTA, J.C., CAPELO, J. H., LOUSÃ, M. & AGUIAR, C. (1993) - "Communautés de Juniperus au
turbinata represents the mature step 01 the system and does not move towards the interior. It can often Portugal ". Colloques phytosociologiques, Vol XXII, pp. 499-511.
be lound in present stabilized dunes recently lormed without podzolic processes. Towards the inland COSTA, J.C. , CAPELO, J. H.;,LOUSÃ, M.; AGUIAR, C. & NETO, C. (1998) - "Biogeogralia de Portugal
the communities 01 Juniperus turbinata are replaced by Janiperus navicu/aris that sometimes can be Continental". Ouercetea. Vol. O, pp. 1-56.
lound in old podzolic dunes. COSTA, J.C.; CAPELO, J. H., NETO, C , EspíRITO-SANTO, D. & LOUSÃ, M. (1997) - "No;as
Fitossociológicas sobre os Tojais do Centro e Sul de Portugal» . Silva Lusitana, Ano V, Vol. 5, n- 2,
Characteristic species 01 association and higher syntaxonamic units: Juniperus turbina ta, Rubia
/ongifolia, Pistacia /entiscus, Rhamnus oleoides, Osyris quadripartita, Atnirrhinum majus ssp. pp.275 - 282. . .. . .
COSTA, J.C., LOUSÃ, M. & PAES, A.P.O. (1996) - "As Comunidades Rlbelnnhas da Bacia
cirrhigerum, Asparagus aphyllus, Corema a/bum, Rhamnus a/aternus, Phillyrea angustifo/ia, Daphne Hidrográlica do Rio Sado (Alentejo Portugal)". Actas do I Colóquio Internacional de Ecologia da
gnidium, Ouercus coccifera, Asparagus acutifo/ius
Vegetação, Évora, pp. 291-320. ,
Accompanying species: Sedum sediforme, Carpobrotus edu/is, C/adonia sp., Pimpinella villosa, Pinus COSTA, J.C., LOUSÃ, M., CAPELO, J., ESPIRITO-SANTO, M. D. , IZCO, J. & ARSENIO, P (2000) -
pinaster, Ca/endu/a a/garbiensis, Dactylis hispanica, Arrhenatherum a/bum, Car/ina corymbosa, Cistus The coastal vegetation 01 the portuguese divisory sector: dunes, cliffs and low-scrub communltles.
sa/vifo/ius, Rosmarinus officina/is, Ha/imium ca/ycinum, He/ichrysum ita/icum ssp. picardi, Armeria
Finisterra, XXXV, 69, pp. 69-93. . . .
pungens ssp. pungens, Ammophi/a arenaria ssp. austra/is, Calluna vu/garis.
COSTA, J. C. (1999) - Excursion to the Tejo and Sado Estuanes. ln Ana IS,abel Correia & ai. Ed. LIVro
de Resumos e Guias de Excursões das V. Jornadas de Taxonomia Botamca, pp. 87-98.
COUTINHO, A.X.P. (1939) - Flora de Portugal. Plantas Vasculares, Livraria Bertrand, Lisboa, 2ª ed. ,
938 p.
Neto, Carlos & aI.
Quercetea 7: 65-81 , 2005
FRANCO, A. & AFONSO, M. da L. R. (1994) - Nova Flora de Ponugal (Continente e Açores), Volume III
(Fascículo I), Escolar Editora, Lisboa, J81 pago ALFA, Lisboa. Portugal
FRANCO, A. (1971) - Nova Flora de Portugal (Continente e Açores) Volume I (L YCOPOOIACEAE -
UMBILlFERAE) , Lisboa, 647 pago Sintra Vegetation and Landscape
FRANCO, A. (1984) - NO'la Flora de Ponugal (Continente e Açores), Volume II (CLETHRACEAE -
COMPOSITAE), Lisboa, 659 pago
GARRETAS, B. D. (1984) - "Datos sobre la vegetaciõn psammofila de las costas Portuguesas". Sandra Mesquita*, Pedro Arsénio**, Mário Lousã*, Tiago Monteiro Henriques* & José Carlos Costa*
Oocuments Phytosociologiques, Instituto di Botanica dell'Università-Camerino, Camerino, Vol. VIII,
p.71-83.
MATEUS, J. E. (1992) - Holocene and present-day ecossystems of the Carvalhal region, southwest
Portugal, Lisboa, Tese de doutoramento, 183 pago
MOREIRA, M. E. S. A. (1987) - «Estudo Fitogeográfico do Ecossistema de Sapal do Estuário do The field trip to Sintra region, as part
Sado», Finisterra, Centro de Estudos Geográficos, Lisboa, Vol XXII, nº 44, pp. 247-296.
of the 48 th IAVS Symposium, aims to
MOREIRA, M.E.S.A. & PSUTY, N. (1993) - «Sedimentação holocénica no Estuário do Sado. Nota
preliminar» , Actas da 3ª Reunião do Quaternário Ibérico, Coimbra, pp. 289 - 297. illustrate well-preserved and
°
MOREIRA, M.E.S.A. (1985) - «A evolução do litoral a partir da análise da rede hidrográfica. exemplo
da Ribeira da Comporta», Lisboa, Actas da I. ª Reunião do Quaternário Ibérico, Vo11 , pp. 297-331
representative plant communities of
Sintra municipality. II consists of two
MOREIRA, M.E.S.A. (1987) - Glossário de Termos Usados em Geomorfologia do Litotal. Centro de short field stops and a 3 km walk
Estudos Geográficos, Estudos de Geografia das Regiões Tropicais, ReI. nº 15, 167 pago (Fig.1).
NETO, C (2002) - "A Flora e a Vegetação das Dunas de Tróia e da Costa de Galé". GuineanA , nº 8..
Leioa, p. 1-269. From a geological point of view, the
NETO, C.S. (1997) - A Flora e a Vegetação dos Meios Palustres do Superdistrito Sadense, Lisboa, study area has a matrix 01 sedimentary
Centro de Estudos Geográficos, 96 p. rocks, cut by intrusive and extrusive
NETO, C.S; CAPELO, J.H. & COSTA, J.C. (1996) - «Sobre a posição fitossocio!ógica dos matos de
magmatic outcrops.
Stauracanthus genistoides (Brot.) Sampaio e Santolina rosmarinifolia L. varo impressa (Hoffmanns.
& Link) Coutinho no Superdistrito Sadense.». Silva Lusitana, Ano IV, Vol. 4, nº 2, pp.255 - 257. The older sedimentary rocks of the
RIVAS-MARTINEZ, S., FERNANDEZ-GONZÁLEZ, J. LOIDI, M.LOUSÃ & A. PENAS (2001) -
territory were lormed during Superior
Syntaxonamical checklist of vascular plant connunities of Spain and Portugal to association levei.
Itinera Geobotanica 14: 5-341. Jurassic, in marine environment. They
RIVAS-MARTíNEZ, S., LOUSÃ, M.; DIAZ, T.E.; FERNANDEZ-GONZALEZ, F. & COSTA, J.C. (1990) - correspond mostly to limestone,
«La Vegetación dei Sur de Portugal (Sado, Alentejo y Algarve)>> , Itinera Geobotanica, Asociacion sometimes with marly leveis or
Espafiola de Fitosociologia (A.E.F.A.), Madrid, Vo1.3, p.5-127. dolomite. The Cretaceous deposition
RIVAS-MARTINEZ, S., DíAZ, H., FERNANDEZ-GONZÁLEZ, F., IZCO J., LOIDI, J., LOUSÃ, M. & environment changed cyclically Iram
PENAS, A (2002) - Vascular plant communities of Spain and Portugal. Addenda to the cheklist of marine to fluvial and lacrustrine. Thus,
2001. /tinera Geobotanica 15: 5-922. - ==--o
K!
----,
10 ai 1m

there are several limestones, marls,

clays and sandstones Irom this period.
Fig. 1 - lIinerary of the excursion
Sintra mountain range is the result of

* Departamento de Protecção de Plantas e de Fitoecologia. Instituto Superior de Agronomia: Tapada

da Ajuda 1349-017 Lisboa. Portugal. mesquita.sandra @sapo.pt; manolousa @lsa.utl.pt; tmh @lsa.utl.pt,
jccosta @isa .. utl.pt.
** Secção Autónoma de Arquitectura Paisagista. Instituto Superior de Agronomia. Tapada da Ajuda
1349-017 Lisboa. Portugal. arseniop @isa.utl.pt.

F; 4
Mesquita & alo Sintra Vegetation and Landscape

an intrusive event occurred about 80 million years ago, called "Maciço Eruptivo de Sintra" (MES), i.e. This lield trip is divided in two parts. ln the morning we will visit an example 01 the most common
Sintra Eruptive Massif. It consists 01 a syenite nucleus surrounded by a long granite ring, occasionally vegetation series in the area, developing on limestone substrata. The places visited are included in the
separated by a band 01 gabbros and diorite. By means 01 contact with rocks dating Irom the Jurassic Olissiponean Super-District (Dividing Portuguese Sector) and Upper Thermomediterranean Lowe r
period , the MES originated a border 01 contact metamorphism, lorming glassy limestone, calcareous Subhumid bioclimatic belts.
schist, etc. This vegetation mosaic corresponds to the potential and seral stages 01 the series Arisaro clusii-
The landscape 01 the westernmost hall 01 the District is dominated by several small volcanoes 01 the Querco broteroi sigmetum (Iigure 3). It is a characteristic and endemic series 01 limestone territories
"Complexo Vulcânico de Lisboa" (CVL) - Lisbon Volcanic Complex. These were lormed during the 01 the Center West 01 mainland Portugal. The mature stage is a Portuguese oak woodland - Arisaro
Superior Cretaceous and the beginning 01 the Tertiary periods, expelling lava Ilows and pyroclastic c/usii-Quercetum bote roi.
materiais in alternate lashion. The resulting basalt complex sits on top 01 the limestone 01 the Upper This woodiand Irequently exhibits an herbaceous border, the community Leucanthemo sy/vatici-
Cretaceous period (Turonian). Cheirolophetum sempervirentis, which also occupies small clearings in the woods where the sunlight is
Also dating Irom the Tertiary period, some sites 01 conglomerates with elements 01 variable nature able to penetrate, as opposed to under closed crown cover where is shady and lew plants thrive.
and size, and also limestone and clays 01 deposition in marine environment can be lound. This series also includes a wild pi um prickly high shrub hedge, Rubo ulmifolii-Prunetum insititioidis, an
Finally, there are some areas 01 sand and shingle 01 ancient beaches and lossil dunes Irom the endemic community. Nowadays, well preserved Quercus faginea subsp. broteroi woodlands are rare
Pleistocene, as well as mobile dunes Iram the Holocene. and, therelore, it is uncommon to lind wild plum hedges in their typical seral position. These
communities have, however, a huge development as separation lences 01 agricultural lields, and
The meteorological stations located in Sintra municipality are classilied as Oceanic Pluviserotinal
therelore are very common in this territory.
Mediterranean Macrobioclimate. Most 01 the region lalls into Upper Thermomediterranean thermotype,
with the exception 01 the Sintra mountain range, which is classilied in the Upper Mesomediterranean ln wetter and deeper soils, the wild plum community is replaced by a laurel hedge - Vinco difformis-
thermotype. As lor the ombrotype, the majority 01 this territory is Lower Sub-humid, with the exception, Lauretum nobilis. This laurel thicket is not very common, appearing typically in hedges close to streams
once again, 01 the Sintra mountain, which belongs to the Upper sub-humid ombrotype, and 01 a small or near rocky and North laced rock walls, shady and often with small water springs.
northern coastal area, which belongs to Dry Upper ombrotype. The lirst seral stage 01 this oak woodland is a high scrub, also endemic to the Center West 01
Sintra municipality has been under intense human inlluence since historical times - which is shown Portugal, Bupleuro fruticosae-Arbufefum unedonis, dominated by strawberry trees and other tall shrubs,
by the countless roman, visigothic and medieval remains currently exhibited in the Museum 01 "São such as Phil/yrea latifolia, Viburnum tinus and Phil/yrea angustifolia.
Miguel de Odrinhas". This area was essentially rural until mid 20 th century, with exception 01 the The second substitution stage is a kermes oak thicket, Melico arrectae-Quercetum cocciferae, a
mountain, which has been lor long a summer leisure destination lor nobleman and the bourgeoisie. dense high shrub community dominated by kermes oak. This is a very common community in the study
Parks and gardens with exotic ornamental vegetation, palaces and glass-houses are abundant in the area. When the anthropic successional pressure (usually cutting or lire) in dry and rocky places ceases
mountain range. The Palace and the Park 01 Pena, built in the 19th century by D. Fernando II, dominate lor enough time, this community may not evolve again to the terminal stage 01 the ecological
the mountain landscape and are an example 01 romantic architecture 01 Sintra. succession, but instead to a wild olive tree woodland (Olea europaea varosylvestris). This is a version 01
During the second hall 01 the 20 th century, Sintra suffered an intense urban development as a result 01 the kermes oak thicket, co-dominated by tall wild olive trees, lorming part 01 secondary woodland. This
the expansion 01 Lisbon suburbs. woodland may eventually evolve to the following stages 01 succession or become permanent.

Due to this development, natural vegetation was Iragmented and conlined to marginal positions in The next seral stage is a meadow of perennial grass dominated by Brachypodium phoenicoides -
hedges, rocky grounds not suitable lor agriculture and inaccessible areas. Phlomido Iycnitidis-Brachypodietum phoenicoides. Tall grasses dominate this community, allowing a
complete covering of the soil ali year round.
Recently, agricultural set aside allowed natural vegetation to re-occupy some 01 areas lormerly
ploughed. Therelore, the lower stages 01 vegetation succession recover nowadays in signilicant areas. Finally, the endemic gorse thicket Salvio sclareoides-Ulicefum densi stands lor the last perennial
The mature stages - mainly climatophilous woodlands - are very rare and only small nuclei in areas stage in the successional process in thin soils. This gorse thicket occurs freq uently in a mosaic pattern
less lavorable to the urban expansion remain. As well as coastal areas, which still have well preserved with the two stages described above, associated to recurring but not very intense wildfires.
natural vegetation. This series includes an annual plant community 01 Helianthemetea gultati vegetation class , as an
extreme regression stage.

67
66
Mesquita & ai. Sintra Vegetation and Landscape

1sI Stop: Cabeceiras da Ribeira de Cabrela (Alcolombal)

ln the afternoon, a walk ove r sea-cliffs is planned. We will pass through several types 01 coastal This place has one 01the lew Quercus faginea subsp. broteroi woodland Ihat can still be lound in the
vegetation, lirstly on limestone c1iffs, then on granite cliffs (Iigure 2). This walk begins in the District 01 Sintra (Arisaro c1usii-Quercetum botero/). The understory 01 oak woodland is nevertheless
Olissiponean Superdistrict and enters the Sintranean Superdistrict (both in the Dividing Portuguese poorly preserved since it has been used lor grazing. However, one can lind the species associaled to
Sector). The Irontier between the Upper Thermornediterranean Lower Sub-humid belt and the Lower this mediterranean oak woodland, namely climbers - Smilax aspera, Hedera maderensis subsp.
Mesomediterranean Upper Sub-humid belt will be crossed later in the day. iberica, Rubia peregrina, Rosa sempervirens, Lonicera implexa - and shade-Ioving herbs or shrubs-
Sintra municipality exhibits two types Viburnum tinus, Coronilla valentina subsp. glauca, Ruscus aculeatus, Iris foetidissima, Vinca difformis.
01 maritime cliffs, according to the type It is also possible to observe some hedge communities, in mosaic pattern with the oak woodland: a
01 rock in which they are cu!. This lact laurel thicket (Vinco difformis-Lauretum nobilis), like the one next to the rock wall Ihat limits the
is paramount to differentiate distinct woodland, and a tall-Iorb community (Leucanthemo silvatici-Cheirolophetum sempervirentis).
complexes 01 plant cornmunities.
Some 01 the seral stages 01 the oak woodland can also be lound in a slope that was most likely burnt
However, the vegetation structure is down a lew years ago. Namely a perennial grassland - Phlomido Iychitidis-Brachypodietum
similar in both cases. It is a lairly sirnple phoenicoides - can be seen in the lower portion 01 the slope, which nowadays is used lor grazing.
geo-complex lormed by a Also adense kermes oak woodland - Melico arrectae-Quercetum cocciferae - in the upper part,
Fig. 2 - Schematic litological prolile 01 the local coastal microgeoseries with only one
cliffs
community and a geoseries with only
one series, 01 which the only climactic stage is a juniper community with kermes oak (Querco
cocciferae-Juniperetum turbina ta e) .

The microgeoseries is lormed by a community 01 herbaceous perennial plants adapted to the extreme
habitat conditions in the steep sea clills. This community develops on rocks, under the inlluence 01
strong salty winds (i.e. splashed by marine salt spray). On limestone (or sandstone) cliffs occurs a
community 01 Limonium spp. - Limonietum multiflori-virgati (Iigure 4). On subslrata other than
limestone, namely syenite, granite or basalt, appears a community 01 roman carnalions (Armeria
pseudarmeria) and rock carrots (Daucus halophillus) - Diantho cintrani-Daucetum halophili, endemic
to Sintra (Iigure 5).

On the other hand, in the clill platform, over thin soils, Ihrives the juniper / kermes oak series - Querco
cocciferae-Junipero turbinatae sigmetum that presents two successional facies. ln limestone substrata,
this series has as seral stages the gorse thicket Salvio sclareoidis-Ulicetum densi, variant 01 Daphne
maritima, and a Brachypodietum meadow, Phlomido Iychnitidis-Brachypodietum phoenicoides (Iigure 3).
ln non-calcareous substrates a dillerent successional facies is present, which has as lirst seral stage a
maritime gorse Ihicket 01 U/ex jussiaei, Daphno maritimi-Ulicetum congesti, lollowed by a perennial
grassland 01 Stipa gigantea, Avenulo sulcatae-Stipetum giganteae (Iigure 5).

80th sucessional facies have as regressive stages extreme communities, which are annual pioneers
01 the class Saginetea maritimae.

Fig. 3 - Arisaro clusii-Querco brotero; sigmetum: 1 Arisaro c1usii-Quercetum broteroi;

2 Bupleuro ruticosae-Arbutetum unedonis; 3 Leucanthemo silvatici-Cheirolophetum sempervirentis;
4 Melico arrectae-Quercetum cocciferae; 5 Phlomido Iychitidis-Brachypodietum phoenicoides;
6 Salvio sclareoides-Ulicetum densi; 7 Rubo ulmifolii-Prunetum insititioidis
Mesquita & ai. Sintra Vegetation and Landscape

dominated by Quercus coccifera and Pistacia lentiscus is present.

The path crosses a small stream with rocky banks and bed with some riverside vegetation , although
poorly preserved. A community 01 tamarisk (Polygono equisetiformis-Tamaricetum africanae) develops
in the rocky banks, where the characteristic habitat conditions 01 temporary streams 01 the driest
Mediterranean territories are partial ly reprodu ced. Contacts with the alluvial terrace - in which the
natural vegetation was destroyed and replaced by agricultural lields - traces 01 ash tree woodlands
(Ficario ranunculoidis-Fraxinetum angustifoliae) or its most common seral stage - hygrophilous
bramble thicket (Lonicero hispanicae-Rubetum ulmifolil) can be observed.

2nd Stop: Moinhos da Torre

ln this site, a community 01 Salvio sclareoidis-Ulicetum densi, gorse thicket dominated by Ulex densus
- species endemic to Portugal - can be observed. The potential distribution area of these gorse
thickets is limited to the limestone soils 01 the Dividing Portuguese Sector, but the majority 01 its current
occurrence area is in Sintra region . This gorse thicket lorms a mos ai c pattern with the perennial
grassland Phlomido Iychitidis-Brachypodietum phoenicoides.

3rd Stop: Walk between Adraga Beach and Roca Cape

This walk begins in the limestone cliff next to Adraga beach, stretching through the abrasion platform,
amongst communities 01 Querco cocciferae-Juniperetum turbinatae dominated by Juniperus turbinata,
Quercus coccifera and Phillyrea angustifolia. ln the steep cliffs over the sea, communities 01 Limonietum
multiflori-virgati can be observed.

Fu rther ahead , a mos ai c pattern 01 juniper community with gorse thicket 01 U/ex densus - Salvio
sc/areoidis-Ulicetum densi, variant 01 Daphne maritima, pillow-shaped communities lew centimeters
high, can be observed. Occasionally, in small patches 01 sand, one can also see communities 01
Armerio welwitschii-Crucianel/etum maritimae, with Helichrysum picardi, Armeria welwitschii and Ononis
ramosissima.

Nearly hallway through the walk, we start the descent to Ribeira da Ursa valley. This is a very small
valley with steep slopes, cut in the cliff by a stream. Hall way through the descent, the limestone
changes into granite and correlative vegetation chang e also lollows.

On the slope above the beach, one can see the communities Diantho cintrani-Daucetum halophili,
local endemic vegetation - i.e., Irom the seacoast Sintranean Superdistrict. We emphasize the
presence 01 Dianthus cintranus subsp. cintranus, Armeria pseudarmeria, Limonium virgatum and
Daucus halophilus.

Climbing to Roca Cape, the seral stages 01 the juniper community in acid plutonic substrata can be
observed: the gorse thicket 01 Ulex jussiaei- Daphno maritimi-Ulicetum congesti- and the perennial
grassland Avenulo sulcatae-Stipetum giganteae.
Mesquita & aI. Sintra Vegetation and Landscape

ln the last stretch of the route, already on syenites , this vegetation is enriched by the presence of
shrubby individuais of pyrenean oak (Quercus pyrenaica).

The walk ends close to Roca Cape, an outstanding 142 meter high headland above the Atlantic
Ocean, in the most westerly place of Continental Europe.

Synthetic tables and relevees of the most representative communities are presented below:

Synthetic table of Arisaro c1usii-Quercetum broteroi 8r. -81. , P. Silva & Rozei ra 1956 (Quercetea
i/ieis, Querceta/ia i/icis, Quercion broteroi, Quercenion brotero!) from LOUSÃ et alo (1994) , 11 relevees:
Characteristics: Quercus faginea subsp. broteroi V, Smi/ax aspera varoa/tissima V, Rubia peregrina
varo /ongifo/ia V, Ruscus acu/eatus IV, Hedera maderensis subsp. iberica IV, Euphorbia charaeias IV,
Arisarum vu/gare var c/usii III, Laurus nobi/is III, Rhamnus a/atemus III, Asparagus aphy//us III, Rosa
sempervirens III, Arbutus unedo II , Phi//yrea /atifo/ia II, Vibumum tinus II, Pistacia /entiscus II, Myrtus
communis II, Coroni//a g/auca II, Jasminum fruticans II, Daphne gnidium II, Osyris alba II, Asp/enium
onopteris II , /ris foetidissima II, Luzu/a forsteri subsp. baetica II, Erica arborea I, Quercus coccifera
subsp. rivasmartinezii I, O/ea europeae varo sy/vestris I, Paeonia broteroi I, Bup/erum rigidum subsp.
panicu/atum I, Carex distachya I, Cepha/antera /ongifo/ia I, Quercus rotundifo/ia t, Quercus suber t,
Pu/icaria odora t , Lonicera imp/exa t , Me/ica minuta subsp. arrecta t, Sanguisorba hybrida t ,
Anemone pa/mata t, Asparagus acutifo/ius t , Hyacintoides hispanica t, Se/agine//a denticu/ata t ;
Companions: Rubus u/mifo/ius IV, Crataegus monogyna subsp. brevispina IV, Lonicera peryclimenum
subsp. hispanica II, Tamus communis II, Ca/amintha baetica II, Brachypodium sy/vaticum II, Origanum
virens II, Sa/via sc/areoides II, Dacty/is hispanica II, Smirium perforatum II, Po/ypodium austra/e II,
Asp/enium trichomanes II, Sedum a/bum II, Prunus spinosa subsp. isititioides I, Vinca difformis I,
Teucrium scorodonia I, Cistus sa/vifo/ius I, Cheir%phus sempervirens I, Brachypodium phoenicoides I,
Arist%chia paucinervis I, Agrimonia eupatoria I, Pteridium aqui/inum I, Arum ita/icum I, Geranium
purpureum I, Asp/enium ceterach I, Ptenospartum tridentatum s.l t , Genista triacanthos t, Piptaterum
mi/eaceum t, C/inopodium vu/gare subsp. arundanum t , Cistus crispus t, Digita/is purpurea t, Urgin ea
maritima t, Ranuncu/us pa/udosus t , Geum sy/vaticum t , Bituminaria bituminosa t , Smimium
o/usatrum t , Daucus crinitus t , Be//is perennis t , A/lium rase um t, Phagna/on saxati/e t , P/antago
/anceo/ata t .

Synthetic table of Melico arrectae-Quercetum cocciferae quercetosum cocciferae 8r.-81. , P. Silva

& Rozei ra 1956 (Quercetea i/icis, Pistaeio /entisei-Rhamneta/ia a/atemi, Asparago a/bi-Rhamnion
o/eoidis) from COSTA et alo (2004) , 19 relevees: Characteristics: Quercus coccifera V, Rhamnus
a/atemus V, Daphne gnidium V, Smi/ax aspera varoaspera V, Lonicera imp/exa V, Rubia peregrina varo
/ongifolia V, Me/ica minuta subsp. arrecta V, Pistacia /entiscus IV, O/ea europaea varo sy/vestris IV,
Asparagus aphy//us IV, Arisarum vu/gare varoe/usii IV, Genista toumefortii III, Osyris alba III, Euphorbia
charaeias III, Arbutus unedo II, Phi//yrea angustifo/ia II, Myrtus eommunis II, Vibumum tinus II, Phi//yrea
 Sintra Vegetation and Landscape
Mesquita & alo
subsp. insititioides V, Vinca difformis V, Rubus u/mifo/ius V, Quercus faginea subsp. broteroi IV, Ruscus
/atifo/ia II , Rhamnus o/eoides II, Coronil/a g/auca II, Ruscus acu/eatus II, Bup/eurum rigidum subsp.
aculeatus IV, Rubia peregrina varo /ongifolia IV, Tamus communis IV, Arbutus unedo III, Viburnum tinus
panicu/atum II, Hedera maderensis subsp. iberica II, Hyacintoides hispanica II, Vinca difformis II,
III, Osyris alba III, Rhamnus a/aternus III, Quercus coccifera III, Euphorbia characias III, Lonicera
Anemone pa/mata II, Scil/a monophyl/os II, Carex hal/erana II, Se/aginel/a denticu/ata II, Laurus nobilis I,
peryclimenum subsp. hispanica III , Crataegus monogyna subsp. brevispina III , Ulmus minor III, Phil/yrea
Bup/eurum fruticosum I, Erica arborea I, Pu/icaria odora I, Quercus /usitanica t , De/phinium pentaginum
/atifo/ia II, Phil/yrea angustifo/ia II, Pistacia lentiscus II , Olea europaea varo sylvestris II, Asparagus
t , Asparagus acutifo/ius t, Carex distachya t; Companions: Brachypodium phoenicoides V, Urginea
aphy/lus II, Asp/enium onopteris II , Luzu/a forsteri subsp baetica II, Arisarum vulgare varo clusii, II,
maritima V, Cistus sa/vifo/ius V, Rubus u/mifo/ius IV, Sa/via sclareoides IV, Teucrium Origanum virens
Castanea sativa II, Fraxinus angustifo/ia II, Cory/us avel/ana II, C/ematis vitalba II, Prunus lusitanica I,
IV, Si/ene /ongici/ia IV, Antirrhinum majus subsp. /inkianum IV, Cheir%phus sempervirens IV, Dactylis
Bup/eurum fruticosum I, Lonicera etrusca I, Me/ica minuta subsp. arrecta I, Hyacinthoides hispanica I,
hispanica IV, Bel/is sy/vestris III, Geranium purpureum III, Prunus spinosa subsp. insititioides II, Cytisus
Quercus robur I, Prunus avium I, Rosa canina I; Companions: Cheir%phus sempervirens IV,
grandif/orus II, U/ex densus II , U/ex jussiaei II, Cistus crispus II , Cistus monspe/iensis II, Tamus
Teucrium scorodonia IV, Origanum virens III , /ris foetidissima III, Brachypodium sylvaticum III, Bryonia
communis II, Erica scoparia II, scorodonia subsp. scorodonia II, Ruta cha/epensis II, Asphode/us
dioica III, Sa/ix atrocinerea II, Pteridium aqui/inum II , Si/ene longici/ia II , Lathyrus clymenum II
/usitanicus II, Arist%chia paucinervis II, Astraga/us /usitanicus II , Lathyrus clymenum II, Thapsia vil/osa
Piptatherum mi/iaceum II, Leucanthemum sy/vaticum I, Polypodium vu/gare I, Cistus sa/vifo/ius I,
II, Crataegus monogyna subsp. brevispina I, Pteridium aqui/inum I, Ca/endu/a suffruticosa subsp.
Rosmarinus officinalis I.
/usitanica I, Clinopodium vu/gare I, Stachys /usitanica I, Piptatherum mi/iaceum I, Bar/ia robertiana I,
Arum ita/icum I, Agrimonia eupatoria t , Sedum sediforme t, Cal/una vu/garis t, Feru/a communis t, Synthetic table 01 Leucanthemo sylvatici-Cheirolophetum sempervirentis J.C. Costa, Ladero, T.E
Smirnium o/usatrum t, Lavatera o/bia t, Bryonia dioica t , Lithodora prostrata subsp. /usitanica t, Díaz, Lousã, Espírito Santo, Vasconcelos, Monteiro & Amor 1993 (Trifo/io-Geranietea, Me/ampyro-
Lobu/aria maritima t, Cynara humi/is t, Crepis versicaria t. Holceta/ia, Origanion virentis, Stachydo lusitanicae-Cheir%phenion sempervirentis), Iram COSTA et alo
(2001), 5 relevees: Characteristics: Cheirolophus sempervirens 5, Teucrium scorodonia 4, Calamintha
baetica 4, Leucanthemum sy/vaticum 3, Clinopodium vu/gare subsp. vu/gare 3, C/inopodium vulgare
Synthetic table 01 Bupleuro fruticosae-Arbutetum unedonis Capelo, J.C. Costa & Rivas-Martínez subsp. arundanum 3, Brachypodium sylvaticum 3, Origanum virens 3, Sedum forsteranum 2, Silene
2002 (Arbuto unedonis-Laurion nobilis, Pistacio /entisci-Rhamnetalia a/a temi, Quercetea ilicis) Iram la tifo/ia 2, Stachys germanica subsp. lusitanica 2, Vinca difformis 2, Heleborus foetidus 2, Hypericum
COSTA et alo (2002), 9 relevees: Characteristics: Arbutus unedo V, Erica arborea V, Vibumum tinus V, perforatum 2, Agrimonia eupatoria 2, Lathyrus sy/vestris 2,Carduus broteroi 1, Geum sy/vaticum 1,
Coronil/a g/auca V, Smi/ax aspera varo aspera V, Phil/yrea angustfo/ia V, Pistacia /entiscus V, Picris spinifera 1, Primu/a vulgaris 1; Companions: Digita/is purpurea 4, Prune/la vulgaris 3, Cistus
Asparagus aphyl/us V, Bup/erum fruticosum IV, Rubia peregrina var. /ongifo/ia IV, Myrtus communis IV, salviifo/ius 3, Antirrhinum majus subsp. /inkianum 3, Salvia sclareoides 3, Geranium purpureum 3, Rubia
Quercus coccifera subsp. coccifera IV, Phil/yrea /atifo/ia IV, Daphne gnidium IV, Rosa sempervirens IV, peregrina varo longifo/ia 3, Luzula forsteri subsp. baetica 3, Conyza bonariensis 2, Coronil/a glauca 2,
Rhamnus a/atemus IV, Vica difformis IV, O/ea europea varo sy/vestris III, Lonicera imp/exa III, Ruscus Bup/eurum fruticosum 2, Myrtus communis 2, Erica arborea 2, /ris foetidissima 2, Bituminaria
acu/eatus III, Arisarum vu/gare varo c/usii III, Osyris alba III, Euphorbia characias III, Quercus faginea bituminosa 2, Rhagadi/us stel/atus 2, Si/ene /ongicilia 2, Arist%chia paucinervis 2, Arum ita/icum 2,
subsp. broteroi II, Rhamnus o/eoides II, Lonicera etrusca II, Laurus nobi/is II, Anemone palmata II, Geranium rotundifolium 2, Carex ha/lerana 2, Be/lis perennis 2, Sanguisorba rupico/a 2, Brachypodium
Bup/erum rigidum subsp. panicu/atum II, Jasminum fruticans II, Genista tournefortii II, Juniperus phoenicoides 2, Thapsia vil/osa 2, Calendu/a suffruticosa subsp. /usitanica 2, Euphorbia characias 2,
turbina ta I; Companions: Rubus u/mifo/ius IV, Teucrium scorodonia IV, Brachypodium phoenicoide IV, Rosa sempervirens 1, Lapsana communis 1, Daucus carota 1, Andrya/a integrifo/ia 1, Centaurium
Crataegus monogyna subsp. brevispina IV, Tamus communis III, Erica scoparia III, Cistus erythreae subsp. grandiflorum 1, Biscutel/a /usitanica 1, Carex flacca 1, Dipsacus comosus 1,
monspeliensis III, Cistus sa/viifo/ius III, Ca/amintha baetica III, Cheir%phus sempervirens III, Pu/icaria Sanguisorba hybrida 1, Piptatherum mi/eaceum 1, Myosotis baetica 1, Oenanthe crocata 1, Vicia sativa
odora III, Geranium purpureum III, Prunus spinosa subsp. insititiodes II, Lonicera peryclimenum subsp. 1.
hispanica II, Lavandu/a /uisieri II, Picris spinifera I, Astraga/us /usitanicus I, Cistus crispus I, Dacty/is
Synthetic table 01 Salvio sc/areoidis-Ulicetum densi Rivas-Martínez, Lousã, Díaz, Fe rnandez-
g/omerata subsp. hispanica I, Cistus a/bidus I, Bryonia dioica I, /ris foetidissima I, Sa/via scareoides I,
González & J.C. Costa 1990 ex Capelo, J.C. Costa, Lousã & Neto 1992 variant of Daphne maritima
Geum sy/vaticum I. (Rosmarinetea, Rosmarineta/ia, Saturejo- Thymbrion capitatae, Serratulo estremadurensis-Thymenion
sylvestris,) lrom COSTA et ai. (2001), 8 relevees: Characteristics: Ulex densus V, Eryngium di/a ta tum
V, Salvia sclareoides V, Anthy/lis vu/neraria subsp. maura IV, P/antago serraria va ro hispanica IV, Carex
Synthetic table 01 Vinco difformis-Lauretum nobilis Capelo & J.C. Costa in J.C. Costa, Lopes,
ha/lerana IV, Cistus monspe/iensis II, Ruta cha/epensis II, Rosmarinus officina/is II , Bartsia aspera I,
Capelo & Lousã 2001 (Arbuto unedonis-Laurion nobilis, Pistacio /entisci-Rhamnetalia a/atemi,
Iberis linitolia subsp. microcarpa I, Daucus crinitus I; Differentials of variant Daphne marítima:
Quercetea i/icis) , Iram COSTA et alo (2002), 9 relevee: Characteristics: Laurus nobilis V, Hedera
Daphne gnidium var.maritima V, Dactylis marina IV, Ca/endula suffruticosa subsp. algarbiensis IV,
maderensis subsp. iberica V, Smilax aspera varo aspera V, Rosa sempervirens V, Prunus spinosa
Mesquita & alo Sintra Vegetation and Landscape

Daucus ha/ophi/us IV, Car/ina corymbosa varo major IV, Ononis natrix subsp. ramoslsslma II I, I, Ruscus acu/eatus I, Genista toumefortii I, Rosa sempervirens I, Me/ica minuta subsp. arrecta I,
He/ichrysum decumbens II; Companions: Brachypodium phoenicoides V, Cistus sa/vifolius V, Hyacintoides hispanica I, Carex hal/erana I, Asparagus acutifo/ius +, Scil/a monophyl/os +, Carex
Asparagus aphyl/us V, Rubia /ongifo/ia IV, Pu/icaria odora IV, Urginea marítima IV, Scabiosa distachya +, Sel/aginel/a denticulata +; Companions: Cistus salvifo/ius V, Brachypodium phoenicoides
atropurpurea III, Echium tubercu/atum III, Cynara humi/is III , Quercus coccifera II, Erica scoparia II , V, Dacty/is marina IV, Ca/endu/a suffruticosa subsp. a/garbiensis IV, Cistus crispus III , U/ex densus III ,
Juniperus turbinata II, P/antago /anceo/ata II, Ho/cus /anatus II " Schoenus nigricans I, Centaurium Eryngium dila ta tum III, Urginea maritima III, Sedum sediforme III, Daucus halophi/us III, U/ex jussiaei
eríthrea subsp. grandif/orum I, Smi/ax aspera varo aspera I, Convo/vu/us a/thaeoides I, Arisarum vu/gare subsp. congestus II, Lonicera peryc/imenum subsp. hispanica II , Cistus monspe/iensis II, Carlina
I, Lonicera imp/exa I, Romu/ea bu/bocodium I, Rhamnus a/atemus I, A/lium roseum I, Lobu/aria maritima corymbosa varo major II, Anthyl/is vu/neraria subsp. maura II, Antirrhinum majus subsp. /inkianum II ,
I, Allium ampe/oprasum I, Cheir%phus sempervirens I, Pistacia /entiscus I, Phil/yrea angustifo/ia I. Cheir%phus sempervirens II, Thapsia vil/osa II, He/ichrysum decumbens II , Armeria pseudarmeria II ,
Carpobrotus edu/is II, Dítrichia viscosa II, /beris /inifo/ia subsp. microcarpa II, P/antago serraria II ,
Synthetic table of Phlomido /ychchitidis-Brachypodietum phoenicoides Br.-BI., P. Silva & Rozeira
Convo/vu/us altheoides II, Lobu/aria maritima II, Limonium virgatum II, Críthmum maritimum II, Echium
1956 (Festuco-Brometea ercti, Brachypodieta/ia phoenicoides, Brachypodion phoenicoides) from
tubercu/atum II, Cynara humi/is II, Euphorbia port/antica I, Bel/is sy/vestris I, Si/ene /ongicilia I, Erica
CAPELO et alo (1993), 7 relevees: Characteristics: Brachypodium phoenicoides V, Dacty/is hispanica
scoparia I, Rubus u/mifolius I, Cal/una vu/garis I, Ha/imium ca/ycinum I, Armeria we/witschii subsp.
V, Eryngium di/a ta tum V, Daucus crinítus V, Urginea maritima V, P/antago serraria varohispanica IV,
cinerea I, Ulex congestus x densus I, Narcissus bu/bucodium subsp. obesus I, Romu/ea bulbucodium I,
Sa/via sc/areoides IV, Ph/omis /ychnitis IV, Sanguisorba spachiana IV, Convo/vu/us a/thaeoides IV,
Ca/amintha baetica I, U/ex europaeus subsp. latebracteatus +, Prunus spinosa subsp. insititioides +,
Asphode/us /usítanicus IV, Centaurium erythraea subsp. eythraea IV, Anthyl/is vu/neraria subsp. maura
Astraga/us /usítanicus +, /ris subbiflora +, Rosmarinus officinalis +, Phagnalon saxati/e +, Stipa gigantea
IV, Allium roseum IV, Orchis íta/ica IV, Ophrys /utea III, Phagna/on saxati/e III, Aceras anthropophorum
+, Schoenus nigricans +, Ho/cus lanatus +, Plantago /anceo/ata +, Halimum calycinum +, Atriplex
III , Anacamptis pyramida/is III, Serapias parvif/ora II, Allium pal/ens II , Lathyrus amphicarpus II, Ophrys
halimus +, Scabiosa atropurpurea +, P/antago lanceolata t, Suaeda vera +.
fusca II, Nepeta tuberosa II, Ajuga iva II, Mantisa/ca sa/mantica II, Pha/aris caeru/escens II, Bituminaria
bituminosa II, Bel/is perennis II, Romu/ea bu/bocodium II, Hypericum perforatum II, Orchis coriophora Synthetic table 01 Daphno maritimi-U/icetum congesti Rivas-Martínez, T.E. Díaz & J.C. Costa ex
subsp. fragans II, Orchis tenthredinifera subsp. praecox I, Orchis mascu/a I, Spiranthes spira/is I, Orchis J.C. Costa, Espírito Santo, Capelo & Lousã in J.C Costa, Lousã, Capelo, Espírito Santo, Izco & Arsénio
tenthredinifera subsp. tenthredinifera G/adio/us ita/icus I, Ph/eum partense susbp. bert%nii I, Bel/is (Cal/uno-U/icetea, Ulieta/ia minorisi, Dacty/o marítimae-U/icion maritim/) Iram COSTA et alo (2001), 16
sy/vestris I, Fritil/aria /usitanica I, Orchis morio I, Ophrys apifera I, Ophrys sc/opax I; Companions: relevees: Characteristics: U/ex jussiaei subsp. congestus V, Daphne gnidium var. maritima V, Dacty/is
Carex hal/erana V, Pu/icaria odora IV, Cynara hymi/is IV, P/antago /anceo/ata IV, Brachypodium marina V, Carlina corymbosa var. major IV, Armeria pseudarmeria III, Cal/una vu/garis III, Avenu/a
distachyon III, Bup/eurum rigidum subsp. panicu/atum III, Atractylis gummifera III, Scorpiurus su/cata II , U/ex europaeus subsp. /atebracteatus I, U/ex minor I, U/ex /atebracteatus x congestus +,
vermicu/atus III, Asparagus aphyl/us III, Anagallis arvensis III, Pal/enis spinosa 3, Trifo/ium angustifo/ium Simethis mattiazii; Companions: Cistus sa/viifolius V, Daucus halophi/us V, Euph orbia port/antica V,
III , Euphorbia exigua III, Scabiosa atropurpurea III, Hyparrhenia sinaica III, Reichardia intermedia III , Ca/endula suffruticosa subsp. algarbiensis V, Carpobrotus edu/is IV, Brachypodium phoenicoides IV,
Astraga/us hamosus II, Campanu/a rapuncu/us II, Leontodon /ongirrostris II, Bromus /anceo/atus II , Cistus crispus IV, Urginea maritima IV, Stipa gigantea III, Thapsia vil/osa III , Oianthus cintranus subsp.
Linum trigynum II , Onobrychis pedicu/aris II, Avena barbata II , Briza maxima II, Car/ina corymbosa II , cintranus III, Lobu/aria maritima III, Juniperus turbina ta II, Lavandula /uisieri II, Pu/icaria odora II ,
Ca/amintha baetica II, Centaurea pul/ata II, C/inopodium vu/gare II, Origanum virens II, Ononis Asparagus aphyl/us II, Lithodora prostrata subsp. /usítanica II, Eryngium di/a ta tum II, Anagal/is monelli
mitissima II, Medicago minima II, Sherardia arvensis II, Crucianel/a angustifolia II, Ga/ium diva rica tum II, var. microphyl/a II, Holcus /anatus II, Agrostis castel/ana II, Asphode/us /usítanicus II, Rubus u/mifo/ius
Fi/ago gal/ica 1, P/antago afra I, Stachys arvensis I, Carduncel/us caeru/eus I, Lathyrus clymenum I, II , Armeria we/witschii subsp. cinerea II, Crucianel/a maritima II, Quercus pyrenaica I, He/ichrysum
Linum strictum 1, Agrostis st%nifera I, Cynodon dacty/on I, Ho/cus /anatus I, Picris echioides I, decumbens I, Pimpinel/a vil/osa I, Lathyrus c/ymenum I, Centaurea sphaerocepha/a I, Halimium
Schoenus nigricans I. ca/ycinum I, He/ichrysum picardi I, Cistus /adanifer I, Quercus coccifera I, Pteridium aqui/inum I,
Schoenus nigricans I, Coincya pseudeurocastrum subsp. cintrana I, Anthyl/is vu/neraria subsp. maura I,
Synthetic table 01 Querco cocciferae-Juniperetum turbinatae (Rivas-Martínez 1975) Rivas-
Bel/is perennis I, /onopsidium acaule I, Rhamnus alatemus I, Smilax aspera varo aspera I, Cuscuta
Martínez, Lousã, T,E. Díaz, Fernández-González & J.C. Costa 1990 (Quercetea i/icis, Pista cio /entisci-
kotschya I, Anthyl/is gerardi +, Elaeoselinum gummiferum +, Spergu/aria australis +, Arenaria montana
Rhamneta/ia a/atemi, Asparago albi-Rhamnion o/eoidis) Irorn COSTA et alo (2001), 19 relevees:
+, Crithmum marítimum +, Sese/i turtuosum +, Beta vu/garis subsp. marítima t, Ononis ramosissima +,
Characteristics: Juniperus turbinata V, Daphne gnidium varo marítima V, Asparagus aphyl/us V, Smilax
Lotus creticus +, Aetheorhiza bulbosa +, Limonium virgatum +, Sedum sediforme +, /beris we/wítschii +,
aspera varo aspera V, Rubia peregrina varo /ongifo/ia V, Quercus coccifera IV, Rhamnus a/atemus IV,
Scrophu/aria frutescens +, Carex hal/erana +.
Pistacia /entiscus IV, Phil/yrea angustifolia IV, O/ea europaea varo sy/vestris III, Arisarum vu/gare var.
c/usii III, Pu/icaria odora III , Rhamnus o/eoides II, Lonicera imp/exa II, Osyris alba II, Myrtus communis Synthetic table of Salvio scJareoidis-Ulicetum densi ulicetosum densi Rivas-Martínez, Lousã,
II , Euphorbia characias II, Bup/eurum rigidum subsp. panicu/atum II, Phil/yrea media I, Coronil/a g/auca Díaz, Fernandez-González & J.C. Costa 1990 ex Capelo, J.C. Costa, Lousã & Neto 1992
Mesquita & aI. Sintra Vegetation and Landscape

(Rosmarinetea, Rosmarineta/ia, Saturejo- Thymbrion capitatae, Serratu/o estremadurensis-Thymenion Armeria we/witschii subsp. we/witschii V, Pancratium maritimum IV, Ar1emisia crithmifo/ia IV, Ononis
sy/vestris,) from CAPELO et alo (2003), 24 relevees: Characteristics: U/ex densus V, Sa/via sc/areoides natrix subsp. ramosissima IV, Ma/co/mia /ittorea IV, Euphorbia por1/antica IV, Sedum sediforme IV,
V, Eryngium di/a ta tum V, Anthy/lisvu/neraria subsp. maura IV, Oaucus crinitus III, P/antago serraria varo Lotus creticus III, Aetheorhiza bulbosa III, Scrophu/aria frutescens III, /beris procumbens III, Cyperus
hispanica II, Bar1sia aspera I, Serratu/a estremadurensis I, Cardunce/lus caeru/eus I, Phagna/on capítatus III, Linaria caesia subsp. decumbens III, Sese/i tortuosum III, Medicago marina II, Anaga/lis
rupestre I, Ruta cha/epensis I, Staehe/ina dubia +, E/aose/inum gummiferum +, /beris /inifo/ia subsp . mone/li varomicrophy/la II, Mathio/la sinuata II, Si/ene nicaensis II, Carex arenaria I, Herniaria maritima I,
microcarpa r, Va/eriana tuberosa r, Cistus monspe/iensis r; Companions: Brachypodium phoenicoides Herniaria ci/io/ata subsp. robusta +, Leontodon arenarius r; Companions: Carpobrotus edu/is V,
V, Asparagus aphy/lus V, Bup/eurum rigidum subsp. panicu/atum V, Oaphne gnidium IV, Carex Ammophi/la arenaria subsp. austra/is IV, Corynephorus canescens varo maritimus III, Verbascum
ha/lerana IV, O/ea europeae varo sy/vestris IV, Quercus coccifera IV, Pu/icaria odora IV, Centaurium /itigiosum III, Vu/pia a/opecuros II, Ca/endu/a suffruticosa subsp. a/garbiensis II, Senecio ga/licus II,
grandiflorum IV, Rhamnus a/aternus III, Oacty/is hispanica III, Cistus crispus III, Atracty/is gummifera III , Si/ene /ittorea II, Ca/ystegia so/dane/la I, Oacty/is marina I, Lobu/aria maritima I, Reichardia gadítana I,
Carlina corymbosa III , Pa/lenis spinosa III, Cistus sa/vifo/ius III, Urginea marítima III, Rhamnus o/eoides Anchusa ca/carea I, Pimpine/la vi/losa I, Corema a/bum I, Otanthus maritimus +, Eryngium maritimum +,
II , Genista tournefor1ii II, Lonicera imp/exa II, Smi/ax aspera varoaspera II, Asphode/us /usitanicus II, Medicago littora/is +, Críthmum maritimum +, Euphorbia terracina r.
Campanu/a rapuncu/us II, A/lium pa/lens II, Cynara humi/is II, Centaurea pu/lata II, P/antago /anceo/ata
Synthetic table of Limonietum multiflori-virgati J.C. Costa & Capelo in J.C. Costa, Capelo, Lousã &
II , Origanum virens II, Ca/amintha baetica II, Lonicera etrusca II, Ho/cus /anatus II, Rubus u/mifo/ius I,
Espírito Santo 1998 (Críthmo -Limonieteae, Crithmo-Limonieta/ia, Crithmo-Oaucion ha/ophili) from Costa
Myr1us communis I, Rubia peregrina varo /ongifo/ia I, Osyris alba I, Phagna/on saxati/e I, Scabiosa
et aI. (1998), 26 relevees: Characteristics: Limonium virgatum V, Limonium mu/tiflorum V, Oacty/is
atropurpurea I, Ph/omis /ychnítis I, Sanguisorba spachiana I, Oe/phynum pentagynum I, Hypericum
marina V, Críthmum marítimum V, P/antago coronopus subsp. occidenta/is V, Oaucus ha/ophi/us V,
perloratum I, Stachys officina/is subsp. a/geriensis I, Anthy/lis gerardi I, Oiftricchia viscosa I, Foenicu/um
Ca/endu/a suffruticosa subsp. a/garbiensis IV, Spergu/aria austra/is III, Armeria we/witschii subsp.
vu/gare subsp. piperítum I, Me/ica minuta subsp. arrecta +, Pistacia /entiscus +, Anacamptis pyramida/is
cinerea III, He/ichrysum decumbens I, Carlina corymbosa varo major +; Companions: Frankenia /aevis
+, Bítuminaria bituminosa +, Quercus /usitanica r, Nepeta tuberosa +, Lithodora /usitanica +, Cynara
V, Beta vu/garis subsp. maritima V, /nu/a crithmoides IV, Leontodon taraxacoides subsp. taraxacoides
carduncu/us +, Schoenus nigricans +, Echium tubercu/atum +, Erica scoparia +, Convo/vu/us
IV, Limonium feru/aceum III, Lotus creticus II, Parapho/is incurva II, Cruciane/la maritima II, Eryngium
a/thaeoides +, Euphorbia characias +, Cheir%phus sempervirens +, Friti/laria /usitanica +, Lathyrus
di/atatum II, Euphorbia por1/antica II , Sedum sediforme I, Carpobrotus edu/is I, Catapodium maritimum I,
c/ymenum +, Picris echioides +, Arist%chia paucinervis r, Vinca difformis r, Asparagus acutifo/ius r,
Romu/ea bu/bocodium I, E/ymus farctus subsp. boreo-atlanticus +, Lobu/aria maritima +, Ononis natrix
Oipsacus comosus r, Se/lagine/la denticu/ata r, Reichardia picrioides r, Si/ene /ongici/ia r, Serapias
subsp. ramosissima +, Sonchus o/eraceus +, Anthy/lis vu/neraria subsp. maura +, Centaurium spicatum
vomeracea r, Achi/lea ageratum r, Asparagus a/bus r, Ajuga iva r, Oaucus maximus r, Euphorbia
+, A/lium ampe/oprasum +, Mesembryanthemum crista/inum r, Mesembryanthemum nodiflorum r,
amygda/oides r, /ris subbiflora r, Ruscus acu/eatus r, Reichardia gaditana r, U/ex jussiaei r,
Mathio/a sinuata r, Trifo/ium angustifo/ium r.
Cepha/antera /ongifo/ia r, Tammus communis r, Car/ina racemosa r.
Synthetic table of Diantho cintrani-Daucetum ha/ophili J.C. Costa, Capelo, Lousã & Espírito Santo
Synthetic table of Avenu/o su/catae-Stipetum giganteae Capelo, J.C. Costa, Lousã & Espírito
1998 (Crithmo -Limonieteae, Crithmo-Limonieta/ia, Críthmo-Oaucion ha/ophili) from Costa et alo (1998) ,
Santo in. J.C. Costa, Capelo, Lousã & Esp írito Santo 2002 (Stipo giganteae-Agrostietea caste/lanae,
26 relevees: Characteristics: Oaucus ha/ophi/us V, Oianthus cintranus V, Oacty/is marina V, Crithmum
Agrostieta/ia castellanae, Agrostio caste/lanae-Stipion gigantaea) from Costa et alo (2002), 9 relevees:
marítimum V, Limonium virgatum V, Armeria pseudarmeria V, P/antago occidenta/is V, Ca/endu/a
Characteristics: Stipa gigantea V, Agrostis caste/lana V, Brachypodium phoenicoides V, Avenu/a
a/garbiensis IV, Spergu/aria austra/is IV, He/ichrysum decumbens IV, Si/ene mariziana II; Companions:
su/cata subsp. su/cata IV, Stachys officina/is subsp. a/geriensis IV, Arrhenatherum a/bum III , Oacty/is
Frankenia /aevis IV, Euphorbia port/antica IV, Carpobrotus edu/is III , Beta marítima III, Lotus creticus II,
hispanica III, Oacty/is g/omerata subsp. /usitanica III, Asphode/us /usitanicus III, Thapsia vi/losa III,
Lobu/aria maritima II, Sedum sediforme II, /nu/a críthmoides II, Cruciane/la marítima I, Juniperus
Avenu/a su/cata subsp. gadítana I, Festuca durandoi I, Ho/cus annus I; Compan ions: Ho/cus /anatus V,
turbina ta I, A/lium ampe/oprasum I.
Erica scoparia III, Agrostis cur1isii III, Thymus vi/losus III , Lavandu/a /uisieri III, Car/ina corymbosa III,
U/ex jussiaei III, Oianthus cintranus subsp. cintranus III, Quercus /usitanica III, Anthy/lis gerardi II ,
Asparagus aphy/lus II, Briza maxima II, Lagurus ovatus II, U/ex minor II, Brachypodium sy/vaticum II ,
Euphorbia characias II, Coyncia pseudeurocastrum subsp. cintrana II, Armeria pseudarmeria I, References
P/antago afra I, P/antago coronopus I,
BRAUN -BLANQUET, J., PI NTO DA SI LVA, A.R. & ROZE IRA, A. (1956) - Résultats de deux excursions
Synthetic table of Armerio we /witschii-Crucianelletum maritimae Br. -BI., Rozeira & P.Silva in J. & géobotanique à travers le Portugal septentrional & moyen II. Chenaies à feu illes caduques
G. Br.-BI., Rozeira & P. Silva 1972 (Ammophi/etea, Cruciane/leta/ia maritimae, He/ichrysion picardil) (Quercion occidenta/e) et chenaies à feuilles persistentes (Quercion faginae) au Portugal. Agron.
Lusít. 18 (3) : 167-234
from COSTA et alo(2001), 22 relevees: Characteristics: Cruciane/la marítima V, He/ichrysum picardi V,
 Mesquita & ai. Sintra Vegetation and Landscape

BRAUN-BLANOUET, J., BRAUN-BLANOUET, G., ROZEIRA, A. & PINTO DA SILVA, A.R. (1956) - FRANCO, J.A. & M.L. ROCHA AFONSO (1994 -1998 - 2001) - Nova Flora de Portugal (Continente e
Résultats de deux excursions géobotanique à travers le Portugal septentrional & moyen IV. Açores). vollll, fase. 1,2,3. Escolar Editora. Lisboa.
Esquisse sur la végétation dunale. Agron. Lusit. 33 (1-4): 217-234. GÉHU, J.M. & S. RIVAS-MARTíNEZ (1981) - Notions fondamentales de phytosociologie. ln
CAPELO, J., J.C. COSTA, J.C, EspíRITO SANTO, M.D. & LOUSÃ, (1993) - As comunidades Syntaxonomie. 5-33. Ed. J.Cramer. Vaduz.
camefíticas dos calcários do Centro-Oeste Português (Serratulo estremadurensis-Thymenion LOUSÃ, M., EspíRITO SANTO, M.D. & COSTA, J.C. (1994) - A vegetação da Serra de Alvaiázere.
sylvestris, suball. nova). ln Guia Geobotânico das XIII Jomadas de Fitossocilogia: 99-118. I.S. Anais Inst. Sup. Agron. 49 (1): 215-233.
Agronomia. Lisboa. RIBEIRO, M.L., RAMALHO, M.M. (1997) - Notícia Explicativa da Carta Geológica Simplificada do
CAPELO, J., COSTA, J.C. & LOUSÃ, M (1994) - Distribuição das séries de vegetação climatófilas da Parque Natural de Sintra-Cascais. Instituto Geológico e Mineiro/Instituto de Conservação da
região de Lisboa segundo pad rões edáficos e mesoclimáticos. Anais do Inst. Sup. Agron. 44 (1): Natureza.
285-301. RIVAS-MARTíNEZ, S. (1976) - Sinfitosociologia, una nueva metodologia para el estudio dei paisaje
CASTROVIEJO, S. et ai. (ed) (1986-1997) - Flora Iberica. 1, 2, 3, 4, 5, 6, 8. Real Jardín BoI. Madrid. vegetal. Anales Inst. Bot. Cavanilles 30: 69-87.
Madrid. RIVAS-MARTíNEZ, S., DíAZ, T.E., FERNANDEZ-GONZÁLEZ, F., IZCO, J., LOIDI , J., LOUSÃ, M. &
COSTA, J.C., AGUIAR , C. , CAPELO, J. , LOUSÃ, M & NETO, C. (1999) - Biogeografia de Portugal PENAS, A. (2002) - Vascular Plant communities of Spain and Portugal. Addenda to Sintaxonomical
Continental. Quercetea O: 5-56 checklist of 2001 (part I, II). Itinera Geobot. 15 (1 , 2): 5-922.
COSTA, J. C. , CAPELO, J., LOUSÃ, M. & EspíRITO SANTO, M.D. (2002) - Aditamentos à vegetação RIVAS-MARTíNEZ, S., FERNÁNDEZ-GONZÁLEZ, F. & LOIDI, J. (1999) - Checklist of plant
do Sector Divisório Português. Silva Lusit. 10 (1): 119-128. communities of Iberian Peninsu la, Balearic and Canary islands to suballiance leveI. /tinera Geobot.
COSTA, J.C., CAPELO, J. & LOUSÃ, M. (1996) - Os bosques de zambujeiro (Olea europaea L. varo 13: 353-451 .
sylvestris Miller): vegetação potencial dos vertissolos das áreas termomediterrânicas da RIVAS-MARTíNEZ, S., FERNANDEZ-GONZÁLEZ, F., LOIDI , J., LOUSÃ, M. & PENAS, A (2001) -
Extremadura portuguesa. Anais do Inst. Sup. Agron. 44 (2): 497-513. Sintaxomical checklist of vascular communities of Spain and Portugal to association leveI. Itinera
COSTA, J.C., CAPELO, J., LOUSÃ, M. & AGUIAR, C. (1994) - Communautées de Juniperus au Geobot. 14: 5-341.
Portugal. Colloques Phytosoc. 22: 499-526. RIVAS-MARTíNEZ, S., LOIDI, J., COSTA, M., DíAZ, T.E. & PENAS, A. (ed.) - Iter Ibericum A.D. MIM.
COSTA, J.C., CAPELO, J., LOUSÃ, M. & EspíRITO SANTO, M.D. (1998) - Sintaxonomia da vegetação (Excursus geobotanicus per Hispaniam et Lusitaniam , ante XLII Symposium Sociatatis
halocasmofítica das arribas marítimas portuguesas (Crithmo-Staticetea Br.-BI. 1947) Itinera Internationalis Scientiae Vegetationis Bilbao mense lulio celebrandu dicti Anni). /tinera Geobol. 13:
Geobot. 11: 227 -247. 5-347.
COSTA J. C. , CAPELO, J., LOUSÃ, M. & EspíRITO SANTO, M.D. (2002)- Os sobreirais do Divisório RIVAS-MARTíNEZ, S., LOUSÃ, M., DíAZ T.E., FERNANDEZ-GONZÁLEZ, F. & COSTA, JC. (1990)-
Português: Asparago aphylli-Quercetum suberis. Quercetea 3: 81-98. La vegetación dei sur de Portugal (Sado, Alentejo y Algarve) /tinera Geobot. 3: 5- 126.
COSTA, J.C., CAPELO, J., LOUSÃ, M. & EspíRITO SANTO, M.D. (2004) - As comunidades de RIVAS-MARTíNEZ, S., A. PENAS & T.E. DíAZ (2004) - Biogeographic map of Europe. Serviços
Asparago albi-Rhamnion oleoidis Rivas Goday ex Rivas-Martínez 1975 do Divisório Português. Cartográficos da Universidad de Léon .
Quercetea 4: 31-43. SAMPAIO, G. (1947) - Flora Portuguesa. Ed 2. Imprensa Moderna. Porto.
COSTA, J.C., CAPELO, J., NETO, C. , EspíRITO SANTO, M.D. & LOUSÃ, M. (1997) - Notas TEIXEIRA, C. (1996) - A evolução do território português no decurso dos tempos geológicos. Rev.
fitosociológicas sobre os tojais do Centro e Sul de Portugal. Silva Lusit. 5 (2): 275-282. Ped. Cult. 28: 111-157.
COSTA, J.C., EspíRITO-SANTO, M.D., LOUSÃ, M., RODRíGUEZ-GONZÁLEZ, P., CAPELO, J. & TUTIN et ai. (1964-1980) - Flora Europaea (I-V). Cambridge University Press.
ARSENIO, P. (2001) - Flora e Vegetação do Divisório Português (Excursão ao Divisório VALDÉS, B., S. TALAVERA & E. GALlANO (1987) - Flora vascular de Andalucía occidental. Vol. 1-3.
Português). 2º Curso Avançado de Fitossociologia, ALFA, I.S. Agronomia. Lisboa. Ketres. ed. Barcelona.
COSTA, ~.C., EspíRITO-SANTO, M.D., LOUSÃ, M., RODRíGUEZ-GONZÁLEZ, P. , CAPELO, J. &
ARSENIO, P. (2002) - Flora e Vegetação do Divisório Português. Excursão Geobotânica ao
Costeiro Português, Olissiponense e Sintrano. Actas VII Simpósio da Associação Ibero-
Macaronésica de Jardins Botânicos: 249-340. I.S. Agronomia. Lisboa.
COSTA, J.C. , LADERO, M., DíAZ, T.E. , LOUSÃ, M., EspíRITO SANTO, M.D., VASCONCELOS, T.,
MONTEIRO, A. & AMOR, A. (1993) - Guia Geobotânico da Excursão das XIII Jomadas de
Fitossociologia: 1- 98. I. S. Agronomia. Lisboa.
COSTA, J.C., LOUSÃ, M., CAPELO, J., EspíRITO SANTO, M.D., IZCO, J. & LADERO, M., (2001) -
The coastal vegetation of the Portuguese Divisory Sector: dunes, cliffs and low-scrub communities.
Finisterra, XXXV, 69: 69-93.
COUTINHO, A.X. PEREIRA (1939) - Flora de Portugal. Bertrand. Lisboa.
FEIO, M, DAVEAU, S. (2004) - ° Relevo de Portugal - Grandes Unidades Regionais. Volume II
Associação Portuguesa de Geomorfólogos.
FRANCO, J.A. (1971-1984) - Nova Flora de Portugal (Continente e Açores). voll e II. Lisboa.
Quercetea 7: 83-94, 2005
ALFA, Lisboa. Portugal

Vegetation and Landscape 01 Serra de Montejunto

Maria Dalila Esp írito Santo', Ilídio Moreira & Patricia Rodríguez González

I. INTRODUCTION
Hall hundred kilometres northern 01 Lisbon and less than thirty Irom the sea rises the harmonious and
small elevation called "Serra de Montejunto", covering a sutiace with more than 1500 hectares. Its
maximum altitude is 666 meters and it is salient a plateau by the 500 meters altitude (Planalto da Quinta
da Serra). The "Serra de Montejunto" is visited by its agreeable vews and by the monumental ruin s,
specially the "Fábrica do Gelo" (ice lactory) built by the Franciscan monks, which explored it between
the XIII and XVII centuries. It is consisted by a net 01 little deep reservoirs, lilled with water in the cold
nights in winter time; the ice, retired and contained in deep wells supplied with straw, remained even in
the summer time and was transported by night in mules and fluvial boats until the Lisbon palaces.

The "Serra de Montejunto" is a small limestone hill mostly 01 Jurassic origin; the dominant soils are
derived 01 hard calcium carbonate rocks some with dolomitical or marl character. The calcareous soils
are predominantly cambisoils and luvisoils.

The disposition northeast - southeast, relatively near the ocean, causes climatic characteristics
different in the two main slopes. ln the down-hill north-northeast the rainlall is higher (rounding about an
hundred milimiters annually) than in the south-southwest. However the temperature is similar in the two
down-hills, the annual median about the 15ºC, the maximum annual median 19-20ºC and the minimum
annual median 10ºC.

The bio-climatic characterization 01 the Serra de Montejunto, according the terminology 01 Rivas-
Martínez et aI. (2002) , can be considered on the Mediterranean pluviseasonal ocienic bioclimate,
thermo-mesomediterranean belt and upper subhumide-lower humide ombrotype (according with the
altitude and the slope). The meteorological stations 01 Proença (northeast- southwest, 207 m altitude)
and Ota (south-southwest, 41 m altitude) have, respectively, subhumide ombrotype and dry ombrotype,
despite having a thermicity index very closed .

According the biogogeographic map 01 Costa et ai. (2002) and Rivas-Martínez et aI. (2004), "Serra de
Montejunto" is included in Maciço-Estremenho superdistrict, one 01 the six recognised within the West-
Estremenho subsector (Dividing Portuguese Sector, Sadensean-Dividing Portuguese Subprovince,
Western Mediterranean Subregion, Mediterranean Region). Floristically, the West-Estremenho
subsector bears the following endemic taxa: Armeria welwitschii subsp. welwitschii, Coincya monensis

- Instituto Superior de Agronomia, Departamento de Protecção das Plantas e de Fitoecologia, Tapada

da Ajuda, 1349-018 Lisboa; dalilaesanto @isG.Jtl.pt
 Vegetation and Landscape 01 Serra de Montejunto
Espírito Santo & aI.

subsp. cintrana, Dianthus cintranus subsp. barbatus, Limonium mu/tiflorum, Saxifraga cintrana and Vlex
jussiae subsp. congestus. By other side, Anthy/lis vulneraria subsp. maura, Bartsia aspera, Cistus The lower belt on the sandstone soils with Asparago aphyllii-Ouerco suberis S., the midle belt on the
albidus, Deschampsia stricta, Delphynium pentaginum, Fumana thymifolia, Genista tournefortii, Phlomis Iresh valleys 01 the limestone soils with Arisaro-Ouerco broteroi S. or on the south slope with Lonicero
Iychnitis, Prunella X intermedia, Prunella vulgaris subsp. estremadurensis, Ouercus X airensis, Salvia implexae-Ouerco rotundifoliae S.; in the upper belt, on the severely karstiled rocky surfaces, a
sclareoides, Siderites hirsuta, Teucrium polium subsp. capitatum, Vlex densus are territorial differentials permanent community with Ouercetum coccifero-airensis shrubland .
(COSTA et aI. 2002). Some European species , whose Portuguese populations are restricted to this
sector, can also be pointed ou!: Inula montana, Koelleria vallesiana, Orobanche latisquama and
Scabiosa turolensis. The seral vegetation types that predominate are Arisaro-Ouerco broteroi S, II. T H E V EG ET AT I O N OF THE ITINERARY
Asparago aphylli-Ouerco suberis S., Lonicero implexae-Ouerco rotundifoliae S. e Viburno tini-Oleo
The itinerary 01 the day is presented in picture 2. The city 01 Lisbon is located in the right side 01 the
sylvestris S.
Tagus river estuary. Leaving Lisbon by At motoraway the river runs off on our right side. The landscape
The karsic hills are predominant in the "Maciço Estremenho"superdistrict landscape. Ouercus until Vila Franca de Xira town is much humanised and with lew interest. However, ln the edges 01 the
rotundifolia, Ouercus x airensis (O. rotundifolia x O. coccifera), Biarum arundanum, Thymus zygis road remains some species 01 the
subsp. sylvestris, Narcissus calcicola,Vlex airensis are some superdistrict differentials. • O' Viburno tini-Oleetum sylvestris', a
sclerophyllous shrub community
At the Montejunto hill, the climatophyllous woodland vegetation is distributed by three belts (picture t):
composed by thermo-
Mediterranean species, among
666m which wild olive (Olea europaea
varo sylvestris) dominates. This
small tree substitutes the
Ouercus spp. in vertic soils 01 the
Olissiponense superdistrict. As a
permanent sunny community it is
accompanied by other species
such as Ouercus coccifera,
Asparagus albus, Rhamnus
alaternus, Pistacia lentiscus,
Daphne gnidium, characteristic
species 01 Asparago albi-
Rhamnetum oleoidis, association
110m that currently lorms adense
kermes growth and replaces the
Vibumo tini-Oleetum sylvestris.
Viburnum tinus and Coronilla
valentina subsp. glauca, also
Qu ercetum coccifero-airensis
characteristic species, plains 01
colour the borderlines 01 the road
Lonicero il11plexae-Querco rotundifoliae S.
Picture 2: The itinerary Lisbon-Serra de Montejunto-Óbidos- in spring.
(;.'t1>
'.·tr - Arisaro-Querco broteroi S.
Lisboa.

~fiJfç·
• " •• M$ Asparago aphylli-Querco suberis S.
, Ali the synthaxonomy follow lhe RI VAS- MARTíNEZ et ai. (2001) criteria
Picture 1: Zonatioll of th e vegetati on ill SeITa de Montejunto.
Espírito Santo & aI. Vegetation and Landscape 01 Serra de Montejunto

ln the "Serra de Montejunto" are present numerous Orchidaceae included in the Convention 01 the
Near Vila Franca de Xira begins the "Lezíria" (marshy land), the more lertile soils 01 Tagus Iloodplain, International Commerce 01 the Threatened Fauna and Flora Species (CITES Convention) as the
territory 01 horses and bulis. The elm-Iorests (Opopanaco chironii-Ulmetum minoris) and the poplar- following: Aceras anthropophorum (L.) Aiton, Anacamptis pyramidalis (L.) L.C.M. Richard ,
arboreal willow lorests (Salici neotrichae-Populetum nigrae) are represented by very small patches 01 Cephalanthera longifolia (L.) Fritsch; Oactilyorhiza insularis (Sommier & Martelli) Landwehr; Epipactis
Fraxinus angustifolia, Populus nigra and Salix neotricha, which are reduced at lines left by the helleborine (L.) Crantz; Neotinia maculata (Des!.) Stearn; Ophrys bombyliflora Link; Ophrys scolopax
agriculturalland use. Cav.; Ophrys speculum Link; Orchis italica Poiret; Serapias strictiflora Welwitsch ex Veiga; Serapias
parviflora ParI.
ln the town Carregado we left the highway to lollow the E1 national road. The presence 01 cork oak
(Quercus suber) gets our attention. This strip belongs to the mesomediterranean stage where the cork-
oak series Asparago aphylly-Querco suberis sigmetum corresponds to the climatophyllous vegetation. IV. THE VEGETATION OF "SERRA DE MONTEJUNTO "
These siliceous lorests, usually, are rich in lianas, shinny-Ieaved shrubs and shade herbaceous plants,
totally disappeared in the area. The intensive use 01 the land with agriculture and grazing and also with From Abrigada we start the transect till the high part 01 Serra de Montejunto (Nossa Senhora das
eucalyptus lorests and urban areas led to the disappearence 01 characteristic vegetation. Neves, 666 m). During the lirst part we drive across an area where the soils are built on siliceous rocks
(sandstone) and where the vegetation series corresponds to that 01 lhe mesomediterranean cork-oak
Arriving to the Abrigada town , we lace to the Montejunto mountain and it is possible to note the effect lorests (Asparago aphyllii-Querco suberis S.)2. The cork-oak lormalions themselves are rare and
01 the last wildlire, typical 01 the Mediterranean regions. The highest belts 01 vegetation at least in an Quercus suber is lound spread in lhe lerritory. These lorests and their seral stages are usually
hall 01 the slope are now destroyed, but we must to go through the hill to understand how wonderful the submitted to inlense anlhropogenic pressure resulting in a severe degradalion. The florislic combination
recovery 01 the Mediterranean vegetation is after a wildlire. and the relative frequency of species in a relevé, made on the stand of the South-eastern slope, is
shown in the following table:

III. THE FLORA OF "SERRA DE MONTEJUNTO " Asparago aphy/li-Quercetum suberis: Characterislics: Quercus suber 3, Quercus faginea subsp.
brote roi 1, Quercus coccifera 2, Phy/lirea angustifolia 1, Rhamnus alaternus 1, Asparagus aphyllus 1,
By its inclusion in the Annex " 01 the Directive 92/43/CEE, should be enhanced the lollowing species: Oaphne gnidium 1, Lonicera implexa t, Smi/ax aspera t, Pistacia lentiscus t, Myrtus communis t,
Silene longicilia (Bro!.) Otth, Arabis sadina (Samp.) Coutinho, Coincya monensis (L.) Greuter & Burdet Osyris alba t , Rubia peregrina t , Euphorbia characias t , Aristolochia paucinervis t . Companions:
cintrana (P. Cou!.) Franco, Juncus valvatus Link, Narcissus calcicola Mendonça. Cistus salviifolius 2, Rubus ulmifolius 1, Genista triacanthos 1, Ulex jussiaei 1, Lavandula luisieri 1,
On account 01 the situation in its meridionallimit and to be one reliquary species it is mentioned also Calamintha baetica 1, Brachypodium phoenicoides 1, Leuzea conifera t, Carex hallerana t,
the Acer monspessulanum L. Sanguisorba minorsubsp. verrucosa t. (M.D. Espírito Santo &J.C. Costa, near the "Casa das Barreiras
Vermelhas", E, ai!. 210 m, 90 m2)
Besides a reliquary oak lorest and some endemic species 01 the calcareous soils common in the
Centre 01 Portugal, there are taxa, Portuguese endemic or rare, but well represented in the mountain Although lhe rural leatures 01 the landscape are here, in the bottom of the hill, we lound an endemic
which must be mentioned (EspíRITO SANTO et ai. , 1992): Centaurea africana Lam., in populations rush-community, the Juncetum acutifloro-valvati:
with small number of individuais in low lormations 01 Quercus lusitanica; Crambe hispanica L. , Juncetum acutif/oro-va/vati: Characteristics: Juncus valvatus 2, Juncus acutiflorus 2, Scirpoides
Brassicaceae, in Portugal, only known in "Maciço Calcáreo Estremenho"; Helianthemun viola ce um holoschoenus 2, Phalaris caerulescens 2, Carex flacca subsp. serrulata 2, Cynodon dactylon 1.
(Cav.) Pers., only known, in Portugal, in Montejunto and "Aire e Candeeiros" hills; Hesperis laciniata Ali ., Companions: Lythrum junceum 2, Mentha suaveo/ens 1, Polypogon monspeliensis 1, Juncus bufonius
lew Irequent and in reduced populations; Hornungia petraea (L.) Reichenb., characteristic 01 the pioneer 1. (M .D. Espírito Santo, C. Pinto Cruz & P. Arsénio, E of "Malhadas", 110m, 4 m2)
association 01 annual herbaceous pre-colonizers 01 calcareous rocks; Koeleria vallesiana (Honckney)
Gaudin, only lound in West-Centre calcareous; Leucanthemun sylvaticum (Hollmanns & Link) Nyman, When we begin to go up limestone soils appear. ln a place called Furadouro it is loreseen the first
iberian endemic in the grove margins in calcareous soils; Saxifraga cintrana Willk., endemic 01 the stop (Picture 3). Here, in edaphohygrophyllous conditions, is represented one of the most original and
West-Centre; Senecio lagascanus DC. subsp. lusitanicus (P. Cout.) P. Silva, endemic 01 the West- characteristic vegetation types of the Maciço-Estremenho superdistrict; it is the "carvalhal" (beech oak
Centre calcareous and actually only known in "Serra de Montejunto"; Senecio minutus (Cav.) DC., forest) of the association Arisaro-Quercetum brote roi, together with some of its seral stages such as
iberian endemic annual plant, rare. "carrascais" (grove of holly oak: Me/ico arrectae-Quercetum cocciferae) , "medronhais" (strawberry-tree

2 The habitats of "Serra de Montejunto" were described in PINTO DA C RUZ & EspíRITO S ANTO, 1999.
 pt

Vegetation and Landscape 01 Serra de Montejunto

Espírito Santo & aI.

grove:. Bup/euro fruticosae-Arbutetum unedonis) , "giestais" (broom-lield: Erico scopariae-Cytisetum Phlomido /ychnitidis-Brachypodietum phoenicoidis: Characteristics: Brachypodium phoenicoides
grandtflon) and dry grasslands (Ph/omido /ychnitidis-Brachypodietum phoenicoidis). These communities 5, Oacty/is glomerata subsp. hispanica 2, Sa/via sclareoides 2, Campanu/a rapunculus +, Eryngium
occupy the mld-slope ln well drained deep soi ls and are represented in the lollowing tables: dila ta tum +, Phlomis Iychnitis +, Plantago lanceo/ata +. Companions: Urginea maritima 1, Carex
hal/erana 1, Cynara humilis 1, Bupleurum rigidum subsp. paniculatum +, Pulicaria odora +, Scorpiurus
Arisaro-Quercefum broferoi: Characteristics: Quercus faginea subsp. broteroi 4, Smi/ax aspera 3, vermiculatus +, Anagal/is mone/li +, Anthyl/is vulneraria subsp. maura +, Avenula sulcata subsp.
Quercus suber 2, Arbutus unedo 2, Myrtus communis 2, Asp/enium onopteris 2, Quercus coccifera 1, occidentalis +, Carduus broteroi +, Carlina corymbosa +, Centaurea pul/ata subsp. baetica +,
Rosa semperVf(ens 1, Phyl/yrea /atifo/ia 1, Phyl/yrea media 1, Rhamnus a/atemus 1, Lonicera etrusca + Sanguisorba minor subsp. verrucosa +, Urginea maritima +. (M. D. Esp írito Santo & J.C. Costa,
O/ea europaea varosy/vestris +, Phyl/yrea angustifo/ia +, Pistacia /entiscus +, Vibumum tinus +, Ruscu~
Furadouro, E, 250 m, 10 m2)
acu/.e~tus +, Hedera maderensis subsp. iberica+, Rubia peregrina +, Euphorbia characias +, /ris
foet~d/sstma +, Ansarum vu/gare +. Companions: Crataegus mongyna subsp. brevispina 1, Rubus
u/mlfo~/us 1, Lomcera peryc/imenum subsp. hispanica 1, Po/ypodium austra/e 1, Brachypodium The clearing community Leucanthemo sylvatici-Cheirolophetum sempervirentis is also a remarkable
sy/vat/cum 1, Carex d/st~chya 1, Tamus communis +, Arist%chia paucinervis +, Ca/amintha baetica+,
characteristic 01 these woodland ecosystems:
Luzu/a forsten +, Teucnum scorodonia +, Umbi/icus rupestris +. (M.D. Espírito Santo & C. Pinto da
Cruz, Furadouro, SE, 230 m, 100 m2). Leucanthemo sy/vatici-Cheirolophetum sempervirentis: Characteristics: Cheirolophus
sempervirens 3, Leucanthemum sy/vaticum 2, Teucrium scorodonia 2, Clinopodium vulgare subsp.
Bu~/euro truticosae-Arbutetum unedonis: Characteristics: Arbutus unedo 3, Laurus nobi/is 2, arundanum 1, Calamintha baetica 1, Origanum virens +, Stachys germanica subsp. lusitanica +,
Phy/llrea /aMolla 2, Rhamnus a/atemus 2, Quercus coccifera 2, Quercus suber 1, Quercus faginea Scrophularia scorodonia +, Centaurea sphaerocephala subsp. lusitanica +, Si/ene latifolia +, Agrimonia
subsp. brotero/ 1, Smt/ax aspera 1, Phy/lirea angustifo/ia 1, Pistacia /entiscus 1, Erica arborea 1, eupatoria +. Companions: Cistus salviifo/ius 2, Anthirrhinum majus subsp. linkianum 1, Coronil/a
Vtbumum t/nus 1, Rosa sempervirens 1, Rubia peregrina +, Lonicera etrusca +, Lonicera peryc/imenum va/entina subsp. glauca 1, Sanguisorba rupicola 1, Oigitalis purpurea 1, Rubia peregrina+, Aristolochia
subsp. peryc/tmenum +, Myrtus communis +, Osyris alba +, Ruscus acu/eatus +, Coronil/a va/entina paucinervis +, Brachypodium phoenicoides +, Carex hal/erana +, Si/ene longicilia +, Thapsia vil/osa +,
subsp. g/auca +, Oaphnegnidium +, Asparagus aphyl/us +, Arisarum vu/gare+, Asp/enium onopteris +. Sa/via sclareoides +, Arabis sadina +, Biscutel/a lusitanica +, Euphorbia characias +, Geranium
Companlons: Rubus u/m/fo//Us 1, Crataegus monogyna subsp. brevispina 1, U/ex airensis +, Erica purpureum +, Pteridium aqui/inum +, Carlina corymbosa +, Lathyrus c/ymenum +. (M.D. Espírito Santo
scopana +, Chetr%phus sempervirens +, Teucrium scorodonia +, Tamus communis +, Cistus
& J.C. Costa, Furadouro, E, 250 m, 20m 2)
sa/vllfo//Us +, Brachypodium phoenicoides +, Urginea maritima +, Po/ypodium austra/e +. (M.D. Espírito
Santo & I. Moreira, Furadouro, E, 250 m, 100 m2) The chasmochomophyte vegetation colonizing the vertical calcareous walls, in earth lilled and
sciophilous lissures, belongs to the Sileno longiciliae-Antirrhinetum /inkiani association; on the small
Melico arrecfae-Quercetu~ cocciferae: Characteristics: Quercus coccifera 4, Arbutus unedo 2, platforms , where there is soil deposition and humidity enough it is developed the Polypodietum serrati;
V/bum um t/nus 2, Lomcera /mp/exa 2, Smi/ax aspera 2, Rubia peregrina 1, Phyl/irea angustifo/ia 1, on the earthy banks , under sciophilous conditions, dominate Selaginel/a denticulate (Selaginel/o
Phy/llrea /aMolla 1, Laurus nobi/is 1, Me/ica minuta subsp. arrecta. Companions: Rubus u/mifo/ius 1
denticulatae-Anogrametum leptophyl/ae):
Tamus communis 1, Car/ina corymbosa subsp . hispanica +, Picris spinifera +, Arrenatherum a/bum +:
A/llum roseum +, Sanguisorba minor subsp. verrucosa +, Avena steri/is subsp. /udoviciana+, Si/eno longiciliae-Antirrhinetum linkiani (Ladero et aI. 1991, synthetic table): Characteristics:
Urspermum p/cro/des +, Brachypodium distachyon +, Leontodon taraxacoides subsp. /ongirostris +, Antirrhinum majus subsp. /inkianum 5, Calendula suffruticosa subsp. lusitanica 4, Sanguisorba
Geramum d/ssectum+. (M. D. Espírito Santo & J.C. Costa, near Fu radouro, SW, 230 m, 60 m2) multicaulis 4, Galium fruticescens 4, Biscutel/a lusitanica 3, Melica minuta 3, Silene /ongicilia 3, Avenula
occidentalis 2, Arabis sadina 2, Ceterach officinarum 2, Lactuca viminea subsp. chondril/iflora 2,
Erico scopariae-Cytisetum grandiflori: Characteristics: Cytisus grandiflorus 2, Pteridium aqui/inum Phagnalon saxatile 2, Umbificus ruspestris 2, Bituminaria bituminosa 2, Asplenium ruta-muraria 1,
3, Enca scopana 2, Enca arborea 1. Companions: Rubus u/mifolius 2, Pinus pinaster 1, Quercus Asp/enium trichomanes subsp. quadrivalens 1, Coincya monensis subsp. cintrana 1, Crambe hispanica
cocc/fera 1, P/stac/a /ent/scu.s 1, Lonicera imp/exa 1, Rosa sempervirens 1, Coronil/a va/entina subsp. 1, Polypodium australe subsp. serrulatum 1, Rumex intermedius subsp. /usitanicus 1, Saxifraga cintrana
g/auca 1, Gemsta toumefortll 1, Cistus sa/viifo/ius 1, Cheir%phus sempervirens +, Origanum vu/gare +, 1, Selaginel/a denticulata 1, Scabiosa turolensis 1. Companions: Sedum album 2, Melica ciliata subsp.
Hypencu~ perforatum +, Ansarum vu/gare +, Arum italicum +, Arist%chia paucinervis+, Silene la tifo/ia magnolii 2, Geranium purpureum 2, Sedum sediforme 2, Oactylis glomerata 1, Lagurus ovatus 1,
+, Senec/O jacobaea +, Bel/is perennis +, Asplenium onopteris +, Geranium purpureum +, Cerastium Anthyl/is vulneraria subsp. maura 1, Hyparrhenia sinaica 1, AI/ium pal/ens 1, Rhamnus alatemus 1,
glomeratum +, Coleostephus myconis +, Rhagadiolus edulis + (M .D. Espírito Santo & J.C. Costa, near Hedera maderensis subsp. iberica 1, Brachypodium distachyon 1, Blakstonia perfoliata 1, Linaria supina
Furadouro, E, 250 m, 40 m2) 1, Avena barbata 1, Sedum forsteranum 1, Crepis vesicaria subsp. haenseleri 1, Arrhenatherum a/bum
1, Torifis nodosa 1, Sideritis hirsuta 1, Sedum brevifolium 1, Hyacinthoides hispanica 1, Piptatherum
Espírito Santo & aI. Vegetation and Landscape 01 Serra de Montejunto

miliaceum 1, Cheirolophus sempervirens 1, Campanula erinus 1, Iberis procumbens subsp. microcarpa Bituminaria bituminosa +, Rumex intermedius subsp. lusitanicus +, Allium pallens subsp. pallens +,
1, Clinopodium vulgare 1. Ophrys sc%pax subsp. sc%pax +, Serapias parviflora +, Oaucus carota +, Gastridium ventricosum +,
Geranium rotundifo/ium+, Sonchus asper subsp. asper +, Lathyrus sphaericus +, Malva hispanica +,
Po/ypodietum serrati: Characteristics: Polypodium australe 1, Briólitos 3, Asplenium trichomanes 1, Campanu/a erinus +, Po/ycarpon tetraphyllum subsp. diphyllum +, Evax pygmaea subsp. ramosissima
Asplenium ceterach 1, Selaginella denticulata +, Umbilicus rupestris +; Companions: Hyacinthoides +, Centaurium erythraea subsp. grandiflorum+, Medicago minima +, Vicia laxiflora +, Vicia tenuiflora +,
hispanica 1, Sedum album +, Saxifraga granulata +. (M .D. Espírito Santo & J.C. Costa, Furadouro, Linum trigynum subsp. tenue +, P/antago bellardii subsp. bellardii. (M.D. Espírito Santo & J.C. Costa,
vertical calcareous wall, in platforms, E, 250 m, 20 m2).
south-western 01 "Moinho do Céu", S, 460 m, 100 m2)

The last wildlire lavours the development 01 a scrub with heather and the endemic gorse U/ex airensis:

U/iei airensis-Erieetum seopariae: Characteristics: U/ex airensis 3, Erica scoparia 2, Genista

triacanthos 1, Lithodora prostrata subsp. /usitanica 1, Serratu/a estremadurensis +. Companions: Cistus
sa/viifo/ius 2, Pinus ha/epensis 1, Quercus coccifera 1, Smi/ax aspera 1, Oaphne gnidium1 , Cistus
crispus 1, Lavandu/a /uisieri 1, Quercus faginea subsp. broteroi fruto +, Quercus rotundifolia fruto +,
Lonicera etrusca +, C/inoprxlium vu/gare subsp. arundanum +, Pistacia /entiscus +, Myrtus communis +,
Cuscuta p/aniflora +, Arist%chia paucinervis +, Po/yga/a monspe/iaca +, Urginea marítima +, Carduus
broteroi +, Brachypodium phoenicoides +, Oacty/is g/omerata subsp. hispanica +, Sa/via sc/areoides +,
Prunella x intermedia +, Chamae/eon gummifer +, Jasione montana subsp. montana +. (M .D. Espírito
Santo &J.C. Costa, between Ramada house and foreslry guard house, W, 300 m, 100 m2)

The seral grassland, in these dried and termophilous conditions, is the Carici depressae-
Hyparrhenietum sinaicae, a meadow that can be also seral 01 the Arisaro-Quercetum broteroi:

Cariei depressae-Hyparrhenietum sinaieae: Characteristics: Hyparrhenia sinaica 3, Oaucus crinitus

2, Oacty/is g/omerata subsp. hispanica 2, Phagna/on saxati/e 1, Arrhenatherum a/bum 1, Ajuga iva 1,
Sa/via sc/areoides 1, Bituminaria bituminosa 1. Companions: Piptatherum mi/eaceum 1, Foeniculum
vulgare subsp. piperitum 1, Brachypodium phoenicoides 1, Urginea maritima 1, Myrtus communis +,
Ca/endula suffruticosa subsp. /usitanica +, Sedum album +, Sedum sediforme +, Oiftrichia viscosa
subsp. viscosa +, Astragalus /usitanicus +, G/adiolus illyricus +, Leuzea conifera +, Carlina corymbosa
subsp. hispanica +, Picris spinifera +, Si/ene /ongicilia +, Biscutella /usitanica +, Bup/eurum rigidum
subsp. panicu/atum +, Scorzonera graminifo/ia +, Vicia bengha/ensis+, Briza maxima +, Campanu/a
Picture 3: 1- Arisaro-Quercetum broteroi, 2- Bupleuro fruticosae-Arbutetum unedonis, 3- Melico erinus +, Brachypodium distachyon +, Sonchus o/eraceus +, Urospermum picroides +, Capsella rubella
arrectae-Quercetum cocciferae, 4- Erico scopariae-Cytisetum grandiflori, 5- Leucanthemo sylvatici-
+, Hornungea petraea +, Th/aspi perto/iatum +. (M.D. Espírito Santo & J.C. Costa, between the house 01
Cheirolophetum sempervirentis, 6- Sileno longiciliae-Antirrhinetum linkiani, 7- Polypodietum serrati.
"Barreiras Vermelhas" and Furadouro, S, 250 m, 10 m2)

The low-scrubland, often Irequent under hard conditions, is a community 01 Teucrio capitati-Thymetum
On lhe same belt, but in the south slope, where the presence 01 karst also contribute to the deliciency sy/vestris, an open lormation established on the limestone crests ali over the hill; it can be considered a
01 water, we can still appreciate a small Iragment 01 the holm oak lorest (Quercus rotundifolia) : seral stage 01 Lonicero imp/exae-Querceto rotundifo/iae S.:
Lonieero imp/exae-Quereetum rotundifo/ia: Characteristics: Quercus rotundifolia 3, Quercus Teuerio eapitati-Thymetum sy/vestris: Characteristics: Thymus zygis subsp. sylvestris 3, U/ex
coccifera 2, Olea europaea subsp. sylvestris 1, Jasminum fruticans 1, Rubia peregrina 1, Smilax aspera airensis 3, Teucrium po/ium subsp. capitatum 2, He/ianthemum vio/aceum 2, Siderites hirsuta 1.
1, Euphorbia characias 1, Brachypodium phoenicoides 1. Companions: Cistus salviifolius 3, Ca/endu/a Companions: Oacty/is g/omerata subsp. hispanica 3, Erygium di/ata tum 1, Bup/eurum rigidum subsp.
suffruticosa subsp. lusitanica 2, Pinus ha/epensis 1, Rosmarinus officina/is 1, Urginea maritima 1, paniculatum 1, Asparagus acutiflorus 1, Carlina corymbosa 1, Carduus brote roi 1, Chamaeleon
Oacty/is g/omerata var. hispanica 1, Centaurea pullata subsp. baetica 1, Sa/via sc/areoides 1, gummifer 1, O/ea europaea var. sy/vestris +, Sedum a/bum+, Melica magnolii subsp. ciliata +, Allium
Ca/amintha baetica 1, C/inopodium vulgare subsp. arundanum 1, Lathyrus sy/vestris 1, Sedum album +,
 Espirito Santo & aI.
Vegetation and Landscape 01 Serra de Montejunto
pal/ens +, Urginea marftima +, Avenu/a su/cata subsp. occidenta/is +, Sa/via sc/areoides +, Stachys
Companions: Cistus sa/viifo/ius 3, He/ianthemum vio/aceum 1, Rosmarinus officinalis 2, U/ex airensis 2,
germamca subsp. /usilamca +. (M.D. Espírito Santo & C. Pinto da Cruz, N 01 "Casal do Chorão" SE
360 m, 50 m2) , , Stipa gigantea 1, Carex ha/lerana 1, Ca/endula suffruticosa subsp. /usitanica 1, Anthirrinum majus
subsp. linkianum 1, Hyacinthoides hispanica 1, Ga/ium fruticescens+, Polypodium austra/e +,
Often, thH seral vegetation 01 the holm oak lorest has a scrub 01 Quercus coccifera and Quercus x Clinopodium vu/gare subsp. arundanum +, He/leborus foetidus +, Urginea maritima +, Brachypodium
alrensls ln Its stages. Quercus x airensis is a hybridum between Quercus coccifera and Quercus phoenicoides +, Cepha/anthera /ongifo/ia +, Epipactis /usitanica+, Thapsia vi/losa var. p/atyphy/los+,
rotundlfolia very Irequent in the limestone hills on the Central 01 the country. On the upper parl 01 the Si/ene /ongici/iat, Geranium purpureumt. (M.D. Espírito Santo & J.C. Costa, on the lop, 660 m, 40 m2)
hlll, where the edaphlc condltlons 01 d.ryness, lack 01 soil and strong wind prevail, there is no possibility
The scarce conditions 01 lhe sai I permits lhe development 01 a varianl 01 lhe thyme low-scrub lhal
to can occur woodland. ln such condltlons the Quercetum coccifero-airensis is a secondary lares!. Very
forms a mosaic with lhe "carrascal"; rich in endemic species ii is eslablished in lhe earlh depressions:
well adapted to the wlidllres and, ln most cases, in more ar less Ilat topographical positions, is parl 01 a
vegetatlon mosalc determlned by the erosion 01 the soil. That is our 2nd stop (Picture 4): Teucrio capitati·Thymetum sylvestris helianthemetosum vio/aceae Capelo & Espírilo Santo
subassoc. nova: Characteristics: Helianthemum vio/aceum 3, Teucrium po/ium subsp. capitatum 2,
Rosmarinus officinalis +, Avenu/a occidentalis +, Senecio doronicum subsp. /usitanicus t , Anthy/lis
vu/neraria subsp. maura +. Companions: Cistus sa/viifo/ius 2, Ca/endu/a suffruticosa subsp. /usitanica 2,
Hedera maderensis subsp. iberica1 , Centaurea sphaerocepha/a subsp. /usitanica 1, Hyacinth oides
hispanica 1, Urginea maritima 1, Dacty/is g/omerata 1, Asphode/us ramosus 1, Carduus brote roi t ,
Thapsia vi/losa t , Narcissus bu/bocodium t, Ornithogalum ortophy/lum subsp. baeticum, Orchis
mascula t , Aceras antrpophora + Pu/icaria odora +, Sedum a/bum +, Sedum forsteranum t , Sedum
sediforme +, Biscutel/a lusitanica t, Euphorbia panicu/ata subsp. we/witschii t, Saxifraga cintrana t,
Silene longici/ia t , Rumex intermedius subsp. lusitanicus t , Ranunculus o/issiponensis subsp.
olissiponensis t, Sanguisorba minorsubsp. verrucosa t , Centranthus ca/citrapaet, Omphalodes /inifolia
t, Arenaria conimbricensis t, Leontodon taraxacoides subsp. longirostris t, Alyssum simplex t,
Euphorbia exigua t, Briza maxima t, Desmazeria rigida t, Ononis pusil/a t, Ononis repens t. (J.
Capelo & M.D. Espírilo Sanlo, on lhe top, 660 m, 40 m2)

The annual paslures , wilh a poor vegetalion cover, have been allered along lhe lime by lhe
nilrilicalion caused by lhe anlhropogenic inlluence and by the wildlires. The Anthyl/ido lusitanicae -
Brachypodietum distachii is annual grassland endemic of lhe Porluguese limeslones:

Anthy/lido /usitanicae . Brachypodietum distachii: Characlerislics: Brachypodium distachyon 3,

Leotodon taraxacoides subsp. longirostris 1, Trifolium campestre 1, Tuberaria guttata 1, Anthyl/is
vu/neraria subsp. /usitanica 1, Trifolium scabrum 1, Vu/pia ciliata 1, Vulpia myuros subsp. sciuroides t ,
Aira caryophyl/ea t, Campanula erinus t, Ononis pusil/a t, Galium diva rica tum t, Linum strictum t ,
Linum trigynum t, Trifo/ium stel/atum t, Blackstonia acuminata t , B/ackstonia perfo/iata subsp.
perfo/iata t, Centaurium erythraea subsp. erythraea t , Desmazeria rigida t, Aira caryophyl/ea subsp .
Picture 4: l - Quercelul1I cocclfero-airellsis, 2- Tellcrio capilati-Thymellll1J sylveslris helianlhemelosu/1J. caryophyl/ea t , Medicago minima t , Petrorhagia nanteui/ii t, P/antago bel/ardii t. Companions:
vlOlaceae , 3- Phlo/1Jldo lychl1llldls-Brachypodiellllll phoenicoidis 4- Anlhy l1id I '1 . B· I I' Centranthus calcitrapae 1, P/antago serraria 1, Anagal/is arvensis 1, Leuzea conifera t, Euphorbia
disla chii. ' o II S 1 al1lcae- w c lypoc leIU/1J.
por/andica t, Cerastium g/omeratum +, Silene gal/ica t , Medicago trunca tu/a t, Sonchus o/eraceus t,
Urospermum picroides t, Geranium purpureum t, Avena barbata subsp. barbata t, Aegilops neglecta
t , Bromus madritensis t , Bromus rubens t, Cynosurus echinatus +, Gaudinia fragi/is t , Lagurus ovatus
Quercetum coccifero-airensis: Characteristics: Quercus coccifera 4, Quercus x airensis 2, Quercus t. (M.D. Espírito Santo & C. Pinlo da Cruz, near lhe "Moinhos do Barreiro Pequeno", W, 340 m, 16 m2).
rotundlfolta fruto +, Rhamnus a/aternus 2, Smi/ax aspera 1, Asp/enium onopteris 1, Euphorbia characias
+, Rubla peregrma +, Osyns alba +, Genista triacanthos +, Scil/a monophyl/us +, Anemone pa/mata +. Above "Nossa Senhora das Neves" we climb to the valley 01 Vila Verde dos Francos. The conlrast in
lhe vegelalion on ils norlh-western lace is nol very high, because of lhe elevaled human dislurbance.

92
93
 Espírito Santo & aI.

Quercetea 7: 95-122, 2005

ALFA, Lisboa, Portugal

REFERENCES
The vegetation of Madeira Island (Portugal). A brief overview and
COSTA, ~.C., EspíRITO SANTO, M.D., LOUSÃ, M., RODRíGUEZ GONZÁLEZ P.M., CAPELO, J. &
ARSENIO, P. (2001) - Excursão ao Divisório Português. Actas do VII Simpósio da Associação excursion guide
(bero-Macaronésica de Jardins Botãnicos: 249-340. I.S. Agronomia. Lisboa.
ESPIRITO-SANTO, M.D. , LADERO, M. & LOUSÃ, M. (1995) - Comunidades Rúpicolas do Parque
~atural das Serras de Alre e Candeeiros. Studia Botanica 14: 13-22. Jorge Capelo' , Miguel Sequeira**, Roberto Jardim"', Sandra Mesquita···· & José Carlos Costa****
ESPIRITO SANTO, M.D. , PROENÇA, L., COSTA, J.C. & VASCONCELOS T. (1992) - Flora da Serra
de Montejunto. DBEB, ISA. Lisboa. 68pp.
LADERO, M., VALLE, C.J. , SANTOS, M.T., AMOR, A. , EspíRITO SANTO, M.D. , LOUSÃ, M. & Foreword
COSTA, J.C. (1991) - Sobre la végetación y Ilora rupícola de las intercalaciones calcareas de los
sectores Divisório portu.guês y Beirense litoral. Candollea 46(1): 53-59. This text aims to be a briel overview 01 vascular plant communities lound in Madeira Island in the
PINTO CRUZ, C. & ESPIRITO SANTO, D. (1999) - Habitats naturais da Serra de Montejunto. perspective 01 syntaxonomy, but encompassing also leatures 01 structure, composition, biogeography,
Quercetea 1: 103-116.
and role as ecosystem/landscape elementary units, General traits 01 soil , bioclimate, plant communities
RIVAS-MARTíNEZ, S., FERNÁNDEZ-GONZÁLEZ, J., LOIDI, J., LOUSÃ, M. & PENAS, A. (2001) _
and respective successional relationships are brielly stated, locusing on specilic sites or transects that
Syntaxonomlcal Checkllst 01 Vascular Plant Communities 01 Spain and Portugal to association
leveI. !tinera GeobotaniCa 14: 5-341. will illustrate madeiran vegetation during the excursion, The text stands as the english companion to the
RIVAS-MARTíNEZ, S., DíAZ, T.E. , FERNÁNDEZ-GONZÁLEZ, F., IZCO, J., LOIDI, J., LOUSÃ, M. & more comprehensive podromus lound in CAPELO (ed,) (2004) - A paisagem vegetal da Ilha da
PENAS, A.. (2002) -. Vascular Plant Communities 01 Spain and Portugal. Addenda to the Madeira. Quercetea 6: 3-200 [The plant landscape 01 Madeira Island] written in the portuguese
syntaxon~mlcal checkllst 01 2001. !tinera G,eobotanica 15(1): 5-432 language, Extensive phytosociological tables, syntaxonomical catalogues, Iloristic and bioclimate data
RIVAS-MAR,TINEZ, S" A PENAS & T.E. DIAZ (2004) - Biogeographic map of Europe, Serviços should also be consulted therelrom, Along the text, relerences to pages or tables in the lormer are
Cartograllcos da Umversldad de Léon,
noted by [rectangular brackets].

Origin and evolution of madeirean flora and vegetation

The islands 01 the Madeiran archipelago lie in the Atlantic Ocean about 978 km southwestwards 01
Lisbon and about 630 Km 011 the atlantic coast 01 Morocco, in the latitudes 33º 10' - 33º 20' N and the
longitudes 16º 10' - 17º 20' W. Being Madeira the largest island (728 square kms) , it reaches 1861 m
above sea levei, The geological origin 01 Madeira lies in the Mid-Tertiary volcanic cycles that gave rise
to the arch 01 atlantic archipelagos spreading Irom 40º to 15º N along the coasts 01 Europe and Alrica,
inlormally known as Macaronesia ('1he Fortunate Islands"): Azores, Madeira, Salvage Islands, Canary
Islands and Cape Verto ln Madeira, the oldest volcanic lormations date Irom the Miocene - 5 to 6 million
yrs, B,P,

• Departamento de Ecotogia, Estação Florestal Nacional. Quinta do Marquês 2780-159 Oeiras. Portugal.
jorge.capelo @efn.com.pt
Deptarmento de Biologia, Universidade da Madeira, Campus da Penteada, 9000 Funchal. Portugal.
. clqueira@ uma.pt
••. Jardim Botânico da Madeira. Funchal. Portugal. robertojardim.sra @gov-mad.pt
•••• Departamento de Protecção de Plantas e de Fitoecologia. Instituto Superior de Agronomia. Tapada da Ajuda
1349-017 Lisboa, Portugal. mesquita.sandra @sapo.pt; jccosta@isa.utl.pt
 The vegetation 01 Madeira Island (Portugal)
Capelo, J. & aI.

The sequential setting 01 the archipelagos and their approximately parallel position in relation to the Sonchus, Perical/is) , Boraginaceae (Echium) , Campanulaceae (Muschia in Madeira, Azorina and
alrican and european mainland are paramount to hypothesis explaining actual "macaronesian" Ilora and Canarina in the Azores and Canary Islands respectively), Scrophulariaceae (Isoplexis) and others such
vegetation leatures, namely colonization, dispersion and speciation events. as Sldentls, Plantago, Aeonium and Euphorbia. These plants exhibit consistent habit and structure
leatures, namely being cauli rosetted (Iong thin woody stems with a single rosette 01 leaves at the top) ,
Traditionally, the extremely original and highly endemic vegetation 01 these islands has been
havlng candelabra hablt or belng monocarpic (plants with a base rosette, dying alter seeds are
interpreted as being basically 01 relictual character Irom the sub-tropical Tertiary vegetation around the
produced). The majority 01 their continental relatives are small herbaceous or even annual plants. These
archaic basin 01 the Tethys Ocean - Artho-Tertiary vegetation s.l. (coarsely the Mediterranean Sea,
plants are lound in peculiar lorest habitats, namely where higher Lauraceae trees are temporarily
spreading eastwards to include the actual Black and Arai seas, reaching the Indian Ocean). ln
removed by natural landslldes or whenever natural openings in the canopy occur ('1rump" species) .
continental areas around the Tethys, several violent disruptive environmental events took place during
Some early authors regard such taxa as relicts Irom the artho-tertiary flora, but molecular and
the mid and late Tertiary period, with dramatic elfects on this vegetation. Tectonic movements 01 the
phylogeographical evidence is summing up to support their derived neo-endemic character. Carlquist
Alrican plate northwards intensilied alpine orogeny, giving rise to both the Mediterranean Sea and
(1974) named thls evolutlonary tendency island woodiness and reported it as universal phenomena in
Alpine geo-synclinal ranges. Several lactors related to the settling 01 a new closed circulation regime 01
most Islands. Therelore, such taxa should be regarded as having continental herbaceous ancestors
oceanic currents in the North Atlantic , gave rise to the regular intra-annual oscillation 01 the system 01
coming Irom early colonization events and alterwards developing woodiness traits and correlativ~
lows, associated with rain storms coming Irom the Atlantic, along with the solidary southernmost sub-
si~apomorphic characters. Molecular studies show that island clades 01 such genera are, in general,
tropical highs (e.g. Azores High). Thus , typical "summer dry-winter rain" Mediterranean bioclimate was
stnctly monophyletic. This implies a single ancestral population to reach the islands - a unique
established at the time. Massive geological and climate disturbance lead to great reduction 01 sub-
colomzat/On event - lollowed by extensive adaptative radiation and inter-island dispersion. Taking on
tropical tethyan vegetation . Moreover, new habitats and migratory pathways were created both lor
account parclmonlous phylogenetlc reconstructions and island ages , it is also possible to trace back the
continental (neo-mediterranean) and paleo-boreal deciduous Iloras. As an example, in the
best extant candidate to taxonomical ancestor 01 groups. ln most groups, circum-mediterranean origin is
Mediterranean area a major disturbance event, known as Messinian Salinity Crisis (Miocene) settled
pos.tulated but some larther ancestors are also lound (american, in genus Perical/is and Bystropogon,
alternate periods 01 marine and dry or lacustrine conditions , due to closing-opening cycles in the
l0r. Instance). Evolutlonary mechanisms lor island woodiness are yet to be se\. Barber et aI. (1996) and
Gibraltar Strait that altered main environmental conditions and lacilitated new coming floras. Also, alpine
Bohle et ai. (2002) proposed some working hypothesis. Selective pressure on ancestors reaching the
ranges started to act as pathways to mountain Iloras in an east-westward lashion. Later, mainland
Islands would have been in the context 01 lorest ecosystems and 01 competition with trees in the
areas sullered Irom the many glaciation cycles during the whole Pleistocene which promoied cycles 01
absence 01 large grazing animais (an absent selective pressure). Reproductive advantage 01 generalist
vegetation cover destruction and recovery Irom reluge areas. Moreover, Holocene history brought along
IndiViduais in. lorest habitats, tending to exclude newcomers with taxonomical and ecological niche
changes in landscape leatures dating back to 6.000 yrs. B.P with the Neolithic period.
afllnltles IS galned iI individuais have greater longevities. Also, elficient pollinization should be based on
Oceanic islands were not alfected by such complex phenomena during the Tertiary period, thus lew generalist insects. Thus, long-lived individuais, bearing inllorescences lor a long time would have
retaining part 01 the paleo-subtropical vegetation in such high latitudes. Further biogeographical .greaterreproductive success. Adding to this, necessary genetic diversity lor such a rapid and intense
relationships , mostly with the Mediterranean area, but also with others as lar as North America, were adaptatlvH radlatlon would be supplied by counter-selection 01 autogamic reproduction, lavoring also
conditioned by long-range dispersai events. Meanwhile, extensive speciation phenomena was taking 10ng-lIved Inflorescences. Hence, island woodiness habits stand lor physiognomical traits lavoring such
place, based both on the ancestors Irom the Tertiary elements and later newcomers Irom the evolutionary pressures.
continental areas. Finally, in spite 01 some uncertainty 01 human presence earlier, lirst human settlers
. Other non-Iorest groups such as Aeonium (Crassulaceae) and Echium (Boraginaceae) also exhibit
reached the islands in the early XV. Madeira Island was reached by the portuguese navigator
Island woodlness patterns. Consistency with the stated hypothesis is, in these cases, harder to sustain.
Gonçalves Zarco, in 1419 A.D . Ever since, both archeophytes and global neophytes are reaching
Madeira. Other important groups encompassing madeiran vegetation are paleo-mediterranean termophillous
sclerophylls (including many sprouters) such as spurges - Euphorbia (Sec\. Pachycladae and
Comparing with lossil records ranging Irom mainland Portugal to Siberia, it is assumed with some
Candelabra) , Olea and Maytenus. These are 01 circum-mediterranean-alrican-arabic alfinities (Rand
certainty, that lorest Ilora - trees , some climbers and a wealth 01 lorest lerns stand lor nowadays
Ilora) and reached the island in several cycles 01 colonization.
remnants 01 Artho-Tertiary Ilora. Examples 01 these in Madeira are: Laurus, Ocotea, Clethra, Apol/onias,
Myrica, Picconia, Dracaena, Persea, Syderoxylon, Prunus sect. Laurocerasus and I/ex. Cosmopolitan Neo-mediterranean elements (including many seeders) such as Teline Cistus Micromeria Sideritis
pteridophytes common in lorests are, e.g, 01 genera Oiplazium, Woodwardia, Oryopteris and Pteris. and Genista reached lhe island later. " ,

The most striking II0ral elements 01 the atlantic islands are woody endemics. Several genera,
subgenera or sections extant only in one archipelago or several are common in the Asteraceae (To/pis,
 The vegelation of Madeira Island (Portugal)
Capelo, J. & aI.
Starting from sea levei, the vegetation series of Madeira are as follows. Some references lo syntaxa
ln short, Madeira's flora can be coarsely grouped in five types: that integrate lhe correspondent geocomplex, viz. other relevant non-forest or forest seral stages, are
Paleo-endemic paleo-subtropical forest flora of tethysian origino also briefly discussed. Example-relevés can be seen in the correspondent descriptions of excursion
1.
Neo-endemic flora with island woodiness physiognomy. stops and consulting the portuguese text in QUERCETEA 6.
2.
3. Paleo-mediterranean xerophytic, sclerophyllous flora. ... .
Neo-mediterranean seeder flora of late-tertiary continental eurosSlbenan ongln.
4. 1. Climatophyllous series
5. Flora introduced by Man.
1.1. Mayteno umbellatae-Oleo maderensis sigmetum [series of madeiran oleaster tree]
d 2 stand for the forest vegetalion class PRUNO HIXAE-LAURETEA NOVOCANARIENSIS.
Types 1 an . I I d (Iype l' Pruno hixae-Lauretalia novocananensls; Iype 2. Inframediterranean, dry stage series on quite termophyllous rocky biotopes with patches of vertisols
endemic lo Madeira and Canary s an S · d II (Andryalo-Ericetalia of
Euphorbion mel/iterae) . Type 4 includes natural foresl hedges an . man es . I RHAMNO and cambisols, from the lower altitudes and sea cliffs of South face, up to 200 m above sea leveI. The
PRUNO-LAURETEA). Type 3 slands for lall-shrub xerophyllc vegelallon c ass . for climax is a low microlorest 01 madeiran oleaster [Olea maderensis] along with other paleo-
CRENULATAE-OLEETEA CERASIFORMIS. Type 5 encompasses several syntaxa slandlng mediterranean sclerophyll shrubs such as Maytenus umbel/ata, Chamaemeles coriacea, Dracaena
draco and Asparagus scoparius. First substitution stage is the scrub community 01 madeiran tree-
vegelalion of nilrogen-rich and dislurbed biotopes.
spurge - Euphorbia piscatoria together with Echium nervosum and Globularia salicina (Euphorbietum
piscatoriae) , lound mostly in moderate depth eroded soils of abandoned agricultural lields. ln leptosols
01 cliffs and rock outcrops , another community can be lound - The Artemisio argenteae-Genistetum
Vegetation belts and vegetation series of Madeira
tenerae, where Genista tenera (pi llow-shaped lorm). Carlina salicitolia, Micromeria varia subsp.

;tB;:~t~~ ::g~::t~:;~t';,:t,~~n~s:: ::~ ::~~~ ~:;t:~~:~;:~';~:~;::~~::~:::r:~:~~:: thymoides and Phagnalon bennetii are dominant. The perennial grassland stage is dominated by
Andropogonae grasses - Cenchro ciliaris-Hyparrhenietum sinaicae. Annual grassland stage is Galactito

~~~~~d:s~r~~:v~~Ihalenli~fln~ IOY:P: :!gfy~:W~:~:~~:t~~::~::~:';:::~'~I::i~':~~:~:~:; ~:;hU~td~

toinentosae-Brachypodietum distachyae. Rock communities in mosaic with the potential and seral
stages are Sedo nudi-Aeonietum glutinosi. Most of the territory 01 this series is nowadays occupied by
800 m.s.m. ege a 10 . M '1 tal (2004) - [see agricultural and urban setllements.
RIVAS-MARTíNEZ (2002) , modeled using geo-stalislics for Madeira by esquI a e . found in
18 and 19]. Three vegetalion series [successlonal elementary umts] are 1.2. Helichryso melaleuci- Sideroxylo marmulanae sigmetum [series of marmulano tree].
~:~~e~~~ean macrobioclimate under more or less severe summer droughl. Two e~~~nr:~:c:~!e~~~~ Microforest meso-xeric series of Sideroxylon marmulano, in inlramediterranean, subhumid bioclimate
forest vegelalion series plus a mountain vegelatlon complex (geoslgm~u~;o~e ~~ramediterranean lo on thin cambisols or leptosols. It occurs mostly in the north face from Oto 80 m.s.m, scatlered in the

:~:~~~d~I~~::and:~~g~~m~~i:~r~~~i~~ T~~~:;~~~ I::(~:~I: s~:ges range from Infralemperale lo south face, above the oleaster series (200-300 m.sm.). Potential communities are dominated by
Sideroxylon marmulano, Maytenus umbel/ata, Globularia salicina and in a few spots by Juniperus
Supralemperale and Irom humid lo ullrahyperhumid. . turbina ta subsp. canariensis. The typical substitution stage is a community 01 Helichrysum melaleucum

N:~:~~:~sscsa~~ ~~~;d:I~~~~~:nsi:~lo~:!a:f ~~~::liS~~::~~:i~~~i~~~~::e ~~~~~:~~~;~ V;~~I~~~;~

and Globularia salicina. On rocky walls, in the scope of the series geocomplex, a Aeonium
glandulosum-dominated community is found (Sinapidendro gymnocalicis-Sedetum brissemoretil).
(bananas, ho~icullure, greenhouse crops) and afforestalions wilh exollc trees. . 1.3. Semeie androgynae-Apollonio barbujanae sigmetum [mediterranean laurel / barbusano-tree
. . . b Ih faces of lhe island, so11 Iypes and lorest series]
[Climatophyllous vegelalion series, Ihe:r ~IIIIUdl~~1 ~:b~:s2 ~~ p~4 summarizes lhe main seral stages
bioclimate slages are summanzed ln lab e , p. ~e-A ol/onio barbujanae sigmetum are considered - The climax is a thermophillous, inlra- and thermomediterranean Lauraceae forest subhumid to humid
of such series. Two syn-facles of Semeie androgyn d .~ Globulario salicinae-Ericetum maderincolae. forest community 01 cambisols, in both faces 01 the island. These micro-Iorests are dominated by
wilh Myrto commums-Hypencetum canan~nsls t~~estW~bOSque) , high-scrub (matagal) , perennial tal/- Apol/onias barbujana, Laurus novocanariensis, Myrica taya and I/ex canariensis. ln understory layers,
Stages consldered are , ln respectlve or er. b ( lo baixo) and annual grassland (arrelvado climbers are dominanl - Semeie androgyna, Smilax canariensis, Smilax pendulina, Hedera maderensis
torbslgrass community (arrelvado Vivaz) , low scru ma subsp. maderensis, Convolvulus massonii and Rubia agostinhoi. Thermophyllous, sometimes spiny
annual)]. shrubs such as Asparagus umbellatus subsp. lowei, Visnea mocanera and Maytenus umbel/ata are also
 The vegetation of Madeira Island (Portugal)
Capelo, J, & aI.

frequent. When compared to temperate communities , where a wealth of ferns, shade-Ioving forbs and community (topographical climax), These heath scrubs can sometimes assume the character of a
secondary forest due to the heaths tree-habit. A second substitution stage is of brooms (Genista tenera,
grasses are found, the Semele-Apol/onietum is poorer, as expected from a mediterranean community
Teime maderensls): Bystropogono punctatae-Telinetum maderensis, Complete destruction of stink
under some summer drought.
laurel forest and ItS woody seral stages gives rise to a grassland - Leontodo longl'r t' -
Two distinct successional facies of the barbusano series can be recognized by their distinct seral O 'th d' os ns
rm opo letum perpusili made up of exotics introduced long ago as weedy archeophytes, Grazing
stages. Thus, in infra-thermomediterranean stages sub-humid stages of South face (300-600 m,s,m,), promotes a perennlal grassland : VIOlo nvmlanae-Agrostietum castel/anae,
an high-scrub community of Hypericum canariense (Myrto communis-Hypericetum canariensis) is the
1,5, Polysticho falcinelli-Erico arboreae sigmetum [high altitude tree-heath series]
forest hedge, sometimes becoming dominant. ln higher altitudes (600-800 m,s,m) and in the North face,
the high shrub stage of the series is an heath community of Erica platycodon subsp, maderincola, Erica Meso-supra temperate, hiperhumid and ultrahyperhumid series of tree-heath forests (Erica arborea) , It
arborea, Myrica fava and xero-thermphyllous shrubs such as Globularia salicina, Teucrium betonicum stretches approxlmately from 1400 to 1650 m,s,m, The forest community is probably a para-climax of a
and Echium nervosum (Globulario salicinae-Ericetum arboreae) , Advanced degradation stages of the forme r Enca arborea -Juniperus cedrus subsp, maderensis (Madei ra juniper) community with scarce
forest can also include the Euphorbietum piscatoriae in both successional facies , understory, namely of nemoral plants and Lauraceae, Madeira juniper was , in the past, extensivel y
logged for tlmber and charcoal and nowadays is a very rare tree. Understory is dominated by a
1.4. Clethro arboreae-Ocoteo toetentis sigmetum [stink-Iaurel temperate forest series]
xeromorphlc lern - Polystlchum falcmellum. Clearings bear the Teucrium francoi and oregano
Infra to mesotemperate, humid to hiperhumid forest series on volcanic andosols of stink-Iaurel (Ocotea (Onganum Vlrens) communltles - Teucno francoi-Origanetum virentis, Violo-Agrostietum castel/anae
foetens) , The climax community is forest that covers very large stretches of both faces of the island grasslands are also common under grazing regimes, The natural hedge is a species-poor communi ty of
(800-1450 m,s,m. south face; 300-1400 North face), Multi-stratified forest up to 30 m high (mesoforest) shrubby Enca platycodon subsp. maderincola, A second hedge or rocky permanent community can be
of hiper-oceanic character with tree-stratum dominated by Ocotea foetens, Laurus novocanariensis, and also recognlzed ln the scope of the associated geocomplex (Argyranthemum montanae-Ericetum
Clethra arborea, Also frequent are: Picconia excelsa, Heberdenia excelsa, Persea indica, Prunus maderensis)
lusitanica subsp, hixa and I/ex perado (Clethro arboreae-Ocoteetum foe ten tis) , The understory has a
1,6. Armerio maderensis-Parafestuco albidae microgeosigmetum [high-altitude rock vegetation
very high diversity of ferns , epiphytic plants, mosses, herbs and forbs. Examples are: Diplazium
complex]
cauda tum, Pteris incompleta, Asplenium onopteris, Dryopteris maderensis, Woodwardia radicans,
Culcita macrocarpa, Festuca donax, Carex lowei and Sibthorpia peregrina, Climbers like Rosa mandonii Ove r 1650 m above sealevel , under supratemperate ultrahyperhumid bioclimate stage, a mosaic of
and Rubus grandifo/ium are also frequent. Natural forest clearings bear tall-forb communities of three communllles occuples most of the higher peaks, according to micro-habitats: i) Armerio
Geranium palma tum, Perical/is aurita and Ranunculus cortusifolius subsp, major (Perical/ido auritae- maderensls-Parafestucetum albidae, in earthy crevices or natural platforms on vertical cliffs; ii)
Geranietum palmatae), Smapldendro frutlscescentls-Aeometum glandulosi in vertical rock walls and iii) Thymetum micantis in
Many microhabitats are found within the forest, namely: epiphytic communities (Davalio canariensis- flat leptosols of pyroclast material.
Polypodietum macaronesic/); communities of shady earthy walls (Sel/aginelo denticulatae-
Cystopteridetum viridulae, Hymenophyl/etum thumbrigensi-maderensis); shady wet rock walls
Excursion program and geobotanical notes on stops and walks
(Aichrysetum divaricati-vil/osae) . The caulirosetled phanerophyte (Euphorbion mel/iferae) communities
associate with catastrophic gravitational landslides within the forest and rocky ground of small streams Table 1- Excursion program to Madeira
(Isoplexido sceptri-Euphorbietum mel/iferae: - dominant Isoplexis ~ceptrum, Sonchus fruticosus,
Muschia wol/astonii, Melanose/inum decipiens) , ln the upper part of permanent streams within the Day Morning (8H30-13H) Afternoon (14H-19H)
forest, contacts with hygrophilous communities are with madeiran elder community (Rhamno glandulosi- 18 Arrival on Funchal ai rport Presentation of Madeira vegetation
Sambucetum lanceolatus/), Mid-strech streams found within the Clethro-Ocotetum are ebony riverine
forests of Persea indica (Oiplazio caudatae-Perseetum indicae), ln lower streches of streams , where silt Presentation of excursion program
substratum is greater, a willow community is found : Scrophulario hirtae-Salicetum canariensis, Hedges 19 Visit to Madei ra Botanical Garden Calheta,
and mantles of riverine gallery forests are olten bramble thickets: Rubio agostinhoi-Rubetum bol/ei,
Ribeira Brava (by bus) Prazeres (two-hours moderate-easy walk)
The natural hedge community and fisrt substitution stage of the stink-Iaurel forest is an tree-heath
community and madeiran blueberry (Vaccinio padifo/i-Ericetum maderincolae). ln some thin soil steep Ponta do Sol (by bus)
cliffs, exposed to permanent fog s and wetling winds the later can be regarded also as a permanent
 The vegetation of Madeira Island (Portugal)
Capelo, J. & al.
July, the 19 th -1. Ribeira Brava stop, 2. Ponta do Sol stop, 3. Calheta stop, 4. Prazeres-Paúl do
20 Montado dos Pessegueiros (six-hou r forest walk, quite hard)
Mar walk (Iow altitude mediterranean vegetation)

21 Levada do Folhadal (5-hour walk, easy-moderate) A transect following along the steep basalt cliffs and valleys facing the sea is made. Natural potential
vegetation of the low altitude area is of the madeiran oleaster woodlands - Mayteno umbellatae-
São Vicente/Seixal (one-hour walk, easy)
Oleetum maderensis [see table 2, p. 74J. Due to human activities and rural and urban settlement, only
22 Pico Areeiro - Pico Ruivo (5-hour walk, moderate, beware of vertigo) Ponta de São small patches of well preserved oleaster forests can be found. Therefore, its main seral stage - the tree-
Lourenço (2 hour walk , easy) spurge community of Euphorbia piscatoria - Euphorbietum piscatoriae is dominant [se e table 1, p. 73J.
Canary-madeiran tall -grass community of Hyparrhenia sinaica - Cenchro ciliaris-Hyparrhenietum
23 Caldeirão Verde (5-hour walk, easy, beware of vertigo) sinaicae occupies the clearings in eroded soil [see table 27, p. 112J. Rosetted crassulaceae
communities with Aeonium glutinosum and Sinapidendron angustifolium can be found in rock outcrops
- Sedo nudi-Aeonietum glutinosi [see table 65, p. 165J.
24 Departure to Lisbon
Data concerning the stop at Ribeira Brava are as follows.

Bioclimate: upper inframediterranean pluviserotinal upper dry euhyperoceanic

Substratum: basalt leptosols and rock outcrops.

3tOCOO 32(){)OO 330000 340000 ~
290000 300000
~L ~ Vegetation series: Mayteno umbellatae-Oleo maderensis sigmetum
~ i

Mosaic of communities (geocomplex):

1) Mayteno umbellatae-Oleetum maderensis (Rhamno crenulatae-Oleetea cerasiformis). [see table

§j '" 2, p. 74J.
~ ]
"" 2) Euphorbietum piscatoriae (Rhamno crenulatae-Oleetea cerasiformis) [see table 1, p. 73J.
3) Cenchro ciliaris-Hyparrhenietum sinaicae (Lygeo sparti-Stipetea tenacissimae) [see table 27, p.
112J
4) Sedo nudi-Aeonietum glutinosi (Greenovio-Aeonietea) [see table 65, p. 165J.
§J
~i
i 5) Galactito tomentosae-Brachypodietum distachyeae (Stellarietea mediae) [see table 49, p. 144].
6) Sonchetum pinnati (extra-forest caulirosetted community) [see relevé in p. 91J.

1) Ribeira Brava, 19/4/2004, 40 m2, basalt, JC, RJ , MS, JCC, SM: 3 Olea maderensis, 1 Maytenus
umbellata, 1 Globularia salicina, + Asparagus scoparius, + Sonchus pinnatus, + Echium
~~~ I
~
+ + + + + 8
nervosum, 1 Euphorbia pisca to ria, + Carlina salicifolia, + Aeonium glutinosum, 2 Opuntia tuna, +
- o
5000 5 000 10000 m
- -= Bituminaria bituminosa.
BO'ooO 340000
290000 300000

2) Ribeira Brava, 19/4/2004, 40 m2, basalt, JC, RJ, MS, JCC, SM: 5 Euphorbia piscatoria, 3
O- IDO 600 - 800
1. Ribeira Brava 6. Levada do Folhadal walk
m 1 4oo-1600
Globularia salicina, 2 Echium nervosum, + Crambe fruticosa , 1 Olea maderensis, + Genista
2. Ponta do Sol 7. São Vicente-Seixal walk 100 - 200 800 - 1000
3. Calheta 8. Pico do Areeiro-Pico Ruivo walk 200 - 400 _ 1 000- 1200 m 16oo- 1800 tenera, + Sideritis candicans, + Plantago arborescens subsp. maderensis, 1 Aeonium glutinosum,
4. Prazeres· Paul do Mar walk 9. Ponta de S, Lourenço walk
400 - 600 m 1200 ·1 400 m ,1850
5. Montado dos Pessegueiros walk 10. Caldeirão Verde wal k 1 Opuntia tuna, + Sinapidendron na angustifolium, + Hyparrhenia sinaica, 1 Sonchus ustulatus
susbp. ustulatus.

Figure 1 - Stops and geobotanical walks in Madeira Island IAVS 2005 Excursion 3) Ribeira Brava, 19/4/2004, 4 m2, basalt, JC, RJ , MS, JCC, SM: 3 Hyparrhenia sinaica, 2 Cenchrus
ciliaris, + Phagnalon benetii, 1 Brachypodium distachyon, + Micromeria varia subsp. thymoides, +
Convolvulus althaeoides.
Capelo, J. & aI. The vegetation 01 Madeira Island (Portugal)
4) Ribeira Brava, 19/4/2004, 8 m2, basalt, JC, RJ, MS, JCC, SM: 3 Aeonium glutinosum, 2 9) Ribeira Brava, 100 m2, JC, MS, JCC, SM: 4 Hypericum canariense, 1 Myrtus communis, 1
Sinapidendron angustifolium, 2 Tolpis suculenta, + Davallia canariensis, + Genista tenera, + G/obu/aria salicina, 1 Maytenus umbellata, (x) Laurus novcanariensis, 1 Optuntia tuna, 2
Carlina salicifolia. Aeomum g/andu/osum, + Hyparrhenia sinaica, 1 Asparagus asparagoides, + Davallia canariensis.
5) Ribeira Brava, 19/4/2004, 2 m2, basalt, JC, RJ , MS, JCC, SM: 3 Brachypodium distachyon, +
Galactites tomentosa, 1 Leonfodon taraxacoides subsp. longirostris, 1 Petrorhagia nanteulii,
Vulpia myurus, + Trifolium stellatum, + Holcus lanatus, 1 Silene gallica.

During the afternoon we lollow a descent transect lacing south, Irom 500 m.s.m. approximately to sea
levei, crossing the potential area 01 barbusano tree lorests (SemeIe androgynae-Apollonietum
barbujanae) , and then meeting again, around 300 m.s.m. the oleaste r series vegetation mosaico

ln the barbusano tree series, the hedge/tall-shrub communities 01 Hypericum canariense - Myrto
communis-Hypericetum canariense - can also be see. Both series share, in the locality, the tree-spurge
community as degradation stage 01 lorest ecosystems.

Correspondent bioclimate stages are thermomediterranean low sub-humid Irom 500 to 300 m.s.m.
and inlramediterranean dry Irom under 300 m.s.m.

Data concerning Prazeres-Paúl do Mar walk, are as lollows.

Bioclimate: lower thermomediterranean lower sub-humid pluviserotinal euhyperoceanic

Substratum: basalt, pyroclast and ash cambisols with "mull"and eutric leptosols.

Série de vegetação: SemeIe androgynae-Apollonio barbujanae sigmetum

Mosaico 01 communities (geocomplex):

7) SemeIe androgynae-Apollonietum barbujanae (Pruno hixae-Lauretea novocanariensis) [See table

10, p. 88].
8) Globulario salicinae-Ericetum arboreae (Pruno-Lauretea) [See table 13, p. 92].
9) Myrto communis-Hypericetum canariense [See table 3, p. 75].

7) S. Vicente, 19/4/2004, 200 m2, basalt, JC, RJ , MS, JCC, SM: 4 Apollonias barbujana, 2 Myrica
faya, 2 Laurus novocanariensis, 1 SemeIe androgyna, 1 Persea indica, 1 Hedera maderensis
subsp. maderensis, 1 Smilax pendulina, + Smilax canariensis, + Arum italicum subsp.
canariensis, 1 lIex canariensis, + Convolvulus massonii, + Asparagus umbellatus subsp. lowei, +
Rubia agostinhoi, 1 Festuca donax, 1 Maytenus umbellata, + G/obularia salicina, + Erica
platycodon subsp. maderincola, + Teucrium betonicum, + Carlina salicifo/ia, + Davalia
cannariensis, + Pericallis aurila, + Pteridium aquilinum, + Hypericum gandulosum, + Teline
maderensis.

8) S. Vicente, 19/4/2004, 40 m2, basalt, JC, RJ , MS, JCC, SM: 4 Erica platycodon subsp.
maderincola, 2 Globularia salicina, 1 Helichrysum melaleucum, + Phyllis nobla, + Erica arborea, 1
Echium nervosusm, 1 Myrtus communis, + Plantago arborescens subsp. maderensis, + P/antago
leiopetala, 1 Rubus ulmufolius. Figure 2 - Sktech 01 transect Irom Prazeres do Paúl do Mar. (numbers are those 01 above relevés).
Capelo, J. & aI. The vegetation 01 Madeira Island (Portugal)

CD
July, the 20th - 5. Montado dos Pessegueiros walk (altitude tree-heath and dense forest
vegetation)

Starting Irom ca. 1500 m.s.m. in the central plateau 01 Madeira (Paúl da Serra), we lollow a descent
through dense tree-heath (Polysticho talcinelli-Ericetum arboreae) , then a large stretch inside the stink
laurel lorest (Clethro arboreae-Ocoteetum toetentis) and finally through barbusano tree lorest (Semeie
androgynae-Apollonietum barbujanae). The descent ranges Irom 1500 m.s.m. to ca. 40 m.s.m . and
crosses ali the bioclimates in their wetter ombrotype variants lound in the North lace 01 the island
(supratemperate to inlramediterranean).

Main leatures 01 vegetation along Montado dos Pessegueiros walk are as lollows.

The altitude plateau has been, since early human colonization, completely cleared Irom lorest cover
and grazed by cattle. Low biomass annual grass communities 01 Vulpia myurus (Leontodo longirostris-
Ornithopetum perpusilii, Helianthemetea guita ti) [see table 30, p.116] are dominant. Nevertheless, in
deeper soils with grealer water availability during summer and under grazing by cows, a perennial grass
community appears (Violo rivinianae-Agrostietum castellanae, Stipo giganteae-Agrostietea) [see table
25, p. 109]. Permanently wet spots have Cariei cedercreutzi-Juncetum etfusi (Molinio-Arrhnatheretea)
[see table 19. p. 101]. Large slretches 01 Ilal vernal pools have annual vegetation associated to
presence 01 water during late winter and spring (lIIecebro verticillati-Lotetum parviflori, Isoeto-
Nanojuncetea) [see table 75, p. 180].

The lirst lew kilornelers go through adense heath-high-scrub 01 secondary Polysticho talcinelli-
Ericetum arboreae (Pruno-Lauretea) [see table 11, p. 89]. This micro-Iorest is the meso-supratemperate
altitude climax 01 Madeira. Its nalural hedge is a species-Iow communily 01 Erica platycodon subsp.
maderincola. Around 1400 m.s.rn., one enters lhe Clethro arboreae-Ocoteetum toetentis (Pruno-
Lauretea) [see table 6, p. 80], which is the climax 01 temperate macrobioclimate under 1400 m.s.m. in
Madeira. ln alternate lashion and due to quite ancient use lor charcoal 01 the lorest we go Ihraugh
several patches 01 Vaccinio padifoli-Ericetum maderincolae (Pruno-Lauretea) [see table 7, p. 82], which
is the laurel lorest lirst substitution stage. ln some crests this high heath scrub appears to be a
permanent community. The later healh-scrub is quite distincl Iram Polysticho-Ericetum arboreae due to
a great wealth 01 nemoral elements and lauraceae in the lormer, absent in the altitude heath-Iorest.

Alter a lew time 01 descent, circa 300 m.s.m., the medilerranean, liana-rich lorest 01 Apollonias
barbujana (Semeie androgynae-Apollonietum barbusanae, Pruno-Lauretea) [se e table 10, p. 88]
appears. On the clifl above the sea small woodland 01 Sideroxylon marmulano can be also seen
(Helichryso melaleuci-Syderoxyletum marmulanae, Rhamno-Oleetea) [see table 4, p. 76].
Figure 3 - Skelch 01 Montado dos Pessegueiros transect: 1 Polysticho talcinelli-Ericetum arboreae; 2
community 01 Erica platycodon subsp. maderincola; 3 Clethro arboreae-Ocoteetum toetentis; 4 Semeie
androgynae-Apollonietum barbujanae; 5 communily 01 Globularia salicina, Echium nervos um and
Helichrysum melaleucum
Capelo, J. & aI. The vegetation 01 Madeira Island (Portugal)

July, the 21th - 6. Levada do Folhadal walk, 7. São Vicente-Seixal walk (dense diverse forest 12) Aichrysetum divaricato-vi/losi
and rock vegetation)
1) Levada do Folhadal, 19/4/2004, 800 m2, basalt, JC, RJ , MS, JCC.: 4 Ocotea foetens, 3 Laurus
6. Levada do Folhadal (stink-Iaurel forest)
novocanariensis, 1 Sibthorpia peregrina, 2 Clethra arborea + Diplazium caudatum, 1 Pteris
Walk lollowing a small water channel (levada) that runs through a pristine lorest 01 Laurus/Ocotea incompleta, 1 Festuca donax, 1 Rubia agostinhoi, 1 Euphorbia longifolia, 1 Myrica faya, 1 Persea
(Clethro-Ocoteetum) [see table 6, p. 80]. Although the lorest itsell has a high tree and understory indica, + Sonchus fruticosus, 1 /lex pera do, + Woodwardia radicans, + Argyranthemum
diversity, many lorest-associated habitats can also be seen. Namely the lorest clearing lorbs community pinnatifidum, + Carex peregrina, + Cedrone/la canariensis, 1 Blechnum spicant, + Erica arborea,
01 Perica/lido auritae-Geranietum palmatae, (Trifolio- Geranietea sanguinel) [see table 33, p. 122]; the 1 Heberdenia excelsa, 1 Picconia excelsa, + Viola odorata, + Rhamnus glandulosa, + Ruscus
laurel lorest high scrub hedge 01 Vaccinio-Ericetum maderincolae [see table 7, p. 82]; the second streptophy/lus, + Phy/lis nobla, + Erica platycodon subsp. maderincola, + Hedera maderensis
substitution stage, the broom hedge 01 Bystropogono puncati-Telinetum maderensis [see table 14, p. subsp. maderensis, + Rubus grandifolius, + Rosa mandonii, + Cirsium latifolium, + Carex lowei ,
94]; the caul irosetled communities which lollow any larger clearings in the crown layer 01 laurel lorest, + Se/laginela denticulata, + Rubus bol/ei, + Dryopteris aitoniana, 1 Prunus lusitanica subsp. hixa,
namely the water channel we walk through (Isoplexido sceptri-Euphorbietum me/liferae) [see table 12, + Dryopteris maderensis, + Aichryson diva rica tum, + Dava/lia canariensis, + Perical/is aurila, +
p. 90]. Small permanent streams running through the laurel lorest exhibit the Rhamno glandulosi- Polystichum setiferum, + Brachyodium sylvaticum, + Duchesnea indica.
Sambucetum lanceolati hygrophilous community [see table 8, p. 84]. ln the channel earthy walls, a lern
2) Levada do Folhadal, 19/4/2004, 40 m2, basalt, JC, RJ , MS, JCC: 5 Geranium palmatum, 2
community - Se/lagine/lo denticulatae-Cystopteridetum viridulae is lound [see table 64, p. 63]. Orier
Perica/lis aurila, 1 Ranunculus corfusifolius subsp . major, 1 Brachypodium sylvaticum, +
spots on earth walls exhibit a community 01 Adiantum reniforme (Adiante tum reniformis) [see table 57,
Origanum virens, + Dactylorhiza foliosa, + Clinopodium vulgare, + Rumex maderensis, + Carex
p. 156]. Small vertical cascades with permanent water lilms may have a Woodwardia radicans
divulsa, + Calamintha sylvatica, + Aichryson divaricatum, + Phyl/is nobla, + Argyranthemum
community [see relevé in p. 164] or, depending on substrata and water regime the extensive decumbent
pinnatifidum, + Viola odorata, + Rosa mandonii, 1 Erysimum bicolor, + Sibthorpia peregrina, +
grass community 01 Deschampsia argentea (Deschampsietum argenteae, Phragmileto-Magnocaricetea)
Selagine/la denticulata, + Succisa pratensis, + Tolpis macrorhiza.
[see table 73, p. 177]. Epiphytic communities may be 01 Davallio canariensis-Polypodietum
macaronesici [see table 62, p. 161] or Hymenophy/letum thumbrigensi-maderensis (exhibiting 3) Levada do Folhadal, 19/4/2004, 40 m2, basalt, JC, RJ , MS, JCC: 4 Erica platycodon subsp.
sometimes Elaphoglossum semicylindricum) [see table 63, p. 162]. The Aichrysetum divaricati-vi/losae, maderincola, 1 Erica arborea, 1 Laurus novocanariensis, 2 Myrica faya, 1 Vaccinium padifolium,
dominated by Aeonium glandulosum and annual crassulaceae can also appear in tree trunks and rock + Sibthorpia peregrina, + Argyranthemum pinnatifidum, + I/ex perado, 1 Rubus bol/ei, + Athyrium
walls [see table 69, p. 169]. filix-foemina, Sel/aginel/a denticulata, + Gennaria diphyl/a, + Cytisus scoparius, + Pteridium
aquilinum.
Bioclimate: lower mesotemperate hyperhumid pluviserotinal euhyperoceanic
4) Levada do Folhadal, 19/4/2004, 20 m2, basalt, JC, RJ , MS, JCC: 5 Teline maderensis, 2
Substratum: deep andosols Iram basalt and pyroclasts, mull-rich.
Argyranthemum pinnatitidum, 1 Phyllis nobla, 2 Genista tenera, 1 Bystropogon punctatus, 1
Vegetation series: Clethro arboreae-Ocoteo foetentis sigmetum Cytisus scoparius, 1 Erica platycodon subsp. maderincola, + Festuca donax, + Rubus
grandifolius, 1 Myrica taya, + Perical/is aurila.
Mosaic 01 communities (geocomplex):
5) Levada do Folhadal, 19/4/2004, 40 m2, basalt, JC, RJ, MS, JCC: 2 Euphorbia longitolia, 1
1) Clethro arboreae-Ocoteetum foetentis
Isoplexis sceptrum, 3 Sonchus fruticosus, 1 Erysimum bicolor, 2 Melanoselinum decipiens, 2
2) Perica/lido aurilae-Geranietum palmatae
Muschia wol/astonii, 1 Clethra arborea, + Teline maderensis, + Erica platycodon subsp.
3) Vaccinio padifoli-Ericetum maderincolae
maderincola, + Rubus bol/ei, + Rubia agostinhoi, 1 Phyl/is nobla, + Succisa pratensis, 1
4) Bystropogoni punctati-Telinetum maderensis
Geranium palma tum, + Perical/is aurita, + Bystropogon punctatus, + Ageratina adenophora, +
5) Isoplexido sceptri-Euphorbietum me/liferae
Argyranthemum pinnatifidum, + Erigeron karvinskianus, + Arachniodes webbianum, +
6) Rhamno glandulosi-Sambucetum lanceolati
Stenogramma pozoi.
7) Selagine/lo denticulatae-Cystopteridetum viridulae
8) Adiantetum reniformis 6) Levada do Folhadal, 19/4/2004, 40 m2, basalt, JC, RJ , MS, JCC: 3 Sambucus lanceolata, 1
9) Community 01 Woodwardia radicans Rhamnus glandulosa, + Euphorbia longifolia, 2 Diplazium cauda tum, 2 Festuca donax, +
10) Davalio canariensis-Polypodietum macaronesici Sibthorpia peregrina, + Pteris incompleta, 1 Rosa mandonii, + Cystopteris diaphana, + Athyrium
11) Deschampsietum argenteae
 The vegetation 01 Madeira Island (Portugal)
Capelo, J. & aI.

filix-foemina, 2 Oenanthe divaricata, 1 Woodwardia radicans, 1 Oeschampsia argentea, +

Ranunculus corfusifolius subsp. major, 1 Bryophyta.

7) Levada do Folhadal, 19/4/2004, 1 m2, basalt, JC, RJ, MS, JCC: 3 Selaginella den ticula ta, 3
Cystopteris viridula, 2 Anogramma leptophylla, + Stenogramma pozoi, 2 Anthocerus sp. , +
Asplenium monanthes.

8) Levada do Folhadal, 19/4/2004, 1 m2, basalt, JC, RJ, MS, JCC: 3 Adiantum reniforme varo
reniforme, 1 Asplenium monanthes, + Asplenium trichomanes, 1 Oavallia canariensis, + Aeonium
glandulosum.

9) Levada do Folhadal, 19/4/2004, 4 m2, basalt, JC, RJ, MS, JCC: 5 Woodwardia radicans, +
Stenogramma pozoi, + Carex lowei.

10) Levada do Folhadal, 19/4/2004, 0,5 m2, on Ocotea foetens, JC, RJ, MS, JCC: 2 Oavallia
canariensis, 2 Polypodium macaronesicum, + Aichryson divaricatum, 2 Usnea sp.

11) Levada do Folhadal, 19/4/2004, 10m2, waterfall on basalt, (Capelo et ai., 1999): 5 Oeschampsia
argentea, 2 Oenanthe diva rica ta, 2 Phillonotis fontana, 1 Stellaria uliginosa, 2 Sellaginella
denticulata, + Woodwardia radicans, + Hypericum grandifolium, + Apium nodiflorum, + Athyrium
filix-foemina, + Conocephalum conicum, 1 Asterella africana, 1 Pellia epiphylla, + Anthoceros sp. ,
+ Phaeoceros sp., Marchantia sp., + Porella canariensis, + Jubula hutchinsiae, + Fontinalis
antipiretica.

12) Levada do Folhadal, 19/4/2004, 4 m2, basalt, JC, RJ , MS, JCC: 3 Aeonium glandulosum, 3
Aichryson villosum, 1 Saxifraga maderensis, + Aichryson diva rica tum, + Galium productum, +
Asplenium trichomanes subsp. quadrivalens, + Bupleurum salicifolium.

Figure 4 - Levada do Folhadal (numbers in ligure are those 01above relevés).

111
110
• Capelo, J. & aI. The vegetation 01 Madeira Is/and (Portugal)

7. São Vicente rock vegetation

Woody vegetation is represented by the Sapotaceae - Sideroxy/on marmu/ano - (He/ichryso-

Sideroxy/etum marmu/anae) [see table 4, p. 76J microforest patches and in a higher position on the ciiff
by Seme/e-Apollonietum forests) [See table 10, p. 88]. Moreover, the most striking vegetation is from
rock walls. These are as follows. Extra-forest caulirosetted community of Sonchus pinnatus (Sonchetum
pinnati, Euphorbion mel/iferae); community of Aeonium g/andu/osum from low altitude in the North face
- Sinapidendro gymnoca/icis-Sedetum brissemoretii [see table 66, p. 166J; pyrociast walls community of
Muschia aurea (Campanu/aceae) (Muschiatum aureae, Grenovio-Aeonietea) [see table 67, p. 167J.
Oeschampsietum argenteae and Euc/adio-Adientetum capillis-veneris on shady damp walls are a/so
present.

[Note: additiona/ information on these stops can be found in pages 34 to 37 of CAPELO et alo(2004) .
See a/so pictures there.J

Figure 5 - S. Vicente rock vegetation: 1 Semeie and~ogynae-Apollonietum barbusanae; 2 Sonchetu~

pinnati; 3 G/obu/ario sa/icinae-Ericetum arboreae; 4 Smapldendro gymnocalicls-Sedetum bnssemoretll,
5 He/ichryso me/a/euci-Sideroxy/etum marmu/anae

113
112
Capelo, J. & aI.
The vegetation 01 Madeira Island (Portugal)

July, the 22th - 8. Pico do Areeiro-Pico Ruivo walk, 9 - Ponta de S. Lourenço walk (mountain
vegetation and sea-cliff vegetation)

8. Pico do Areeiro-Pico Ruivo

Mountain walk along rock walls with spectacular sights. Starting Irom Pico do Areeiro (1818 m.s.m.)
we lollow a path to reach Pico Ruivo (1861 m.s.m.). Near Pico do Areeiro climatophyllous woody
vegetation is only represented by a quite diverse shrub community with brooms (Tefine maderensis,
Genista tenera). heaths (Erica maderensis), pride-ol-Madeira (Echium candicans) and the very
abundant neophyte Cytisus scoparius - Argyranthemo montanae-Ericetum maderensis [see table 15, p.
95J. Rock crevices with some soil, exhibit a perennial grass community 01 Oeschampsia maderensis,
Parafestuca albida and Armeria maderensis (Armerio maderensis-Parafestucetm albidae, Stipo-
Agrostietea) [see table 26, p. 11 OJ. Deeper soil small plateaus in steep vertical clifls also have the same
community in a version dominated by grasses that was , until recent times, grazed by goats. Arare
menaced bird endemic 01 Madeira (Pterodroma madeira) nests exclusively on some 01 these plateaus.
Rock walls are occupied by Sinapidendro frutescentis-Aeonietum glandulosi (Grenovio-Aeonietea) [see
table 69, p. 169J crassulaceae community. Flat incoherent pyroclast plattorms have mat-forming
communities 01 Thymus micans (c.f. Thymus caespititius) - Thymetum micantis [see table 24, p. 108J.
Approaching Pico Ruivo, a tree heath woodland of Erica arborea (Polysticho falcinefli-Ericetum
arboreae, Pruno-Lauretea) [see table 11 , p. 89J with very large trees, is lound. Tall-Iorb communities
lound there are in shady semi-nitrogen-rich clearings and are 01 Vicio capreolatae-Odontitetum
hoflianae (Geranio purpureae-Cardaminetea hirsutae) [see table 31 , p. 118J. ln non-nitrophyllous forest
clearings occurs the Teucrio francoi-Origanetum virentis (Trifolio-Geranietea) [table 34, p. 123J. A linal
stretch away Irom Pico Ruivo to Achada do Teixeira is along grasslands 01 Violo-Agrostietum
casteflanae.

~i~ure 6 - Sketch 01 mountain vegetation: 1 Armerio maderensis-Parafestucetm albidae;

rgyranthemo montanae-Encetum maderensis; 3 Sinapidendro frutescentis-Aeonietum glandulosi
Capelo, J. & aI.
The vegetation 01 Madeira Island (Portugal)

9. Ponta de S. Lourenço

The Ponta de S. Lourenço lorms a long '~ail " at the extreme eastern end 01 the Island, terminating in
two small islets. It is an area 01 low altitudes with rounded slopes under the strong inlluence 01
permanent desiccating winds and salt spray. It is almost void 01 tall woody vegetation due to early
agricultural use (wheat, Irom the XV th century on) and probable low dry to semiarid ombroclimate
(thermotype is inlramediterranean). Although being made 01 the oldest volcanic materiais 01 Madeira,
the Pleistocene (or perhaps even late Tertiary) crisis lead to some deposition 01 sand by winds, during
regression 01 sea-Ievel episodes 01 colder glacial and stadial periods. Therelore an area 01 consolidated
paleo-dunes (grimaldian) can be seen at Piedade overtopping volcanites. Plant lossil structures 01
woody plants 01 uncertain origin can also be seen there -rhyzoconcretions-. The sand originated
probably Irom a sea-bottom made 01 rem ains 01 ancient coral reels, being thereafter 01 calcareous
nature. Salt spray intensity, topography and irregular nitrogen deposition (by birds and human activity)
are the main lactors determining vegetation pattern.

Dune vegetation is represented by Euphorbio paraliae-Lotetum glauci, a species-poor branch 01

Ammophilletea vegetation class in Canaries and Madeira (Euphorbio paraliae-Lotion glaucl) [See table
70, p. 173). Sea-cliff rocky habitats heavy sprayed by salt wind are occupied by an original madeiran
version 01 Crithmo-Staticetea vegetation - the Chrithmo maritimae-Helichrysetum obconicae [see table
71, p. 174J. Where inlluence 01 nitrogen is higher - platforms where sea birds dwell- a woody nitrogen-
salt adapted community appears - Calendulo maderenis-Suaedetum verae - dominated by endemics,
e.g. Calundula maderensis and Argyranthemum pinnatifidum subsp. succulentum [see table 37, p. 127).
Depending on combination 01 proportion 01 nitrogen, sodium chloride, biotype and phenology, several
ruderal communities appear. Nitrogen, salt and semi-arid biotopes are occupied by the succulent
Senecio incrassatae-Mesembryanthemetum cristalini [see table 48, 142J. Spring communities 01
moderate nitrogen soil content are dominated by annuals 01 quite large biomass - Galactito 0"0 .~ .

tomentosae-Brachypodietum distachyae [see table 49, p. 144]. Poorer and thinner soils have the grass-
dominated Lino stricti-Stipetum capensis [see table 52, p. 149J. Spots where locally organic debris
accumulate, have the highly nitrophyllous thistle community Scolymo maculatae-Cynarietum
ferocissimae. [ see table 35, p. 125J. Figure 7 - Sketch 01 Ponta de S. Lourenço: 1 Chrithmo maritimae-Helichrysetum obconicae; 2
Calendulo maderenis-Suaedetum verae; 3 Senecio incrassatae-Mesembryanthemetum cristalini; 4
Suaeda vera community, 5 Euphorbio paraliae-Lotetum glauci
Capelo, J. & aI.
The vegelalion 01 Madeira Island (Portugal)
July, the 23th -10. Caldeirão Verde walk (forest, epiphytic and waterfall vegetation)

A walk along a pleasanl levada waler course 01 greal beauly Ihrough lhe lauraceae loresl (Clethro-
Ocoteetum) and ils respeclive seral slages. Caulirrosetled (lsoplexido sceptri-Euphorbietum me/liferae)
and hygrophyllous vegelalion are also very exuberan!. The waler course runs Ihrough vertical clifts
several hundred melers high, wilh speclacular view over wooded valleys and clifts. II ends in a lhe
botlom 01 a greal circular pil wilh a beaulilul walerfall (Caldeirão Verde - lhe Green Cauderon). Along
lhe walk, several small cascades wilh walls covered by Oeschampsietum argenteae (Phragmito-
Magnocaricetea) [see lable 73, p. 177J are seen. This communily is associaled lo permanenl slow
laminar Ilow 01 waler. II has also a weallh 01 bryophyles. AI lhe botlom 01 cascades, a dislincl
communily is seen - lhe Peucedano lowei-Oenanthetum divaricatae [see lable 74, p. 178J, where
endemic umbellilerae 01 richer subslrala dominale. Epiphilic lern communilies are abundan!. The mosl
slriking are lhe Se/laginelo denticulatae-Cystopteridetum viridulae, Hymenophy/letum thumbrigensi-
maderensis (wilh Trichomanes speciosum, Hymenophy/lum thumbrigense, H. maderensis, H. wilson;;
and somelimes Elaphoglossum semicylindricum). Communilies on damp walls 01 Woodwardia radicans
bel ong probably lo a large-area associalion, also presenl in Azores and allanlic hyperoceanic Iberian
Peninsula (lo r relevés and lable relerences see walk # 6).

The walk ends by a walerlall wilh an elder communily which slands lor riverine vegelalion 01 upper-
strech slreams (Rhamno glandulosi-Sambucetum lanceolatl) [see lable 8, p. 84J.

Figure 8 C Id . - V d
-. a elrao er e walerfall: 1 Rhamno glandulosi-Sambucetum lanceolati; 2 Oeschampsietum
argenteae, 3 Woodwardla radlcans community
 The vegetation of Madeira Island (Portugal)
Capelo, J. & aI.
COSTA, J.C., EspíRITO SANTO, M. D., JARDIM, R. , SEQUEIRA, M, & RIVAS-MARTíNEZ, S. (2003)
Aknowledgements
- A classe Polygono-Poetea annuae Rivas-Martínez 1975 no Arquipélago da Madeira. Libro de
To Professor M. F. Lousã and Senior Researcher M. D. Espírito-Santo for stimulus and enthusiasm. resúmens das XIX Jornadas de Fitosociologia- Congreso International de Fitosociologia: pp117.
To Professor Carlos Aguiar for exalted discussion over a future common frame of macaronesian COSTA, J.C., CAPELO, J., JARDIM, R. , SEQUEIRA, M., LOUSÃ, M., EspíRITO SANTO, M.D. &
RIVAS-MARTíNEZ, S. (2004) - A vegetação da Madeira VII: A classe Molinio-Arrhenatheretea
vegetation.
Tüxen 1937 e Isoeto-Nanojuncetea Br.-BI. & Tüxen 1937 ex Westhoff, Dijck & Passchier. Silva
To Professor Salvador Rivas-Martínez for sharp scientific criticism, unconditional friendship and Lusitana 11 (2): 251-256.
guidance. COSTA, J., CAPELO, J., JARDIM, R., SEQUEIRA, M., EspíRITO SANTO, M.D., LOUSÃ, M.,
FONTINHA, S., AGUIAR, C. & RIVAS-MARTíNEZ, S. (2004) - Catálogo sintaxonómico e florístico
das comunidades vegetais da Madeira e Porto Santo. in J. Capelo (ed.) A paisagem vegetal da Ilha
da Madeira. Quercetea 6: 61-185.
COSTA, J., CAPELO, J., JARDIM, R. & SEQUEIRA, M. (2004) - Catálogo florístico do Arquipélago da
References
Madeira. in J. Capelo (ed.) A paisagem vegetal da Ilha da Madeira. Quercetea 6: 187-200.
BARBER, J.C., FRANCISCO-ORTEGA, J., SANTOS-GUERRA, A., TURNER, K.G. & JANSEN, R.K. DANTON, Ph. & GUITIONNEAU, G.G. (2000) - Comptes rendu du voyage d'étude à Madere de La
(2002) Origin of Macaronesian Sideritis L. (Lamioideae: Lamiaceae) inferred from nuclear and Société Botanique de France. Le Journal Botanique de la Société Botanique de France. 11: 5-19.
chloroplast sequence datasets. Molecular Phylogenetics and Evolution 23 : 293-306. DE FOUCAULT, B. (2000) - Notes phytosociologiques sur la végétation observée lors du voyage à
BOHLE, U.-R., HILGER, H.H. & MARTIN, W.F. (1996) Island colonization and evolution of the insular Madere de la Société botanique de France (Juin 1999). Le Journal Botanique de la Société
woody habit in Echium L. (Boraginaceae). Proc. Nat!. Acad. Sei. USA 93: 11740 -11745. Botanique de France. 11: 21-28.
CARLQUIST, S. (1974) - Island biology. Columbia University Press, New York, Nev: York, USA DEL ARCO, M & RODRíGUEZ, O. (2003)- Las comunidades vegetale de Gran Cana ria. ln O.
CAPELO, J., COSTA, J.C., JARDIM, R., SEQUEIRA, M., AGUIAR, C. & LOUSA, M. (2003) - The RODRíGUEZ (ed.) Apuntes sobre Flora y vegetación de Gran Canaria (Guia de la excursión
vegetation of Madeira II: Woody caulirosetted communities of evergreen forest clearings: geobatánica de las XIX Jornadas de Fitosociología y Simposio Internacional de La FIP 2003): 71 -
Euphorbion melliferae alI. nova. Silva Lusitana 11 (1): 111-113. _ 134.
CAPELO, J., COSTA, J.C., JARDIM, R., SEQUEIRA, M., AGUIAR, C. & LOUSA, M. (2003) - The EspíRITO SANTO, M.D., COSTA, J.C., JARDIM, R. & SEQUEIRA, M. (2004) - A vegetação da
vegetation of Madeira III: Oiplazio caudati-Perseetum indicae asso nova and Rhamno glandulosi- Madeira VI: Comunidades nitrófilas dos campos agrícolas, dos pousios e das suas margens. Silva
Sambucetum lanceolati asso nova: two new hygrophillic forest association from Madeira Island. Lusitana 11 (2): 241-251 .
Silva Lusitana 11 (1): 113-116. , FONTINHA, S & CARVALHO, J.A. (1995) - Evaluation of the vascular Flora of Madeira's extreme east.
CAPELO, J" COSTA, J.C., JARDIM, R. , SEQUEIRA, M., AGUIAR, C. & RIVAS-MARTINEZ, S. (2003) - Boletim Museu Municipal Funchal, Sup. Nº 4: 23-275
The vegetation of Madeira VIII: Advances on the phytosociological survey of the Madeira FONTINHA, S. & JARDIM, R. (1999) - Notes on vascular Flora of Porto Santo's islets. Portugalia Acta
Archipelago. Silva Lusitana 11 (2): 256-263. Biologica, Série B, 18: 169-177.
CAPELO, J., COSTA, J.C., LOUSÃ, M., FONTINHA, S., JARDIM, R., SEQUEIRA, M. & RIVAS- HANSEN, A. & SUNDING, P. (1993) - Checklist of vascular plants of Macaronesia. Revised edition, in
MARTíNEZ, S. (2000) - Vegetação da Madeira (Portugal). I - Aproximação à tipologia Sommerfeltia 17: 1-295.
fitossocilógica. Silva Lusitana 7 (2): 257-282. JARDIM, R. , FONTINHA, S. & FERNANDES, F. (1998) - Pico Branco: a peculiar floristic site on Porto
CAPELO, J., SEQUEIRA, M., JARDIM, R., COSTA, J.C. & MESQUITA, S. (2004) - Guia da Excursão Santo Island. Boletim Museu Municipal Funchal, 50 (285): 43-57.
Geobotânica dos V Encontros ALFA 2004 à Ilha da Madeira. in J. Capelo (ed.) A paisagem vegetal JARDIM, R., SEQUEIRA, M., CAPELO, J., AGUIAR, C. , COSTA, J.C. , EspíRITO SANTO, M.D. &
da Ilha da Madeira. Quercetea 6: 5-45. LOUSÃ, M. (2003) - The vegetation of Madeira IV: Coastal vegetation of Porto Santo Island
COSTA, J.C., RIVAS-MARTíNEZ, S., CAPELO, J., FONTINHA, S., JARDIM, R. , SEQUEIRA, M. & (Archipelag of Madeira). Silva Lusitana 11 (1): 116-120.
LOUSÃ, M. (2000) - As comunidades florestais da Ilha da Madeira: composição _florística e JARDIM, R., SEQUEIRA, M., CAPELO, J., AGUIAR, C. , COSTA, J.C., EspíRITO SANTO, M.D &
contribuição para a sua conservação. /I Jornadas Florestais Insulares: pp. 51. Dlrecçao Regional LOUSÃ, M. (2003) - The vegetation of Madeira V: Lino stricti-Stipetum capensis, asso nova and
de Florestas. Funchal. Vicio costei-Echietum plantagini, asso nova, two new semi-nitrophylous associations from Porto
COSTA, J.C. , RIVAS-MARTíNEZ, S., CAPELO, J., LOUSÃ, M., FONTINHA, S., JARDIM, R. & Santo Island (Archipelag of Madeira). Silva Lusitana 11 (1): 120-123.
th
SEQUEIRA, M. (2001) - Bioclimate and vegetation series of Madeira Islands. Abstracts 44 do MENEZES, C.A. (1914) - Flora da Madeira. Funchal.
IAVS Symposium pp 218. Freising-Weihenstephan.
Capelo, J. & aI.

MESQUITA, S., CAPELO, J. & SOUSA, J. (2004) - Bioclimatologia da Ilha da Madeira: abordagem Quercetea 7: 123-173, 2005
numérica. in J. Capelo (ed.) A paisagem vegetal da Ilha da Madeira. Quercetea 6: 47-59. ALFA, Lisboa. Portugal
PRESS, J & SHORT, M. (1994) - Flora of Madeira. Natural History Museum. Londres.
RIVAS-MARTíNEZ, S., DíAZ, T.E., FERNÁNDEZ-GONZÁLEZ, F. , IZCO, J., LOIDI, J., LOUSÃ, M. & Azores Centrallslands Vegetation and Flora
PENAS, A. ed.- Vascular plant communities 01 Spain and Portugal . Addenda to syntaxonomical Field guide
checklist 01 2001. Part I. Itinera Geobotanica 15 (1): 2-432.
RIVAS-MARTíNEZ, S., DíAZ, T.E., FERNÁNDEZ-GONZÁLEZ, F. , IZCO, J., LOIDI, J., LOUSÃ, M. & Eduardo Dias', Cândida Mendes', Cecilia Melo', Dinis Pereira' , Rui Elias'
PENAS, A. ed.- Vascular plant communities 01 Spain and Portugal . Addenda to syntaxonomical
checklist 01 2001. Part II. Itinera Geobotanica 15 (2): 433-922.
RIVAS-MARTíNEZ, S., WILDPRET, W. , DEL ARCO, M., RODRíGUEZ, O., PÉREZ DE PAZ, P.L.,
GARCIA-GALLO, A., ACEBES , J.R., DíAZ, T.E. & FERNÁNDEZ-GONZÁLEZ, F. (1993) - Las INTRODUCTION
comunidades vegetales de la Isla de Tenerile (Islas Canarias). Itinera Geobotanica 7: 169-374.
SEQUEIRA, M., JARDIM, R., CAPELO, J., COSTA, J.C., LOUSÃ, M., RIVAS-MARTíNEZ, S. & LOCATION: Azores belongs to the biogeogralic region 01 Macaronesia, in North Atlantic Ocean, at
FONTINHA, S. (2000) - Estudo litossociológico da Madeira - Implicações no ordenamento. /I hall distance Irom Europe and America. Their nine islands extend in a width 01 615Km , between 36º 39"
longitude N and 25º to 31" latitude W. Belonging to Portugal, they constitute, politically, an autonomous
Jornadas Florestais Insulares: pp. 58. Funchal.
SEQUEIRA, M., CAPELO, J., COSTA, J.C. & JARDIM, R. (2001) - Contribuição para o estudo region.
taxonómico de Teucrium scorodonia L. sensu latissimo - Sectio Scorodonia (Hill) Schreber - em AREA: The total area is close to 2304 Km2, irregularly distributed by the nine islands: the three large r
Portugal. Colóquio de Fitotaxonomia. pp7. ISA. islands, S. Miguel, Pico and Terceira, include 69% 01 the total area 01 the archipelago.
SJÓGREN, E. (1972) - Vascular Plants Communities 01 Madeira. Boletim Museu Municipal do Funchal
ELEVATION: Irom Oto the 2351 meters 01 altitude.
26: 42-125.
VIEIRA, R. (1992) - Flora da Madeira. O interesse das plantas endémicas macaronésicas. Colecção VEGETATION: A total 01 six major vegetation types have been recognized , in agreement with the
Natureza e Paisagem nº 11. Serviço Nacional de Parques, Reservas e Conservação da Natureza. altitude, each one with several types 01 communities. The total inventory 01 the vegetation is still
Lisboa. incomplete and just about twenty communities were described. Among the most signilicant types, are
the coastal communities, the humid areas (Iagoons and salt grasslands, bogs, wet grasslands, streams
and lagoons), lorests (temperate Atlantic, subtropical, 01 clouds), Atlantic heathlands, grasslands and
lormations 01 recent lavas. The native vegetable communities' dominant species are, in majority,
endemics.

FLORA: The natural Ilora are constituted lor about 300 vascular plants with 27% 01 endemics and
about 1000 knowned tallophytes (mosses, lichens, etc.) , with an endemicity percentage close of the 5%.
However, for this last group, the total value is still ignored because the lack of studies.

GEOGRAPHY

Azores are an archipelago of nine volcanic origin islands and several islanders, located in Atlantic
North Oceano Biogeograficaly, bel ong to Macaronésia, designation of the medium-Atlantic archipelagos
of Azores, Madeira, Canaries and Cab Verde. 01 this region, Azores are the northwest, the more distant
more the mainland and, geological, the most recent. Important factor is its dispersion lor a vast area 01
the ocean, 6 I 5 km 01 distance Irom Santa Maria to the Corvo. The lirst and the closest of the European
continent, at 1400 Km Irom Lisbon is equality the oldest island, Sta. Maria. Corvo, on the other hand, is

, Departamento de Ciências Agrárias e Ambiente. Universidade dos Açores. Apartado 387 9700 Angra
do Heroísmo. Portugal. eduardomfdias @c1ix.pt;
Dias, Eduardo et aI. Azores Central lslands Vegetation and Flora

just 1600 Km lrom Newloundland, in the American continen!. The area and altitude 01 each one 01 the concentration 01 richness and lead to a translorm 01-almost alllandscape; in the second largest island _
nine islands are very distinc!. Corvo, being the smallest, with only 17 Km2, reaches 780 m 01 altitude, Pico - with poor soils, lields 01 recent lavas and the Irequent volcanic activities, it has smaller agricul tural
while São Miguel, the largest, with 746 Km2, has an altitude about 1103 m. The largest elevation is the occupation and a low population density, without important urban centres, being, still today, an island
Mountain 01 Pico Island, with 235 1 m, an isolated volcanic cone 01 extraordinary dimensions. Al though with vast natural spaces.
they're volcanic origin , Azores has a diversilied orography caused by many volcanic lorms and
ln the magnilicent Mountain 01 Pico is located the highest point 01 Azores. ln its slopes, appears a
materiais that modelled and stil l model the surface . The oldest areas, usually with a mantle 01 piroclastic
group 01 volcanic and still active apparels, Irom which Ilowed vast lields 01 basaltic lava, some 01
materiais, spreads Irom a large cal dera volcano, present round lorms with deep valleys and deep water
historical eruptions. They constitute privileged places 01 vegetation and endemic species, as the
lines in the high lands and alluvial deposits and coluvial in the low lands. The most recent areas, usually
Mistério da Prainha, in lirst plan .
modelled by eruptions 01 basaltic lavas, show a topography drawn by the volcanic lorms, Irom small
cones to the extensive lavas flows. No doubts, that the dominant lorms 01 the Azorean landscape are As general pattern, the distribution 01 the Azorean village can be seen as a ring around the iSland,
the large calderas, imposing volcanic structures that are in each island origin, in alignment along the Ileeing Irom the sea rigidities and rude conditions 01 the altitude. Like this, the altitudinal zonation is
main tectonic lines E-W. Most 01 the soils are 01 the type" andossol", but the" litossols" occupy a explained in the actual landscape: alter the coast, usually unoccupied, appear the agricultural lands 01
signilicant surface. Some evolved soils -vertissols- has also been identilied in the oldest deposits. The annual cultu res (in past, the wheat) or medite rranean (as the vineyard), the one that is lollowed the
complete classilication 01 the azorean soils and cartography are incompleted. The natural vegetation is, villages, in a parallel coast li ne; depending on the reliel, they are lollowed ag ricultural lields again or,
actually, restricted to the less lertile soils 01 organic type on lavas (histossols), lields 01 volcanic ashes more Irequently, permanent pastures. ln the hillsides North, always colder, is also possible lind the
(inceptissols) or 01 recent lavas streams. The azorean climate is strongly oceanic, with small temperate Iruit trees here or, in alternative, in the poor soils, the production lorests; the high areas 01 the
temperature change, high precipitation and humidity values. The average annual temperature is about mountain are lelt with thei r bushes and indigenous lorests, as source 01 natural resources, in matter
17.5 C, being February the coldest month, with medi um val ues 01 13.8C 01 temperature, in the areas noble wood (as the cedar) and lirewoods. The integration in U.E, leading to high production policies and
close to the coas!. Frosts are rare below 600 m 01 altitude, but night Irosts can happen above 1000 m. the avaibilility 01 machineries and others production lacilities, creates deep alterations in this model,
ln opposite, precipitation has an important difference Irom E island to the W, Irom 710 mm/year to 1592 appearing dissonant occupation lorms with the local ecological resources.
mm/year in Santa Cruz 01 the Flores Island. This effect controls the vegetation lormations at low
ln the most typical pattern 01 the Azorean landscape, the populations extend in a ring around the
altitudes as it correspond the presence or absence 01 a summery time. At the altitude areas has little
island at some distances 01 the coast, as the most effective lorm 01 use 01 the soil, leaving the low
importance, since the values here are always high lor the typology 01 these communities. More
lands, 01 better soils and hotter climate, lor the annual cultures (in past, the wheat) and the ones 01
important is the air humidity with annual values close to 75-80% HR at the coas!. The saturation values
measured altitude, 01 wettest and windiest climate , lor the permanent cultu res (in rule, the pastures).
are reach ed in many days 01 the year, in the high lands.
The high lands and the mountains we re lelt wi th the natural place settings, as source 01 resources
The climatic lacto r 01 major importance lor the azorean natural communities is the wind . The dominant (Iirewoods, wood, water, etc.).
winds Irequently reach high inlensity; appear Irom the quadrant S-W, with high values 01 humidity. The
wind, determin e the structure and the lorest communities' distribution. The natural landscape 01Azores
on ly met the man's translormation five centuries ago, when they were discover by the Portuguese VEGETATION
caravels . According to the historie descriptions, these islands were, essentially, covered with dense
The total knowledge 01 the Azorean vegetation is still incomplete. The classic understandings 01
lorests. The lirst man changes , even belore the land occupation , were the cattle introduction. ln the lirst
altitudinal zonation have been changed by recent studies that proved the exposed to the winds and the
years, the colonization was centred in selected areas, always close to coas!.
soil's humidity as the main lactors lor the communities' distribution. So, while the presence 01 the"
The extensive lorests we re essentially explored in the begin lor collects-wood, firewood, Iruits. The placic" in altitude soils (horizon 01 deposition 01 oxides 01 iron and magnesium, that turn them
several periods 01 economic richness 01 Azores, lirst pastry and urzela, alter the wheat, then the orange impermeable) originates the existence 01 vast wet areas in the endorreic plateaus, the characteristic
and, linally, 01 mil k, they show a crescent translormation 01 the natural landscape in agricultural areas , lormation 01 seas 01 clouds, above 700 m 01 altitude, lor the wet winds, allows the existence 01
Irom the coast to inland . subtropical lorests (cloud lorests). On the other hand, there's inlormation that, belore the human
presence, the best soils would have imposing lorests, with more than 20 meters height (attested by the
The historie events, the environment 01 each island (geology, reliel, area, resources , etc.), the
bars 01 indigenous wood lound). With the agricultural occupation 01 these soils, are only possible lind
historical course and the vaiue 01 the soils determi ne the translormation 01 the landscape and the
today forests with little more than 10 meters heigh!. The vegetation coastal presents three main types 01
human geography, acquiring specilic patterns in each one. For instance, while in the largest island with
communities: the vegetation 01dunes, dominated by Spartina versicolor. The other two, more Irequent,
better resources in soils-São Miguel -the initial distribution 01 properties , lor noble people, took the
Dias, Eduardo et aI. Azores Centrallslands Vegetation and Flora
are the vegetation 01 drained lavas and the one 01cliffs. The lirst and extremely rich in endemic species, Rumex or Ranunculus; and the wet grasslands 01 margins 01 ponds or bogs, with some beautilul
with a microzonation Irom the strip 01 small halophytes, as Spergularia azorica, close to the line 01 species as orchids, Myosotis and Alisma. These communities are, today, difficult to lind for the
water, going by the halophytes 01 large size, as Azorina vidalii or Euphorbia azorica, to the dense pressures 01 marginal pasturing and the progress 01 lorage species, initially sowed in the pastures. The
bushes 01 Erica azorica. ln the second type, the dominant specie depends 01 the clilf's geomorphology, wet areas (Irequent in Azores) are a complex group 01 communities. However, its study and
but the Festuca petraeae and the Erica azorica are, lor norm, important elements. The azorean lorests classilication are not already made. Among the communities already recognized are evidenced , lor its
belong to the macaronesian element, in its more Atlantic species and constitute relic elements 01 the ecological importance: I-The bogs , with important areas in Terceira and Flores islands, are classilied
Tertiary period in the European continent, 01 subtropical conditions, denominated by species" alive habitats, 01 great importance in the conservation 01 the water resources. Is interesting to verily the
lossils" per-glaciers. While Lauraceae lamily possesses great importance in the lormations 01 the other singularity 01 these lormations in these latitudes, they are geographically isolated, lor with typical
archipelagos , is, in Azores, substituted by more elements wet-tempe rate, as the I/ex, Vaccinium, Taxus elements just 01 the boreal areas; II-The helophytes bushes, with important endemic species, as the
and Juniperus species. However, the type 01 leal laurilolia is still the dominant and most 01 these lorests Plathantera micrantha or Oaboecia azorica. Just as the wet grasslands, also these suffer strong
belong to the type Laurisilva. Recent studies could individualize lour types 01 azorean lorests. ln the low pressures of the pasturing, being in extinction risk; III-The lagoons, a lot of times azorean symbols, are
lands, the mesic lorest lormations 01 Myrica faya-Picconia azorica, are structurally, poor. The areas 01 oligotrophics endemics communities of Isoetes azorica habitat. Its total natural patrimony, in matter the
productive soils and sheltered had a rich Laurisilva mesic which don't remain alive examples. The wet microscopic communities, is almost unknown and, in many cases, they could be extinct lor the recent
Lau risilva is the richest lorest 01 Azores. It grows to stocking altitude and it can reach more than 10 eutraphization 01 some 01 them or lor other pressu res that it has been taking, besides, to its total
meters height in the examples that remain. It possesses a large structural complexity; until seven disappearance.
different straits and great diversity 01 species , including rare endemics. The dominants are Laurus
azorica, Vaccinium cyiidraceum, Frangula azorica and Erica azorica. These lormations are already The azorean hyper humid Laurisilva grows only in altitude, in conditions 01 saturation of atmospheric
extremely rare, lor the great search 01 their resources, meeting small stains in São Miguel, Terceira and humidity and 01 the soil, because it lives in a permanent sea of clouds. Although the trees dominant are
Pico. The" lorest 01 clouds" is a hyperhumid Laurisilva, dominated by Laurus azorica, I/ex perado ssp. almost the sam e 01 the other laurisilvas, they constitute the strict habitat 01 many other species, as the
azorica and Juniperus brevifolia, more than the live meters and, structurally, complex and dense, being rare heyiphils (epiphytes on leaves). Its role in the collects is important in the water resources 01 these
taken almost impenetrable. Is difficult to deline strata, because most 01 the individuais are epiphytes or islands. However, its demand lor sheltered places has been taking to the disappearance 01 extensive
areas in the azorean mountains.
they are born in trunks died above the soil, which is permanently soaked 01 water collected by the cup 01
the trees. Therelore, the mosses, that collect ali 01 the available surfaces, take an important paper in the
cycle 01 the nutrients, above the soil, and in the control 01 the water's cycle. 01 this lormation, important
FLORA
unaffected areas exist. The cedar lorest is a mountain lormation, tolerant to the stress 01 the water's
permanent saturation in the soil. ln its typical structure, is lormed by a lorest 01 Juniperus brevifolia and Since the lirst botanical explorations 01 Azores, beginning sec. XIX centaury, it has been recognized,
a moss carpet (Sphagnum sp.) in the soil. Is an endemic lormation 01 Azores, with high mosses for indigenous flora 01 Azores, about 300 vascular plants. Its actual flora contains 1007 plants being, lor
diversity The sub-alpine lormations can only be lound in the Mountain 01 Pico, in the Pico Island, abcve that, the increment 01 707 done by plants introduced by man, most in the last lifty years .
the 1900 m. They are 01 bush scattered , dominated by Cal/una vulgaris, Oaboecia azo rica and Erica
azorica. Some species, in great extinction risk, are restricted 01 this community. The colonizer The largest biological diversities are reached in the endemic lorests 01 Laurisilva mesic. Its structural
communities 01 lavas are, basically, lormations 01 Erica azorica. The ecological effects 01 the insularity diversity, with more than seven strata, high covering 01 epiphytes, climbing plants and lerns give an
are very intense in the colonization processo Therelore, a group different Irom lormations doesn't exist, appearance 01 sub-tropical lorest. Associate to this elevated structural diversity there are numerous
but an aggregation 01 the local species with larger tolerance to the water's stress, with predominance 01 habitats of rare or in extinction danger animal and vegetable species. The areas that remain 01 these
Erica azorica and Myrsine africana. The presence 01 a carpet 01 mosses is dependent 01 the local water formations are extremely small and they are, usually, in private properties.
balance. The species classilied as endemics has been suffering important revisions, by two reasons: taxonomy
ln good conditions, the succession process can be, surprisingly, last and, in 400 years, to lorm a and better knowledge 01 other areas. Curiously, its total numbers have been maintaining stable around
mature lorest. Pico Island is particularly rich in these lormations, lor the youth 01 many 01 their lavas, but the 70 plants (tax of endemicity 01 23%). A numbers of the indigenous plants of Azores are common
these can ais o be lound in Faial and in Terceira. Three types of natural grasslands were recognized , in with the archipelago 01 the Madeira, being able to consider as a natural nursery 01 Azores. Typically,
Azores: the coastal grasslands, of Festuca, Polypogon and Juncus; the mountain grasslands, very rich Azores has few species for lamily, which is an indicator 01 a drap species tax: 81 lam ilies exist lor the
in endemic species, Iram their dominant ones-Holcus rigidus, Festuca petraea, Oeschampsia fo/iosa 300 vascular planls. 01 lhe 41 , that possess endemic plants , 17 are just represented by one species
and 80% don't have more Ihan two species.
and Luzula purpureo-splendens-rare but spectacular and gigantic herbaceous, like Lactuca, Angelica,
Dias, Eduardo et aI.
Azores Centrallslands Vegetation and Flora
The high insularity degree and lhe suave ecological condilions can be lhe main causes 01 lhis
3 - The. adaptation to actual production has been causing strong damages in lhe natural remain's
speclrum. There exisls just one endemic genus-Azorina-wilh a single species. Most 01 lhe endemic
communltles. Wlth the construction 01 roads and the needs 01 water, highways have tom many natural
species is" alive lossils", philogenelicaly primilive, relaled wilh lhe dominanl lamilies in lhe Tertiary
stams and most 01 the ponds and lakes meet in danger by lhe direct extraction 01 waler even some
European Ilora, partially exlinguished during the glaciations. I d .. F ' are
a rea y mlssmg. or the same process, it has been verilying the bogs drying.
The delicale flowers 01 lhe endemic planls suffer peculiar adaplations lo lhe insular land, as iI is the
4. The coas tal vegetation is threalened by the construction 01 summer vacation houses or tourist and
loss 01 slrong colours, lhe decrease 01 lhe size or lhe attraclion 01 polinizer and more generalislic lor the associated social structures.
insecls, as in the Ilower 01 To/pis azorica. Here also lhe own Iruits losl lhe dispersion capacily the long
dislances, lypical 01 lhis type 01 pi anIs, as lorm 01 assuring lhat lhe wind doesn't take them outside 01 The interest lor the tradilions, a lot 01 times with commercial ends, it came to accentuate the value and
the island. the ~earch 01 many species accelerating its extinction danger. Recently, that search relapsed on the
(Jumpe:us. brevlfolia), what has been taking to the illegal discount 01 hundreds 01 trees 01 this species,
The lerns are an inleresting group 01 the Azorean Ilora, witch deserves relerence. It exist about 80
puttmg ln nsk not only some 01 their populations and varieties, as well as natural communities where is
differenl lerns, but only 53 can be considered indigenous. Some, relics 01 the Tertiary, are very rare the dominant species.
species in the European continent, bul they can be easily lound in Azores, like Trichomanes and
Hymenophy/um. Strong elemenls in lhis group are the paleolropical tendencies with, lor instance, lhe . 5. ~ne 01 the important causes 01 the disappearance 01 species, in the last years, is related wilh the
lamily Lycopodisceae. Acording wilh the last revision, 15601 the 300 plants natives can be considered mvaslon 01 exotic species, introduced in Azores and turned aggressive. Pittosporum undu/atum in the
rare and, 01 these, an important numbers has not been lound in the last years . Therelore, the number 01 !ow lands, Carpobrotus edu/is in the coastal vegetation and Hedychium gardnerianum in the wet areas
species in extinction danger is very high and superior to 60, including important endemics as Myosotis, It has been conquering wide areas. A group 01 more than 10 exotic species can be considered a~
Ammi, Chaerophyl/um, Lactuca, Perical/is, Lotus or Euphrasia. The bryoflora (mosses and liverworts) is aggressive invaders lor the natural vegetation .
rich, comparatively with the vascular Ilora. There are classilied about 450 species, wilh a tax 01
6. Equall!., some introduced animais put in serious danger some species or cause signilicant
endemicity 01 5%, but new species are added ali the years. ln this group is present a large
alteratlons ln communities. For instance, the rabbit is pressing in serious danger some endemic
litogeographic spectrum: striclly boreal species, as the group 01 Sphagnum, they appear with other
herbaceous and cause disturbances in the effective lorest trees.
typically tropical; they are some curious species 01 bipolar distribulion equally Uust in small points 01
Northern and South Hemisphere). 7. The direct exploration 01 species or resources, Irom the natural areas, as the cut 01 trees-as the
Juniperus brevifo/ia-or collecting the peat in São Miguel (called, leiva) lor the pineapples productiO~
ln the most hidden parts 01 lhe azorean mountains is slill possible to lind areas no disturbed by the cause irreversible alterations.
mano There, some communities live in its purer lorm. Only in those conditions, some communities seem
to survive, as lhe wel grasslands (in lirst plan that has jusl been lound in the gapes between the natural 8. Genetic contamination, lor introduction 01 species or similar varieties, inter-islands, Irom or out 01
lorests and the oligolrophic lagoons. the archlpelago.

9. Diseases and curses introduced by the ungovemable importation 01 great amounts 01 alive
matenals (e.g. decorative plants, Iruit bowls, seeds, green lorages, etc.).
THREATS
The most dramatic destruction 01 the lorests is its mechanical discount lor substitution lor other,
The largest threats to the azorean vegelal palrimony can be synthesized as:
usuall y. paslure, This conduct has been translorming important areas in the last years, doesn't allow any
1 - Historic aggressions. ln the past, some exploration lorms took some species, lo a reduced number practetlon 01 the natural biodiversity. However, and because the natural areas that still subsist are
01 individuais. This number can be below lhe natural capacity 01lhe species in surviving. ln these cases, always in poor lands, the obtained incames 01 these translormations are very red~ced. '
direcl conservation actions , lor manipulalion 01 seeds, become necessary.

2 - The recent development 01 these islands, accenluated by lhe arrival 01 pressures and economic
supports 01 the European Communily, molivaled lhe occupation 01 poor soils areas, wilh natural
vegetation until then. ln lhe lasl twenly years, lhe areas 01 natural vegelation were reduced in 50%,
mainly lor pastures.
Dias, Eduardo et ai. Azores Centrallslands Vegetation and Flora

MAIN VEGETATlON TYPES AND ECOLOGY VEGETATION OF ROCKY COAST

INTRODUCTION This community extends on whole Azorean coast, with the exception 01 small places, lor
geomorphologic reasons (sands and tulls - geological lormations usually 01 origin in underwater
There are still in Azores signilicant areas with total degree 01 purity as the Mountain 01 Pico (Pico
eruptions, the case 01 the islanders) or human disturbance (garbage/trash deposits, constructions etc.).
island), Sta. Bárbara (Terceira island), Morro Alto (Flores islands) among other places, housing almost
the totality 01 Azorean ecosystems in it's original state. Actually, the most common lorm 01 Azorean coast is lava exposed to sea abrasion. As global
characteristics, we can consider these settlements always 01 low covering (up to 50%, but usually below
Azorean vascular includes 1007 species, including 300 indigenous species and 70 endemic Irom
the 30%), lormed lor Spergu/aria azorica, Euphorbia azorica, Corema a/bum ssp. azorica, Juncus
Azores and Macaronesia. About 50% 01 natural species are in extinction danger. It also exists in Azores
acutus, Festuca petraea, Asp/enium marinum and Crithmum maritimum. The width 01 these
275 species 01 mosses and 175liverworts. communities' Iloristic diversity is big, appearing lrom very poor lormations and always 01 low density in
The vegetation 01 the Azores is lor a signilicant part determined by the Atlantic climate. Because the areas 01 strong hydrodynamic and compact lavas, to rich lormations and 01 considerable structural
Azores are part 01 the Atlantic climate system (mild winters and relatively cool summers with heavy rains diversity (bushes, big and small herbaceous and micro chamaephyte).
scattered throughout the year), the vegetation has a constant supply 01 water. The Azorean Ilora is not
only characterised by the usual Atlantic plant species, but also by Mediterranean, tropical and a wide
range 01 cosmopolitan species. HALOPHYTIC GRASSLAND

Many plant species occurring on the islands are exotic and were introduced by the early colonisers as These are endemic lormations 01 Azores but nowadays very rare, due to the lact that the habitat
lood resources. They thrive well in the warm and humid climate and were able to spread quickly in the develops at lavou rabi e places to traditional agriculture.
absence 01 their natural enemies. The ginger plant (Hedychium gardenerarum) that originates Irom the An intense agricultural occupation delegated us small stains 01 these lormations in inaccessible or
Himalayas out-competes many endemic species plants in their natural habitat and has become a marginal situations, lor that, its distribution, structures and Iloristic composition, as well as its extension
signilicant threat to the survival 01 many 01 them. The Hydrangea macrophy/la was introduced 150 years earth-inside and interaction lorms with other communities, will always be unknown.
ago and since cattle do not graze it, it was used as a natural border in pastures. The natural vegetation
above 700 m is adense humid laurel-juniper shrub-Iorest. The islands 01 Pico, Faial and Terceira have Under this classilication are rocky coastal lormations, occupied by dense herbaceous in a relatively
the largest remnants 01 these unique native lorests. rich soil substrata, with Irequent disturbances, that impede the development 01 arborescent
communities. More Irequent species are Po/ypogon maritimus, Gaudinia spp., Sonchus tenerrimus,
Chrithmum maritimum, Briza maxima, Festuca petreae, Daucus carota ssp. azoricus and Lotus creticus.
COASTAL VEGETATION

The coastal area constitutes the strip Irom the area 01 the tides up to where, lor diflerent means, it COASTALSHRUBLANDS
became preponderant, the permanent deposition 01 sea salts in terrestrial communities.
The coastal shrubs appear as the last covered areas in the coast zonation and, sometimes include the
Coas tal vegetation includes sharp hillsides, lava Ilow lields and sand dunes. transition lor interior lormations. 3 diflerent communities 01 coastal shrublands were described: Erica
The landscape space occupied with this vegetation is very narrow. These biotopes, due to their shrublands, Faya tree shrublands and mixed shrublands.
geographical position and ecological Iragility have been suflering an extensive alteration 01 their original Erica shrublands: communities constituted almost totally (or totally) by Erica azorica with medium to
communities in result 01 the human action. The occupation 01 these soils is due to the richness 01 these high density. These bushes lorms Iloristic poor communities, with 1,5 m 01 maximum height, mono
lormations, but mainly lor the climatic conditions: these are the only places with Mediterranean stratilied, with ali 01 the individuais 01 the same size in places exposed to the sea winds.
tendencies.
Faya tree shrublands: dominated by Myrica taya and is characterized by Iloristic and structural poor
communities, also mono stratilied, in atmospheres with a certain shelter degree, in poor substrata
(recent lava) and with summer hydrologic stress. ln suave environmental conditions this Faya tree
shrubs can grow to larger complexity and Iloristic richness.
 Dias, Eduardo et ai. Azores Central lslands Vegetation and Flora

Mixed shrublands: ln peculiar conditions, 01 accentuated topography (recent lavas 01 aa type), Halophytic lormations 01 Juncus in loamy soils: Whenever these rare communities were identilied,
I '
probably with watery line at low depth, it may occur complex communities in a mosaic distribution. ln the similar ecological conditions were lound: irregular deposits 01 sands above a superficial mantle 01
highest parts 01 the lava, dense shrubs 01 Erica azorica, Myrica faya, Juniperus brevifolia and eventually basaltic lavas , at the tide levei, but protected by higher lava shrubs lormations that allows the passage
Myrsine africana, appears (as well as Pittosporum undulatum); in the lowest parts 01 lava is dominated of salt water at punctual conditions: high tides or storms; this system has also a Iresh water supply,
by bushes 01 Corema album, Silene vulgaris spp. marine and Chritmum maritinum, with herbaceous usually due torrential regime riversides or fresh water nascen!. With halophytic characteristics, much
elements 01 Festuca petraea, Carex hochstetteriana, Daucus carota ssp. azorica, among others. These more marked than previous Juncus communitie, and, for that, is a low-denslty blotope. ln more
very particular lormations are 01 great interest, but very rare and in extinction danger. expressive forms (Lages do Pico) has a high vegetal formation dominated by Juncus maritimus,
Solidago sempervirens, Apium nodiflorum and Carex spp., among others.

DUNES VEGETATION Humid grassland (in coastal lagoons): The vegetation here considered constitutes the remaining
communities' 01 the original structure 01 the Paúl 01 Praia da Vitória. Humid grassland occurs in the
The Azorean coasts have a weak capacity lor the development 01 natural sand beaches and still less lowest areas 01 Paúl, in lands permanently soaked and submits to immersion during a part 01 the year.
lor the lormation 01 dunes. Most 01 the places with sand beaches are restricted to small re-entrance 01 The communication with the sea is, still maintained through tubes, allowing a periodic flow of salt water,
the coas!. The "wide" extensions 01 sand dunes as in the bay at Praia da Vitória (in Terceira island) are unloaded, in the high tide, directly to the nucleus 01 the Paúl. On the other hand, Iresh water coming
extremely rare in Azores. from water hillsides draining and Irom the raise of the hydrologic levei in the winter also enters the
The Dune vegetalion includes: system. The Soaked Grasslands, here considered, are locating in permanently soaked area. The
dominant species are herbaceous, in a grassland structure. The most Irequent specles are Paspalum
(1) The beach, understood by the strip 01 sand immediately above the tides line to the beginning the distichum, Alisma lanceolata, Apium nodiflorum and Lythrum hyssopifolium.
dune Iron!. The area Ihat sulfers lhe direct effect 01 lhe tides doesn'l possess terrestrial vegetable
settlements. The superior part 01 the beach possesses very scattered settlements 01 Salsola kali; (2) Coastal lagoons of Salty Waters: Of the old lagoon salty waters 01 Paúl da Praia (and where it was
dune, where the recent uses 01 this area (beach used areas, port construction, etc.) provoked lhe lished eéls and sardines), it just remains a small permanent puddle and the ditches through where is
disappearance 01 the characteristic continuity 01 the dune system. Other pressures, by chance older done waters flows and re-flows. These correspond to areas with high quantity of deposits and with
(pasture and exotic plant introduction, lor example Cynodon dactylon) , destroyed great part 01 these draining waters. The superior stratum of vegetation is always emerged (Scirpus maritimus, Juncus
dune communilies. These lormalions are dominaled by Arundo donax, Salsola kali, Lotus creticus and acutus ssp. multibracteatus) and the inferior, frequently immersed (Paspalum distichum, Polypogon
Tamarix gallica. monspeliensis, Ruppia sp.). Other example 01 these coastallagoons can be seen in S. Jorge Island.

WETLAND COAST VEGETATION NATURALGRASSLANDS

These are very restricted systems in Azores. They are under intensive human pressure because 01 its Grasslands are natural communities, not implanted nor maintained artilicially, they are not used lor
location in places with great potential lor urban structures constructions. Most 01 these biotopes are domestic cattle pasturing and are dominated, by spontaneous or even endemic species 01 Azores.
associated to dune systems, as it is lhe case 01 lhe bay 01 the Praia da Vitória. They are dominated by • Coast grassland (described at coast vegetation)
Juncus spp. and other species as Solidago sempervirens and Scirpus maritimus.
Wet hillside grassland (described at wetland vegetation)
Several Wetland Coast Vegetation identilied in Azores:
Holcus grassland, this grassland community constitutes Azorean endemic lormations. The dominanl
Halophytic lormations 01 Juncus in oozy soils: Although not belonging properly to the dune systems, species is Holcus rigidus, with a clear adaptation to slope deposits conditions. Typically, these
these communities appear Irequently associated to them. ln Azores, these lormations occur in dune communities grow in the base 01 big mountains, being limited to the strip 01 slope deposits. These
depressions where line alluvium deposits were accumulated above the sand. The only lound place, up lormations are rare, and only recognized in a reduced number of places. Its typical form was found ln
to now, 01 this biotope was in a humid dune area 01 Riviera (Terceira island). Il's possible to distinguish the base of slopes, as in Rocha 01 Juncal (Terceira), Caldeira 01 Faial (Faial) or Caldeira 01 Stª Bárbara
lhe sequence 01 hygrophytes settlements, halophytes, xeric halophytes, and interstitial (out 01 the (Terceira). ln this last one, lorms vast grassland in the base 01 the valley Ihat surrounds the mountaln
prolile, beach). ln terms 01 floristic, in these lormation develops species like Danthonia decumbens, almost completely. This grassland type contains rich mosaics especially in wet organic matter
Cyperus longus, Scirpus maritimus, Spartina versicolor and Atriplex spp., among others. sediments, where Mentha aquatica, Tolpis azorica, Sphagnum palustre, Eleocharis multicaulis, Juncus

132 133
 Azores Cenlrallslands Vegelalion and Flora
Dias, Eduardo el aI.
Ihen, wilh probable slralum in Iheir walers.
effuses dominale, and slony mosaics of certain drainage, where grows Agroslis gracililaxa, Pteridium
aqui/inum, Poa trivia/is, Eriea azoriea, B/echnum spieant and Ho/eus /anatus. We enhance lhe presence Small Oligolrophic Lagoon: These small lagoons are in greal number in Azores. It's frequenl lo see
of some rare endemic species like Rumex azoricus, Laetuea watsoniana, Perical/is ma/vitolia and Ihese slandlng waler formalions suffering human provoked unbalances (infilling, eulrophyzalion and
Leontodon rigens. dlsappearance for ruplure of lhe impermeable layer), anolher show infilling signs or even disappearanc
for nalural reasons. e
Fesluca grassland is, probably, lhe mosl European elemenl of Azorean grassland. These grasslands
are well known Ihrow whole Europe, from lhe sub Medilerranean alpine lo lhe Siberian areas. ln The young lagoons possess , in lhe general, a low covering degree and a low florislic diversily.
Azores , only Iwo species represenls lhes e rich genera in Europe - wilh, ai leasl, 170 speCies. The Iwo Communllles of Llttorella unlflora and / or Isoe.les azo rica dominale lhe deeper zones, while less deeper
species of Festuea genera in Azores are endemic, and allhough, ecologicaliy, very differenl in Iheir parts. has compacl formallons of Llttorel/a untflora, Eleocharis palustris and Cal/itriehe stagna/is, ai lhe
habilal (Festuea petraea, coaslal; Festuea jubata, from mounlain), and rarely in conlacled (wilh lhe margln Juncus spp. and Eleocharis spp. dominant.
exceplion Flores and Corvo islands, in lhe ravines wilh more Ihan 700 m, Ihal connecls coasl and
Developed lagoons and puddles possess a larger florislic diversily, frequenlly associaled lo peal
mounlain), Ihey can be considered physiographic similar. The halophylic grassland, wilh windy/marine
formallon (unilke lhe youlhs). Llttorel/a untflora loses it's importance giving up place lo lhe E/eo h .
erosion/deposilion of lhe soil, where Festuca petraea develops forming regular cushions and promoles a/u t . d Ca/UI ne
. he stagnalis or /soetes azoriea. The submerged margin area possesses e ansa
soil fixalion. By olher hand mounlain grasslands, wilh windy/hydrologic erosion/deposilion of soil , where P s ns an
com~unlly domlnaled by Cyperus alternito/ius, Cyperus /ongus, Juncus articu/atus and E/eoeharis
Festuca juba ta also forms similar regular cushions, wilh obvious soil fixalion funclions. Associaled lo
multleaulls.
Festuca jubata (mounlain grassland) luffs of are, low herbaceous species as Poa trivia/is, Lysimaehia
azoriea, Centaurium seil/oides, Potentil/a sp. , Selaginel/a kraussiana and Holcus /anatus; in lhe spaces ln certain lagoons, carpels of Sphagnum spp. nol possessing lhe described settlemenls dominale lhe
among luffs, corresponding lo waler flowing lines are implanled Juncus effusus, Eleocharis spp. adJacenl sub-coasl area, bul ii is already a pealbog Ihal prolongs lhe hydrological basin of lhe lagoon.
Sphagnum spp. Thiscompl~x ?roup is today slrongl~ allered by lhe human inlervenlion, due lo exploralion of waler,
Deschampsia grassland is, certainly, a vegelable formalion of greal ecological inleresl in Azores. Ils allerallon of hillsldes basln of lhe slandlng waler formalion , pollulion and vegelalion cover alleralion.
dislribulion, allhough very reslricled, ii is coherenl wilh specific ecological condilions. This habilal on lhe
Slream (ribeiras) : The slreams, in Azores generally flow lhe sea. This happens due lo lhe
highesl mounlain above ecological limil of lhe Irees. II is easy lo nolice Ihal, in Ihese exlreme c1imale
geomorphology of lhe Islands, formed by mounlains of hillsides.
condilions (especialiy slrong winds and cold lemperalures), few olher species can accompany
Oeschampsia toliosa. The one Ihal do, use lhe cushions of Oesehampsia toliosa as sheller, as several For obvious reasons, Ihese waler lines suffered, since very early, slrong inlervenlions of lhe Man: Ihey
bryophyles, and olher like Se/aginel/a kraussiana, Centaurium seil/oides, Ho/cus rigidus and Potentil/a wer~ lhe flrsl source of energy (Iogelher wilh lhe winds) and of drinking waler. Bul Ihey were, also, lhe
spp. easlesl garbage conducls, lhe easiesl roads for Irunks Iransport and lhe easiesl source of waler for lhe
domeslic animais. Therefore, since the beginning of lhe colonizalion, Ihese habilats were,
progresslvely, altered, in their communities' constilulion, and in Iheir hydrology.
WETLANDS There are 3 Iypes of slreams: (1) Permanenl creeks corresponding to mountain slreams associated
These areas possess in Azores a greal ecological importance and dimensiono These habilals lo nalural communities Ihis arehydrologic regulators and, for Ihat, with permanent flows, although with
correspond lo areas whose soils, ai leasl periodically, are soaked or emerged by waler, for enough lime some seasonal vanatlon. ln thls mountain habitat, usually on a solid rocky bed, ii settles a peculiar
lo cause effecls in lhe local communilies' composilion. commumly of bryophyles, in a continuous carpet, formed by species of helophylic and hydrophytes
.Slanding Waler Welland: We include in Ihis group big lakes, wilh are moslly caldera lakes of n.atu~e. (Heteroclad~u~. heteropterum, Jubu/a hutchinsiae, Fissidens rivularis, Rhynehostegium
npanoldes and Schlstldlum nvulare) strongly fixed lo lhe substratum. (2) Permanent streams with the
oligolrophic nalure, as well as lhe small ponds, permanenl puddles of differenl Irophy slalus and lhe
developmenl of nalural communities witch are exlremely rare, nowadays, in the global of Azores. These
complex group of waler lines.
were Idenllfled lo Sla Maria, S. Jorge and S. Miguel islands. Some species have always been found
Large Oligolrophic Lake: Ihese lakes, allhough being predominanl elemenls of lhe Azorean landscape as~o.cla~ed lo Ihese slreams and, for that, can be used as indicative: Equisetum teIma teia, Nasturtium !

(S. Miguel and Flores island), lhe mosl explored nalural resource by lourism and an exlremely importanl offlcmalts and Apium nodiflorum. (3) Temporary slreams are without natural vegetation in their great
agglomerale of nalural palrimony, Ihey are, very superficially known in Iheir ecology and Iheir vegelalion maJonty.
was never described in full delail, excepl in some punclual examples. These lakes have big volumes of
Peatlands: The vegetation of peatlands areas develops at permanent or periodically flooded places;
waler, usually deep (for example Lagoa da Caldeira Negra, in Flores island has 108 m of deplh) and,
Dias, Eduardo et aI. Azores Cenlrallslands Vegetalion and Flora

these types 01 communities are dependent 01 this water, and origins peat lormation. exposilion, ii dominales lhe Laurus azorica, Vaccinium cy/indraceum, I/ex perado ssp. azorica or Erica
azorica.
Peatlands are Irequent in high altitude mountain areas, without or with little human disturbance,
where , several types 01 wetland communities may occur in an inter-dependent way. The development 01 Ombrotrophic peallands:
this natural vegetation is related with the high precipitation indexes and the lormation 01 placic, an
Here lhe slatus 01 nulritious available in lhe water Ihal leeds lhe pealland depends, primarily 01 lhe
impermeable iron soil horizon, or the existence 01 compact rocks on the surface.
rainwaler, lhe pH is low. Allhough could exist olher species, lhe best adapled lo Ihese condilions are
Minerotrophic peatlands: lhe genera Sphagnum spp. ln this group we include ali peatland Iypes, whose involving area
lopography makes Ihem dependenl 01 rainwater.
The status 01 nutritious existent in the water that leeds the peatland is signilicantly higher than the
existent in the water 01 the rain; this is due essentially to the external supply 01 nutrients, coming lor Blanked Sphagnum peatland characterized lor covering vast areas in mountain picks and in high
lateral translers 01 water or emergencies 01 water Irom depth. ln here we include dystrophic poals and plateaus, extending, per times, lor the adjacent hillsides, in areas with high leveis 01 rain and a
temporary pools. impermeable subtract. This blanket peat lormation is geomorphologically different Irom other sphagnum
peatbogs but is not possible to lind any Iloristic differences.
Dystrophic pools occur in peatlands with rich and acid water, and are Ilorist poor.
Juniperus lorested peatland is one 01 the most interesting habitat in Azores . The great singularity 01
The temporary pools are extemporary and their vegetation is dominated by Littorel/a unif/ora, this lormation is due; to have as dominant arborescenl species the endemic Juniperus brevifolia, and
Callitriche stagnalis and /soetes azorica.
Ihis genus is little known as representative 01 these lormations. Juniperus lorested peatland occurs is
Wet High Grasslands, dominated by Juncus effusus, Rumex cong/omeratus, E/eocharis multicaulis very wind exposed a place, that's why lew other species occur in this extreme clime situation, where
and Dryopteris azorica or Osmunda rega/is, in permanently soaked environmental. rarely, individuais 01 I/ex perado ssp. azorica and Vaccinium cy/indraceum emerge. ln the high
herbaceous levei, Cu/cita macrocarpa and Dryopteris spp. can be present, although as little developed
Wet hillside Grassland are located in a smooth hillside above an impermeable subtract with
individuais, as well as Juncus effusus.
Hydrocoty/e vu/garis, Lysimachia azorica, Ga/ium palustre and Sphagnum spp.
Raised bogs are theoretically the most evolved peatbog lormations, to witch, in Azores, ali bogs
Grassland in water nascent witch is extremely rich in Iloristic terms. lormations will tendo Nowadays these are rare type 01 Sphagnum dominate community given the
Basin Sphagnum peatbog are small current peatland that has result lrom the inlilling 01 a puddle or extreme conditions their evolution needs to reach this state. Raised bogs are very sensitive to external
pond in an endorreic basin. This peatbog is homogeny, with lhe waler levei mainlained above or ai lhe disturbances.
surface, whal has as consequence a very conslant and poor Iloristic cover, lotally dominaling by These bogs have a higher ombrotrophic plateau, Iloristic poor, and lateral margins, in a lower levei,
Sphagnum spp., wilh presence 01 lew other species as E/eocharis multicau/is. ln the margins, due to which might have some subsidiary sources 01 nutrients Irom external waters (beside rain), and 50 richer
nutrienls that are dragged throw lhe hillsides; dominale hummock communilies 01 Po/ytrivhum
in species.
commune, Cal/una vu/garis, among olhers.

Transilion Sphagnum pealbog, are similar lo basin lormalion bul this develops in large basin areas.
This is lhe mosl common Iype 01 Azorean Sphagnum communilies. ln Ihese Iransilion peatbogs lhe NATURAL FOREST VEGETATION
micro reliel is more evidenl and dispersed Ihrow the enlire pealbog surface. The accenluated struclure The Azores Natural Forest find theme self's in a dilluse ecological position with a strict relation with
01 hummock and hollow allows actives mineralization in hummocks and promole the progress 01 the other types 01 Azorean vegetation .
numerous other species. The hummocks show several vascular species: Po/ytrichum commune,
Cal/una vu/garis Potentil/a sp., Lysimachia azorica, B/echnum spicant, Lotus u/iginosus, Agrostis Recent studies contradict the opinion 01 some scientist that visited the Azores islands in the last
graci/i/axa, Deschampsia fo/iosa, Luzu/a purpureo-sp/endens, Ho/cus /anatus and several bryophyle century, which delended the non-existence 01 lorest lormations in this archipelago. The historical
species. ln lhe hollows, appear E/eocharis multicau/is and Hydrocoty/e vu/garis, besides several descriptions show that dilferent lorest types covered most pari 01 the island's territory. Nowadays most
Sphagnum species. 01 this lormations, especially in low altitudes, were destroyed, but we still can distinguish several types
01 lorests with different ecological tendencies; paleomediterranean, macaronesian, subtropical mountain
Hillside loresled pealland are, probably, lhe mosl alypical situation and lhe more characlerislic 01 and norlh-atlantic lorests, that correspond to unique sintaxas with there own structure and dynamic
Azorean peatland. It's an endemic lormalion 01 this archipelago. The habitai is lormed by two differenl processo
communilies: (1) a continuous carpel 01 Sphagnum spp. ; (2) Iree cover, where, depending on lhe
Dias, Eduardo et aI. Azores Centrallslands Vegetation and Flora

• Fayal Forest's permanent water soil saturation, resulting in poor habitat lor plants germination. Most 01 the seedlings
establish in the hummocks zones where they lind more suitable conditions lor growing.
These lormations are very rare nowadays and we can consider theme almost extinct. Their natural
areas are now occupied by vine culture or introduce vegetation, predominantly Pittosporum woods. The environmental conditions are characterized by humid wind exposition and high precipitation.
They develop in incipient recent coastal lava soils, rich in potassium, with moderate climate conditions Ilex Forest's - also called "cloud lorests" are adapted to extreme conditions 01 atmospheric humidity
(Iow wind exposure, meso-mediterranean precipitation values, medium temperatures). The structure and permanent cloud cover. They develop in acid lavas, between altitudes 01 700 and 900 m, normally
and Iloristic composition is very simple, dominated by Myrica faya and Picconia azorica. Some north exposed. The environmental conditions are characterized by an extreme wetness, high
endemics are associated with the herbaceous layer like Carex hochstetteriana and Po/ypodium precipitation and occult precipitation, low wind exposure and high hill shade. The ground cover is
azoricum. lormed by lorest peat, resulting Irom low lealed mineralization in result 01 the permanent water soil
"Laurilolia" Forest's saturation. ln lace 01 these specilic conditions this lorest's only exist in some islands, where they are
very rare.
These lormations are dominated by laureai species and they develop in moderate climate conditions
especially due to wind exposure. They can be divided in three sublormations: This lorest lormation was a complex interior structure, dominated by horizontal mosaic 01
hummocks/hol/ows and the dynamics process area very related with the hol/ows wetness and the
"Laurilolia" Mesic Forest's - are constitute by a high Iloristic diversity, being the natural lorest stand concentration 01 biological activity in the hummock.
with higher number 01 codominant tree species (Laurus azorica, Frangu/a azorica, Picconia azorica e
Myrica faya). The herbaceous layer is dominat by the pteridophytes Oip/asium cauda tum, Oryopteris lIex lorests show a high Iloristic diversity. The emergent canopy is dominated by lIex pera do spp.
azorica, Oryopteris crispifo/ia. It's also present some rare herbaceous species like Bel/is azorica and azorica and the canopy shows highs cover percentage 01 Juniperus brevifo/ia and Vaccinium
P/atanthera micrantha or P. azorica. cy/indraceum. The herbaceous layer is dominated by the pteridophytes Cu/cita macrocarpa and
Dryopteris aemu/a It's also common the presence 01 Trichomanes speciosum carpets. The epiphytic
The environmental conditions are characterized by low wind exposition, high precipitation, low layer is well developed and most 01 the trees are covered by epiphytic species like Hymenophyl/um
atmospheric humidity with no water soil saturation. tunbrigense and E/aphog/ossum semicy/indraceum.
Nowadays this type 01 lorest is very rare, being the invasion 01 introduce species and pasture Juniperus Forest's - develops in extreme conditions 01 wetness and wind exposition. They are typical
occupation the principal lactors 01 disappearance. lormations 01 expose mountaintops. This type 01 lormations can be dividing in two types:
"Laurilolia" Humid Forest's - with a high Iloristic and structural richness, this lorests is dominated by Juniperus woodland - is lormed by a continuous and homogeneous layer 01 Juniperus brevifolia, that
Laurus azorica, I/ex perado spp. azorica, Erica azorica and Frangu/a azorica. The herbaceous layer is creates shadowed underwood habitat and stops the development 01 Sphagnum spp. peatland., allowing
lormed especially by pteridophytes showing an unusual diversity 01 lern species (Cu/cita macrocarpa, only the grow 01 understory shadow adapted species.
Oryopteris azorica, Oryopteris affinis, Pteris incompleta and Oip/azium caudatum).
The epiphytic bryocommunity it's very characteristic Irom this lorest lormation, and it lorms a
The environmental conditions are characterized by low wind exposition, high precipitation, high bryophitic sleeve over the J. brevifolia branches.
atmospheric humidity with water soil saturation in some periods 01 the year.
Juniperus Wodland with Peat bog - it's a transition community between Juniperus Woodland and
Nowadays this type 01 lorest is also very rare. Juniperus Forested Peat Bog. It distinguishes Irom the anterior lormation Irom having a sparse canopy
Laurilolia" Hiper-Humid Forest's - in present times this is the most common laurel lorest, because it's 01 J. brevifolia permitting the development 01 Sphagnum spp. peat bog carpet.
the type that develops in higher altitude, where human impacts are less signilicant. It differs Irom the Erica Forest's - are adapted to severe conditions 01 wind exposition and low soil hydrological
other laurel lorests in the composition 01 the dominant species. Erica azorica disappears and gives availability. This community is very poor in species. It has a woodland structure, lormed by a
place to Vaccinium cy/indraceum, but the other dominant species maintain the same. The herbaceous homogeneous canopy 01 Erica azorica. The understory scrub vegetation is sparse mainly composed by
layer is lormed by Oryopteris azorica, Cu/cita macrocarpa and it appears Irequently Trichomanes Laurus azorica and Myrsine africana. ln the herbaceous layer its relevant the presence 01 Pteridium
speciosum. aquilinum and Oryopteris azorica. The herbaceous-bryophyte layer is well developed and dominated by
It shows a less vertical structure complexity, but in stead it has a larger horizontal structure complexity, Thuidium tamariscinum.
with the lormation 01 hummocks/hol/ows. The hol/ows are lormed by running water, creating a zone with
Dias, Eduardo et ai. Azores Centrallslands Vegetation and Flora

This formation develops on mountain picks, very exposed to the wet winds, on pomitic impermeable
SHRUBLANDS
substratum. This habitat distribution are a is restricted, because nor ali mountainous have such high
Vegetable formations, dominated by chamaephyte or microphanerophyte, mono stratify, dense or wetnwss. These bushes make the transition space between soaked shrublands, Juniperus Forests and
scaltered, with possible presence of trees emerging, in a casual or scaltered way. The shrubs can be Deschampsia Grassland.
low, when dominant species are true chamaephyte (as Cal/una vu/garis, Oaboecia azo rica and Thymus
caespititius); or arborescent, when the dominant species are microphanerophyte (Hypericum fo/iosum) Pedologic Mature shrubland
or mesophanerophyte of restricted size (Juniperus brevifo/ia, Erica azorica, Laurus azorica and Myrica These shrublands were found in circumstances where, for predominantly soillimitations, the vegetable
faya). ln Azores there are several of shrubland habitats types, following described. community limited in its structural complexity, forming low shrublands in response to these limitations.
Their presence is due to local conditions, wetness in inhospitable geological substratum that will need
several thousands of years to allow a more complex colonization.
Local stress shrubland habitats
Mountain arborescent shrubland appears in sheltered mountainous hillsides, but with great entrance
Observed circumstances show that the evolutionary potentiality of shrubs formations is annulled by a of water, of hillside drainage or fog water interception. ln it's structure possesses an irregular Sphagnum
permanent or periodic external factor that transforms the bus h dominated habitat in the local potential carpet spp. under a community of Laurus azorica, I/ex perado ssp. azorica, Vaccinium cy/indraceum; in
formation or opens possibility for an extremely slow temporary evolution. ln this case, are, for instance, the most exposed places, the vegetation is more open, prevailing the Juniperus brevifo/ia and Cal/una
the coasts formations on high to medium slope hillsides, typically covered by a almost pure bus h vu/garis, in more rocky subtract Erica azorica may dominate.
formation of Erica azorica witch density and height depends of salty winds intensity. These formations
should be considered as mature, for the semi-permanent winds action, and for the age and demography Calluna wet shrubland: this community constitutes the mature form of Cal/una vu/garis shrubland
that some possess along the Azorean coasts. (Cal/unetum, heathlands) in Azores.

Lajido shrubland
Coastal shrublands:

Erica shrublands (described in coast vegetation) The term lajido is used at Pico Island to designate flat basaltic compact lava surface where the
penetration of the roots is impossible, as well as water and deposit surface retention. Their surface is
Faya tree shrublands (described in coast vegetation) almost nude of vegetation, especially is more exposed parts. The only species installed are growing in
Mixed shrublands (described in coas! vegetation) lava fissures being, for that, a scaltered and liltle developed community. Species like Erica azorica,
Myrsine africana, Laurus azorica, Cal/una vu/garis, Oaboecia azorica, Thymus caespititius may occur.
Alluvium shrublands:
Volcanic sand shrubland
Appear in the base of hills or scarps, on fine slope deposits, corresponding to places with some
disturbance. This habitat possesses some ecological similarity with described Holcus Grassland, with The presence of bushes in this substratum is quite rare, in Azores. This substratum is of easy
which, per times , it forms mosaics, but in this case it constitutes a less disturbed situation. Alluviums colonization, reason why is almost ali occupied with cultures or with degraded by intense exotic cover.
shrublands are open communities with reduce structuring. Two cases were inventoried: There are 3 volcanic sand shrubland found, in Azores: Graciosa, (Pico Timão); Pico (sand mountain
formation); and Faial (Cabeço do Fogo).
Erica mixed shrublands occurs on acid gravei slope deposits, and is dominated by Erica azorica and
Juniperus brevifo/ia, it's a open vegetal community, with scaltered grassland mosaics. ln this grassland Predominant species are Thymus caespititius, Oaboecia azorica, Racomitrium /anuginosum,
species like Ho/cus rigidus, Festuca petraea, To/pis azorica, Huperzia se/ago ssp. se/ago and P/atanthera micrantha and Centaurium scil/oides; in costal volcanic sands Corema a/bum is identified.
Lycopodium cernuum develops. Successional Scrubs
Euphorbia shrublands These groups of formations correspond to primary and secondary succession scrubs. The pioneer
Euphorbia stygiana ecology of, an Azorean endemic, is still very badly known. Their settlements are scrubs developed in recent lava flows. The colonization velocity and the typology of these communities
very small and dispersed for several habitats. However, this formation occurs with some regularity in showa great variability. The colonization processes varies not only with the age, but also with the type
slopes or deposits of mountain slopes, giving the indication that prefers habitats with some disturbance. of subtract material, structure and morphology, the local bioclimatic conditions and trom island to island.
There are no specific colonization communities, the floristic composition is very similar with the climaxes
Mountain shrublands communities, and however they have important structural differences. This effect is possibly related with
Dias, Eduardo el ai. Azores Cenlral Islands Vegelation and Flora

lhe insular elfecls, in particular florislic poverty of lhe islands. The zoom effecls seem lo acl as one FIELD TRIP INTREPERTATION SHEET
import succession processo

1 PICO ISLAND

142 143
Dias, Eduardo el ai.
Azores Cenlrallslands Vegelalion and Flora
1 - COAST VEGETATION OF CACHORRO forest. ln less dry areas more complex communily will develop wilh Picconia azorica and Laurus
azonca.
The coasl vegelalion of Pico Island ai Cachorro - Lajido área has a zonal dislribulion, as expecled.
The firsl delerminanl faclor is lhe hydrodynamics of lhe ocean, inducing lhe sall spray and wave's On lhe pahoehoe lava fields , lhe deep cliffs are usually absenl of vegelalion (3), on lhe lop fiai, lhe
invasion of lhe lerreslrial coasl areas. These coaslal areas of Cachorro are facing North, where lhe lerreslnal vegelalion slar wilh lhe communily of Spergu/aria azo rica and Crithmum maritimum very
ocean slorm are inlense ali lhe year, wilh slrong winds and high waves. II is common lhe waves sparce ln lhe flssure (4). The second zone is a communilie of Festuca juba ta wilh annual grasses (5).
achieve 7 melers high and lhe sall-waler can washes areas above 20 melers slopes. The erosion is Very olten Ihese lava can have already permanenl humidily in lhe fissures. Then a lall herbs communily
very inlense and only because of ils basallic composilion ii can be so swallow. Mosl of lhe North face can develop wilh Foenicu/um vu/gare and Daucus carota ssp. azorica,
coasls are deep slopes (achieving 1.000 melers), cause by erosion pressure. Even here we can only
find swallow areas in deep sellzer bays. The waves erosion can also be undersland from lhe exlended Before lhe bushes zone (6), ii is frequenl lhe formalion of wel condilion, were a communily of Juncus
zonalion in Ihis coasl, wilh lhe shrub vegelalion only presenl very far from lhe coasl line, and a large and SO/Idago sempelwens developed. ln lhe upper faces of lava, Erica azorica and Myrica faya slart lo
grow.
"desert" zone in lhe fronl line.

The second faclor is lhe balance belween lhe sall in lhe subslrala and lhe fresh waler. Sall spray The nexl slage is adense shrub formalion (7), wilh Erica azorica, Myrica taya, Juniperus brevito/ia,
deposils produce lhe halophylic condilions, bul ais o change lhe pH condilions, acid in mosl of lhe island Corema a/bum (some places), and Myrsine africana. Under layer, Carex hochsteteriana is common.
soils. The fresh waler can be abundanl in lhe coasl areas as a laminar flux, developing small ponds in
lhe winlertime, and ii has one effecl of washoul lhe sall and minimized lhe waler slress. ln some Soullh
swallow coasl areas, lhe foresl vegelalion can achieve almosl lhe sea line by Ihis effect. ln here, ii has
one importanl effecI, as lhe lavas slruclure, drained from lhe mounlain, in ils layers, importanl amounl
of fresh waler. II gives lhe possibility of lhe shrub vegelalion arriving so far in lhe coasl line, wilh lhe
presence of Iypical mounlain species, like Myrsine africana, Juniperus brevifolia or Lysimachia azorica.
II also explains a fronl line of wel vegelalion alter lhe shrubs, wilh Juncus spp.

The Ihird faclor is lhe geologic slruclure of lhe rocks bed. Here, mosl of lhe coasl is of pahoehoe
basallic lavas flow. They gave a platform almosl fiaI when Ihey flow, and lhe sea erosion broken lhe
fronl as a slrail rocky cliff. They are very difficull lo be colonized unlil Ihey have fresh waler near lhe
surface, and mosl of lhe fronl areas have low-densily vegelalion, only in lhe fissures.

Some areas have aa lavas, like Cachorro, and lhe vegelalion communilies are quile different. Very
easy lo colonize, Ihese lava rich nulrienls, allow lichen communilies in lhe exposed surface and dense
vascular communilies in lhe fissures. They relain also, as lhe erosion of waves slopped, large amounls
of clay and a halophilic grass can become slablished, dominaled by Festuca petraea.

ln very small shelled areas, lhe coasl can be build up wilh rock deposils of differenl sizes. Here xero-
halophilic communilies can be dominanl in lhe fronl line. Bul lhe shrub vegelalion arrived very fasl,
usually wilh Myrica faya or, in some islands, wilh Dracaena draco.

ln lhe Cachorro - Lajido area we can find, on lhe aa lava beds, a firsl communily of Spergularia
azorica sparce in lhe fissures (1), wilh Asplenium marinum and Atrip/ex spp. On second line (2) a
communily of Festuca petraea slarts in lhe fissures and covers ali lhe areas as lhe dislance lo sea line
increase.

The Iransilion belween Ihese communilies and lhe shrub zone slarts wilh lhe presence of Erica
azorica shrubs, sparse in lhe grass carpe!. The fronl of dense shrub zone is usually wilh Erica azorica.
Myrica faya , in lhe dry areas can dominale from large areas inland, increase in size unlil arriving lo a
Dias, Eduardo et aI.
Azares Centrallslands Vegetation and Flora

2 - THE LAURUS FOREST OF AZORES

At the middle altitude on Azores, the climate conditions are excellent lor most plants grow and similar
to a temperate greenhouse. The average temperature is about 18ºC, with low variation along the year,
the rain is about 1500 mm, distributed by ali season, and no stress is usually presen!.

It should also be notice that there are no naturallarge vertebrates on Azores, so the animal pressure
over the natural vegetation was very small. Even the radiation is smooth with very lew days on the year
with clean sky. Most 01 the days, in lact, there are a cloud cover.

ln these condition, under developed soils and if no local disturbance arrived (Iike lava Ilow or Ilooding)
the potential vegetation is a tall Laurus lores!. It is a very complex lormation, achieving, nowadays 20
meters , with more than 7 stract and 300% 01 cover. We have been called 01 mesic Lau rilolia lorest and
it belongs to the classical Laurisilva lormations. The number 01 areas nowadays with this community is
very small and can only be found in inaccessible sites. The good soils have been a major attraction lor
larming.
They have one emergent canopy (1) 01 one species dependent 01 the regional conditions : in less
altitude places it is usually Myrica faya, and a continuous under canopy 01 Laurus azorica, Fragu/a
azorica and Picconia azorica (2). ln the higher altitudes the emergent canopy can be 01 /lex perado ssp.
azorica and Juniperus brevifo/ia been present in the continuous canopy.

A second, less dense canopy, of young trees of Laurus and Picconia (3) can be present, usually
supporting climbers like Rubus and Ruppia agostinhoi. A third stract 01 shrubs (4) , includes Myrsine
africana, Hedera azorica, Vaccinium cy/indraceum and most 01 small plants 01 the dominant trees.
Lower it started the stract of lerns, with large ferns making adense cover (5) with Cu/cita macrocarpa,
Dryopteris azorica and Athyrium fi/ix-femina and a second strat 01 small lerns (6), like Dryopteris
aemu/a, B/echnhum spicant, Phy/litis sc%pendrium and, some times, even Trichomanes speciosum,
associated with tall herbs, like Carex vu/cani, C. peregrine.

At this levei, the lorest is more or less impenetrable, as the dense lerns and climbs are associated to a
dense cover 01 dead trunks and branches. The absence of large vertebrates results in adense cover
near the soil. As the old description said , it is possible to walk in the lorest 01 Azores without to walk on
the soils.
The low stract of vascular is formed with slender plants like Be/lis azorica, P/antanhtera micrantha or
Prune/la vu/garis. Below them, a carpet 01 bryophytes usually covers almost ali the soil.

The epiphytic communities have also dense mousses carpet (7) with lerns like Po/ypodium azoricum,
E/aphog/ossum semicy/indricum and Hymenophy/lum spp. (8). ln very shelter conditions, spectacular
curtains 01 Neckera can be presen!.

Ali 01 these lormations are endangered, most 01 ali by the exotic plants like Pittosporum undu/atum.
This Australian tree has arrived to ali 01 them and a large number have been completely transformed as
this tree become dominant or replace ali the natural vegetation.

146 147
 Azores Centrallslands Vegetation and Flora
Dias, Eduardo et aI.
3 - PICO ISLAND CENTRAL PLATEAU LAVA VEGETATION AND ECOLOGY

Azorean volcanic islands are testimony of successive eruptions that were in their origino Over time,
lava fields undergo progressive changes. Wind and water wear down the volcanic rock, starting with the
softer materiais. Streams with a steep gradient can sometimes cut through a lava pile. Heating and
cooling by sun and seasonal changes further expand and contract the lava masses, aiding in their
break-up. These processes move very slowly. Eventually men activities can interfere in these natural
succession mechanisms altering lava structure and plants re-established in the nutrient-rich volcanic
soil.

ln Pico island central plateau landscape the vegetation develops above rich nutrient and impermeable
basaltic lava. Its surface has a wrinkled or ropy-Iooking surface appearance. This lava is the substrate
of very different vegetation mosaics, what can be due to the fact that it is a extremely irregular lava
subtract, with quite different environmental conditions associated to each part of the relief. Besides the
impermeable nature of this lava there are some ruptures from where the water flows. But main water
fluxes on this formation occur through the space existent between different layers that result from
different eruptions.

The vegetation zonation of these lava fields varies with environmental factors like height of the
substratum, exposition to the wind, and wetness degree but it also depends and changes with human
activities and soil uses intensity. Generally speaking is possible to distinguish at least 4 types of
vegetation communities occupying different heights of the lava substrate.

(1) ln raised parts of the lava, corresponding to less wet conditions, there are dense mosaics of
forested formations. ln wind exposed places this forest is dominated with Juniperus brevifolia and
Vaccinium cylyndraceum with scaltered herbaceous species like Tolpis azorica, Rubia agostinhoi and
Holcus rigidus. On other hand, wind protected places on high zones of the field lava has different forest
dominant species, like Laurus azorica, Frangula azorica and I/ex perado ssp. azorica with a lower fern
stratum dominated by Drypotris azorica. ln the middle slope of the lava species like Cal/una vulgaris and
Erica azorica dominate with herbaceous stratum of Lysimachia azorica, Holcus rigidus and Hypericum
foliosum, in lighter expose places. Nowadays in Pico middle high pastures or natural grassland pastured
areas, these forest formations were pushed to the higher parts of untransformed lava, and represent the
remain of original forest formation that survived landscape human transformation. Pico pastures are
extremely rich on vegetal species, due to these dense lacy of hedges in human landscapes.

(2) ln the intermediate zone of lava, the line of water is visible but the mosaics grow in small
hummocks above the water leveI. ln here herbaceous dominate the vegetal community. Hypericum
foliosum and Anthoxanthum odora tum are present but scattered. The dominant genus is Carex. ln more
wet formation Carex nigra is the commonest.

(3) ln the lower area of the undulated lava, water is accumulated and conditions are favourable to
peatbogs occurrence, in witch Sphagnum species form adense carpet, enhancing Sphagnum
auriculatum and Sphagnum subnitens. Other species seam adapted to these extreme wet conditions,
like Lycopodiel/a inundata, Scirpus fluitans, Eleocharis multicaulis, among others. The transformed lava
 Dias, Eduardo et aI.
Azares Centrallslands Vegetation and Flora
lields or the pastured . natural areas tend to loose the peatbogs and replace them with lens, more 4 - VEGETATION OF HISTORIC LAVA FLOWS MISTÉRIOS DA PRAINHA
herbaceous commumtles due to the nutrient enrichmenl 01 waler and direct animal Irampling on
Sphagnum sensltlve carpe!. Thls changing can be seen in places like margins 01 Capitão lagoon, and Mislérios da Prainha is a large carpet 01 lava lrom one hisloric eruption (1572) 01 "Bocas do Fogo"
PICO da Urze. On lhe olher hand, places wilh more recent (and less translormed) natural lava is Paul's volcanos. II spread Iram the higher part 01 lhe island in a large carpel until the northern coastline.
lagoon and Caveira. "Mistérios" means in Portuguese mysleries, and was used to name the historie eruption, as the people
didn'l understand whal was happening.
(4) ln some large and deep basin zone on the lava lields, waler can be accumulated in such quantities
thal mlghl lorm small pools where hydrophytes like Litorel/a uniflora, Potamogeton polygonifolius, ln these lava lields, differenl lava colonization process can be lound as Ihey have differenl ecologic
among others, can grow. conditions Iram the wet high mounlains unlil the drier lowlands. On the high part, the lava Ilow lormed a
plateau and Iwo different lava Iypes can be lound: the aa and the pahoehoe lava. The lirsl had Ilowed
This lava lields are, biologically, exlremely important, once iI supports several protecled and rare as waves concentric to the chimney, with a very rough surface, generally clinkery and lull 01 liss ures
Azorean Habitats. II is also habitat to several protected species: Arceuthobium azoricum Erica azorica where the roots can grow. The seconds, pahoehoe, Ilows tend lo be relatively Ihin, Iram a lew inches lo
Frangula azorica, among others, and also endemic species, lar example I/ex perado ssp.'azorica, Tolpi~ a lew leet thick and with a smooth, billowy, ar ropy surface. ln this very difficult surface lar plants
azonca and Rubia agostinhoi. ln Ihese lava studies we lound pealbogs with more than 2 m deep, colonization and more stress toleranl ones are dominant here.
representlng an exlremely import waler retention slruclure, especially in islands with a limited water
resources. The aa lava waves are colonized by Iwo different communities, side by side, one in the top 01 lava
wave, and lhe other in lhe valley. The lirst one is a slress tolerant, associale to dries subtracl but windy
and humid air conditions (1). Erica azorica, Cal/una vulgaris are the dominant shrubs. The Oeschampsia
\.. '
" foliosa and Holcus rigidus communities can cover lhe mosl exposed areas. Some rare species are
associaled lo Ihis habitat, like Planthantera micrantha and Palhinhea cemua ar Euphrasia azorica.

The depressions (2) are a dillerent habilal, very wel, somelimes Ilooded, and rich in nulrients Iram
chemical erosion 01 lava surface, sheller and wormed. II has a high-densily communily 01 Juniperus
brevifolia -I/ex perado ssp. azorica, with almosl ali lhe species 01 lhe malure loresl and some olhers 01
the colonization slage, like Huperzia spp. The rare species 01 Ruppia agostinhoi and Euphorbia
stygiana can be quile abundanl here, and iI has some 01 lhe rare populalion 01 Oaphne laureola . The
bryophytes cave r ali the subtract and mosl 01 lhe endemic species can be lound. The dense cover 01
lerns, including lhe Culcita macrocarpa have lhe same size as the vascular shrubs, as ali are limited by
lhe high 01 lhe near summit 01 lava wave: ali 01 this dense and complex communily, similar lo one ellin
loresl, are unable to grow on very exposed condilion.

The pahoehoe lava has a much low vegelation slage. The dense surface 01 lava can only be cover by
bryophytes and lichen. Mosl 01 lhe higher vegetalion is rooted in lhe lissures. The dominant species are
similar to the one 01 upper part 01 Mounlain, with Cal/una vulgaris, Oaboecia azorica, and Thymus
caespititius. The black rock can became very warm in the summer time, and an annual community can
be lound in the small depressions 01 the Ilal surface with Agrostis gracililaxa.

The volcano chimney, "Bocas", has slill the main composition 01 piroclast deposits at surface. Adense
heath 01 Erica azorica is the dominanl colonize r communily (4) with Oescampsia foliosa and Festuca
juba ta.

150
151
 Azores Centrallslands Vegetation and Flora
Dias, Eduardo et ai.
5 - WET SYSTEM OF CAVEIRO

Caveiro volcano systems are located in the highest point 01 the East side 01 Pico Island, achieve 1065
meters 01 altitude. It is a complex 01 lavas Ilows and dome volcano's over an older pomitic deposi!. This
one is impermeable to water by placic lormation. On these conditions, the volcano slopes works as a
giant water collector Irom the clouds and rain and conduct it to the valleys, were it arrives rich in
nutrients and lorm mires.

By the orography 01 the island, with the high Pico Mountain, at West, these are become the meeting
point 01 the clouds transport by the trade winds. These mountains are almost ali the year emerging in
clouds and the precipitation arrives to values 01 3500 ml/year. But the etfect 01 intercept the clouds
water and log increase these values 3 or 4 times. Meteorology station at the summit 01 Terceira Island,
at the sam e conditions, have collecting data 01 12.000 ml/year as lhe total 01 water collecled by lhe
syslem. Under these condilions lhe environmenl is ali-time saturate and lhe mineralizalion is nol
possible, lead lo accumulation 01 pea!. The lorest communilies can only grow on the lava slopes,
developing a special case 01 loresled bogs. The Ilat areas wilh water deposits have Sphagnum bogs lo
len, depending 01 nutrients slalus in high dystrophic waler. The transilion zone belween the lorested
bogs and lhe open bogs has a mixed lormalion 01 Juniperus brevifo/ia, an endemic tree thal can support
permanenl immersion, with tall nitrophylous herbs, using lhe disturbance gaps. Ali the system is
interconnected and support periodic dislurbance with lhe storms. Dieback processes were describe lor
the Juniperus bogs communilies and are under quantitalive description. The palches areas are
important relugees lor rare nitrophylous endemic pi anIs and seems to be an import source 01 nutrients
lor the open bogs, as a related change in sphagnum density have been observed.

These lorests syslems only depend 01 water and nutrients Irom clouds can very much be considered
as ellin cloud-Iores!. The high summits and the directly exposed slopes to trade winds have a Juniperus
brevifo/ia peat bog. Depending 01 lhe dieback stage, lhes e communities can have ditferent densily 01
sphagnum species, almost absenl under lhe mature stage 01 the lores!. This lorest has adense
endemic epiphy1ic community associaled that covers ali the trunks and branches where most "soils"
processes take place.

On the shelter slopes 01 the lava Ilow and dome, a wet lorest (1) dominated by lIex perado ssp.
azorica as emergent canopy and a under layer 01 Laurus azorica, Vaccinium cy/indraceum and
Juniperus brevifo/ia, are established. These lormations have a mosaic structure, as the depression 01
the lava is lilled wilh lorest peat, and the trees and shrubs can nol rooled. Only the lerns and mosses
can occupied the margin 01 Ihese areas. ln some cases, Irees to root use the tree like Cu/cita
macrocarpa trunk, given one kind 01 lorested hummock. A dens epiphy1ic community cover ali lhe upper
surfaces, including grasses carpets 01 Luzu/a purpureo-sp/endens, Carex vu/cani and Carex peregrine
and Ho/cus rigidus. The lerns are also abundant with large lerns like Dryopteris azorica, or the sensilive
lerns 01 Hymenophillum spp. and E/aphog/ossum semicy/indricum. ln the clirnbers, Hedera azo rica and
Smi/ax divaricata are presen!.

ln the base 01 slopes, a transition community is present, dominaled by Juniperus brevifo/ia (2). It
changes in calenal etfect Irom high density with lorest peat to open shrubs rooled in hummocks 01
Dias, Eduardo et aI. Azores Cenlrallslands Vegetalion and Flora

Polytrichum spp. on a Sphagnum carpe!. The water erosion open large gaps, where rare endemic herbs FIELD TRIP INTREPERTATION SHEET
are associated, like Angelica lignescens, Lactuca watsoniana and Ammi trifoliatum.

The flat areas, Ilooded, have complex communities 01 Sphagnum bogs and lens (3) with a distribution 2 SÃO JORGE ISLAND
seem to be related wilh lhe nulrienls slalus. Carex spp. , Eleocharis multicaulis and endemic Agrostis
(graxililaxa and azorica) are the dominanl species in lhe lens.

Dyslrophic ponds and slreams are also presenl (4). Juncus eftusus, Sphagnum spp. , Eleocharis
palustris and Potamogeton polygonifolius are the dominant species on Ihis community.
Dias, Eduardo et aI. Azores Centrallslands Vegetation and Flora

6 - VEGETATION ZONATION OF FAJÃ DOS CUBRES LAGOON- COASTAL WETLANDS BII - Submerge hydrophytes community develops below the lower limit 01 tidal excursion and it's
composed by a broad band 01 Ruppia maritima. On the eastern par! 01 the lagoon, the surface water it
Coas tal wetlands results Irom particular conditions and they have a very restricted distribution in the
cover with a mass 01 Iloating green weed, mainly Enteromorpha intestina/is and species 01
Azorean archipelago. Nowadays the rarity 01 this systems it's even more pronounced in result 01 there
Chaetomorpha. Upon it's decay, the resulting detritus sinks to create the lagoon Iloor 01 thick, black mud
intense human destruction. The lagoons 01 Fajã dos Cubres and Fajã de Santo Cristo in the island 01
that is a impor! leeding ground lor aquatic birds, especially visitors species.
São Jorge are two rare examples.

ln the Nor!h coast 01 the island 01 São Jorge, one can see two lajãs, or coastal platforms. The
coastline that encompasses this lajãs is a geological continuum that also includes clifts and cobble The lagoon it's a impor!ant habitat lor birds including marine birds (e.g. Ca/onectris diomedea, Stema
shores as hydrological basins 01 a lew streams (ribeiras) and unccounted groundwater outflows that dougallii, Stema hirundo and Larus cachinnans at/antis), aquatic visitor birds (e.g. Ardea cinerea, Fulica
reach the sea in this region. This two platforms where lormed by massive landslides resulting Irom the atra, Tringa sp.), aquatic resident birds (e.g. Gallinago gallinago).
ear!hquake 01 1757. The landlorm was shaped by a combination 01 marine and Ireshwater erosion.

The lagoons in both lajãs have several similarities; they are separated Irom the sea by boulder
rampar!s and they received constant Ireshwater input Irom groundwater reservoirs. However the
hydrology and ecology 01 the two lagoons are lundamentally dillerenl. The Santo Cristo lagoon is
essentially marine with the prevailing inlluence Irom the sea, and in opposite the Fajã dos Cubres
lagoon is most estuarine, largely Ireshwater-dominate, and with interesting zonation 01 halophyte and
hygrophyte vegetation communities.

The Fajã dos Cubres lagoon can be divided in two zones with difterent hydrology's; lhe western
section (Zone I). bordered on one side by seaward boulder rampar!. This par! is more inlluenced by the
tidal cycle and reaches higher values 01 salinity (surface salinily 6% - 15%). The eastern section (Zone
II) is separate Irom the western par! by a central islet, so consequentially this par! is little aftected by
tidal cycle, showing lower salinity values (6%-0%) . This section it's very aftected by groundwater
entrance.

The environmental gradient 01 salinity and water depth creates the principal lactors that aftect the
zonation 01 plants in the margins 01 this ~agoon. The vegetation zonation profile (W- E) can be
described as lollow (Figure 2):

A) Halophytic vegetation:

A.I - Halophytic community develops in the boulder rampa r! exposed to high hydro dynamism. It's a
sparse community 01 halophytic species dominated by the micro-camephytic Crithmum maritimum

Ali - Halophytic community in the inner par! 01 the boulder rampar!, exposed to low hydro dynamism.
It's a sparse community 01 halophytic species like; Juncus acutus, Euphorbia azorica, Oaucus carota,
Atrip/ex hasta ta, Lotus comicu/atus. ln the eastern zone this community also occurs but with more
Foenicu/um vu/gare and Lythrum hyssopifo/ia. The lerns Cyrtomium fa/catum and Asp/enium marinum
exploit the crevices between the rocks.

B) Hydrophytes vegetation:

BI - Emergent hydrophytes community develops in the margin 01 the lagoon lorming adense belt 01
Juncus acutus with a low percentage 01 So/idago sempervirens.
Dias, Eduardo et ai. Azores Centrallslands Vegetation and Flora

7 - THE NATURAL GRASSLANDS OF PICO DA ESPERANÇA - ST. JORGE

The natural grasslands 01 Pico da Esperança Reserve is an unique case at Azores. Grass lormation is
not rare at natural areas, but only in small extension and specilic habitats. Sut in this Reserve, ali the
natural areas are cover with endemic grass communities. As lar as our results can take, the explanation
is in the volcanic history. Ali 01 this area was build up in historic times by volcanic deposit 01 basaltic
ashes. They are nutrients rich and give a neutral pH to the soils solution. Sut at this altitude 01 about
1.000m and on the absolute summit 01 the island, the wind is very strong. Any shrub or tree that
attempted grow on this area is vanished any by the winds as the root are only in sand. The trade winds
gave large amount 01 rain to these mountains, and there is a permanent water Ilow. Ali the communities
are organized by these two lactors: Oeschampsia or Festuca wind resistant communities on the more
exposed slopes, Agrostis or Ho/cus on the shelter ones; Grassy lormation on the slopes and bogs
lormation on platforms, associate with ponds in the crater bottom or valley. These unique communities
expor! nutrients in the run-ofl waters and at the discontinuities 01 the slopes (Iike the road) and
exuberant communities 01 large endemic herbs can appear, on the shelter si de 01 the mountain.
Species very rare can have here large populations or even the largest population. This is the case 01
Trifolium trifo/iatum, Chaerophil/um azoricum, Perical/is ma/vifolia, Euphrasia azorica or Scabiosa
nitens.

The Deschampsia fo/iosa communities are typical 01 the mountains summit 01 Azores. Here have his
largest patch, in the very exposed slopes. It is very interesting as the D. fo/iosa cushions can move
during the years, pushed by the wind. The more exposed side 01 the cushion dies every year and new
grows started at opposite side. Setween two cushions 01 Deschampsia, a less tolerant species take
place, like Scabiosa nitens, Leontodon fi/ii, To/pis azorica or P/antago /anceo/ata.

This structure gave a mos ai c aspect to the communities, with the Deschampsia leaves given a look
like ocean waves when under windy conditions.

The Festuca or Ho/cus spp. communities only developed at the shelter slopes, II has also a mosaic
structure, associated with water run-ofl with a net 01 channels. Each cushion 01 grasses can also have
endemic herbs associated, like Rannuncu/us cortusifo/ius, Scabiosa nitens, Ammi trifoliatum or Rumex
azoricus.
Dias, Eduardo et ai . Azores Centrallslands Vegetation and Flora

FIELD TRIP INTREPERTATION SHEET

3 TERCEIRA ISLAND
Dias, Eduardo et aI.
Azores Centrallslands Vegetation and Flora
8 - PEATLAND COMPLEX OF CRIAÇÕES

The ecological conditions in Criações (Terceira island), are lavourable to the development 01 wet
vegetation complexes, especially mire vegetation or other communities directly dependent 01 these.

The soils 01 these place are andosols with placic (delined by an accumulation 01 iron, responsible lor
limiting water drainage), with high amount 01 organic matter, developed Irom volcanic pyroclastic
material, under a wet temperate Atlantic climate. Criações landscape, correspond to a mire complex,
where ali peatland units are connected by a hydrological net. Several types 01 peatland can be identilied
in the area, the bogs are surrounded by a narrow strip 01 mesotrophic lens and the bogs itsell by mixed
deciduous lorest. Tamujo peatbog (protected by Habitat Directive with the designation 01 transition
peatbog) is the largest pure sphagnum lormation and is the result 01 an inlilling sequence Irom open
water to an acid peat ecosystem. Blanket bogs are also common in the area. The most important
peatland type in terms 01 area and biodiversity is lorested lormations. ln lact, generally speaking, local
mires are signilicant rich in Iloristic diversity and have a complex vegetation structure.

The Pico X mire complex contains plant communities that are typical lor Azorean peatland. The
mountain basin, due to the high entrances 01 water (precipitation and log) and its surface retention, is
dominated by Sphagnum communities lorming extent areas 01 blanket bogs. ln the hills due to more
extreme ecological conditions (wind and water surface movement) the predominant lormation is
forested peatland dominated by Juniperus brevifolia, in the middle hill water quantity is smaller, allowing
some organic matter decay and nutrients became available to vascular plants, forming richer and
denser humid lorests with Laurus azorica, lIex perado spp. azorica and Juniperus brevifolia. ln the basin
01 Pico X complex predominate Flat bogs and lew small island 01 fens.

FiaI bogs are generally characterized by a zonation vegetation gradient, which varies Irom the margin
to the interior 01 the bog. Edges are hummock communilies dominated, where nutrients dragged Irom
the lill allow the development 01 communities 01 Polytrichum commune, Calluna vulgaris, or, less
Irequent Erica azorica and Juniperus brevifolia. Herbaceous communities like Juncus eftusus occupy
the intermediate zone. The centre 01 the bog is Ilal and lew scattered species occur like Eleocharis
multicaulis and Prunella vulgaris. Tamujo peatbog, is a Ilotation moss carpet above a lake, where, due
to the stage 01 succession, dimension and an inlilling origin has an atypical Ilat structure prevailing.
These Sphagnum communities are extremely dependenl 01 waler "collector's" species existence in the
highest parts (inside cloud lorest) in Azores the most ejficient are Erica azorica e Juniperus brevifolia.

Fens are richer in nutrients and less acid than bogs, so shrubs and grasses are plentilul (in density
and diversity), and mainly consisl 01 Calluna vulgaris, Erica azorica, Juncus eftusus, Juncus bulbosus,
Eleocharis multicaulis. The bog moss layer consists mainly 01 Sphagnum palustre, S. centrale, S.
subnitens, S. capillifolium, Leucobryum glaucum and Campylopus cetaceus.

This complex, mostly state property owned, plays an imporlant role in the Terceira island landscape
including waler quality bUllering, water discharge and recharge, nutrient and sediment retention and is
habitat lor a wide variely 01 rare and prolected plant like 8ellis azorica and Trichomanes speciosum
among others.
Dias, Eduardo et ai.
Azores Centrallslands Vegetation and Flora
9 - MICRO STRUCTURE ANO ZONATION OF A HUMMOCKI HOLLOW / LAWN/ / POOL GRADIENT
lN A SPHAGNUMPEATBOG

Sphagnum peatbog surface isn't completely Ilat, it possesses a typical microstructure characterized by
a wavy reliel 01 variable compass, Irom some centimetres to some ~eters. The surface presents a
mosaic 01 pools, hollows (or lawns) and hummocks in various stages 01 inlill, degeneration or building
up. Each 01 these microstructure types are associated to distinct ecological conditions, witch are
rellected in an evident vegetation zonation. ln lact, each microlorm behaves as natural vegetation
entities with specilic mosaics 01 vegetation patches and their own micro successions. ln the complex
micro topography 01 bogs, associations are represented only as Iragments dispersed on microlorms 01
the same type.

The raised areas 01 the peatland are called hummocks, and there's above water table. These
structures are higher than lawns and hollows by about 40 to 80 cm and thus they have drier conditions.
Sphagnum species colonizing plateaus and hummocks grow in dense colonies that allow eflicient water
retention and water supply. The most common species in hummocks are Sphagnum palustre,
Sphagnum centrale and Sphagnum papil/osum. Drier conditions lound on hummocks lavour the
presence 01 shrubs and trees as well as other mosses such as Polytrichum commune,
Pseudoscioropodium purum, Thuidium tamariscinum and lichens species (Iike Cladonia portentosa) .
However, large plateaus can lorm wide-open are as 01 trees and shrubs. Under low water table
conditions, vascular plants can lorm adense cover with a sparse moss layer. ln Azores the most
Irequent woody species in hummocks are Cal/una vulgaris; Juniperus brevifolia and Oaboecia azorica,
associated with herbaceous like Luzula purpureo-splendens, Blechnum spicant and Oryopteris spp ..

The peatbog depressions, where the water levei is ciose or in the surface, are called lawns or hollows
depending on the area they cover. Lawns cover large surfaces, while hollows are small depressions.
Plant communities 01 lawns and hollows are less diverse and more scattered, dominated by Sphagnum
mosses and herbaceous Irom witch we can point out Sphagnum auricula tum, Sphagnum subnitens,
Juncus effusus, Eleocharis multicaulis and Hydrocotyle vulgaris.

Frequently, in the margins 01 these systems, it might be lormed some dystrophic pools, that may
become, in situations 01 some slope, permanent water lines. ln pools, water table is above surface, is
nutrient rich, but also very acido These microstructure supports sparse and low plant biodiversity,
species like Sphagnum auricula tum, Sphagnum cuspida tum, Juncus bulbosus, Scirpus fluitans, among
others are the most Irequent.
 r
Dias, Eduardo et ai.
Azores Central lslands Vegetation and Flora
10 - ECOLOGY ANO VEGETATION ZONATION OF A MOUNTAIN WATER STREAM DOMINETED
BY "LAURIFÓLlA" FOREST merge~ with compact communities or grasses as: Festuca juba ta and Ho/cus rigidus. Se/aginel/a
krausslana has a scaUered dlstnbutlon ln the whole area, lorming a continuous rug Irom the top to the
The hydrological net 01 mountain areas 01 most 01 Azorean islands, is characterized with high base 01 the slope, Indlcatmg the presence 01 high wetness.
structural complexity, high Iloristic and biological diversity, lorming ecological corridors 01 larger
structural wealth, comparing with its marginoWater line vegetation , although it seems a mere floristic Along the in.termediate area 01 .the slopes, small stains appear with the small herbaceous species:
component, it constitutes an essential system lor the Iluvial ecosystems, lomenting biodiversity and Leontodon ftlll, e B/echnum splcant. Near the slope basin there is Sibthorpia europaea Luzu/
purpureo-sp/endens and Cu/cita macrocarpa, which grows in rocky blooming covered with a'line ~
biological productivity, constituting alimentary maUer lor the aquatic systems, keeping sediments 01 the
hydrological erosion , storing nutritious Irom lixiviation (working as biological lilter 01 nutrients and 01 layer. The moss stratum is domin.ated by small discontinues carpets 01 Mnium spp, Scapania spp. :~~
several pollutant substances), mainly, when the adjacent terrestrial systems are responsible lor high the IIverwort Conocepha/um Contcum. Thls type 01 slope lormation occurs at altitude (above 600 m)
where vegetatlon has no (or very liUle) human disturbance.
inputs, as it is verilied in medium/high altitude landscape, where it is high covering with pastoral
systems. The water supply lor these riversides, is due practically to the precipitation, thick log and 2. With Rubus spp. - it stands out 01 the previous vegelable unit, by the signilicant covering 01 Rubus
peatlands, these last ones , play a part 01 extreme importance thoroughly regulation 01 the flows, which spp., resultlng ln a I/onstlc Impovenshment, given the aggressiveness 01 the invasion, lorming a
is described, by Mendes (1998) . The geological structure, delining topographical relief's, litologic mlcrohabltat lavourab~e to others p/agues and becoming too inhospitab/e lor auloch thonous species.
discontinuities and other accidents, inlluences water lines, in it's profile and in the losses or flow The emergence 01 thls type 01 nverslde s/ope lormation is associated to disturbances in the natural
earnings. On the other hand, most 01 the drainage lines are situated , prelerentially, along geologic vegetatlon at lower altitudes (especially between 500-600 m). The most evident disturbances are'
Iracture or in discontinuity areas. The combination 01 these lactors with the plateau situation, structures deposlts 01 garbage, erosion 01 the slopes and passage 01 roads and walking trai/s. .
adense hydrological net, but 01 reduced extensions and in very small basins. These lormations are
associated to elevated precipitation recharges, including the occult, which it receives. . Wat.er bed - n~t ali water tine courses profile possess the community characteristics 01 the base 01
nverslde, the pena/pme Heteroc/adion - Cardamine lormation. The development 01 those riversides
Several studies that took place in GEVA (Applied Vegetal Ecology Study Group) demonstrate the base community's, is due to the geomorph%gy, condition lor the existence 01 small is/ands, with small,
existence 01 several ecological/lloristic types 01 water lines in Azorean islands. Their distinction is based but conllnuous I/owlng, where It deve/ops a bryophyte communities 01 Heteroc/adion hutchinsiae and
in several environmental elements 01 the hydrological nêt like water regime and vegetation zonation. herbace~us vegetation with the domain of Cardamine caldeirarum, where jointly, appears the species
Here lollows the vegetation description 01 a Mountain riverside dominated by a laurilolia Forest, where 3 Slbthorpla europaea, Carex vu/cani, Lysimachia azorica, Sanicula azorica and, rarely, the Bel/is azorica.
distinct zones may be distinguished.
The b~ophyte community 01 the basin 01 the water tine is occupied by: Fissidens spp., Mnium spp,
(1) Gallery 01 natural vegetation: margin 01 the water line, with a variable reliel, this can be more or Scapanta nemorosa, Rlcardla spp. and Conocephalum conicum.
less accentuated, depending on the combination 01 the erosive ellort 01 the running water, 01 the
consistence 01 the materiais and 01 the age 01 the structure. The gallery ends below, near the lotic zone. ln the most torrenlial regime, this community disappears, because emerging substratum became
scaUered or even wlthout any vegetation.
This gallery presents a dominance 01 the lollowing species: lIex perado ssp. azoríca, Laurus azorica
Vaccinium cy/indraceum and Hypericum fo/iosum. The structure 01 this gallery is very complex, and is
possible to distinguish an emerging arboreal stratum, that allows the interception 01 thick log, creating
ideal conditions lor the development 01 a sub-Iorest, where other species seUle down; a high incipient
herbaceous stratum, where it can be lound: Dryopteris azorica, B/echnum spicant, Woodwardia
radicans, Cu/cita macrocarpa; a grassy herbaceous stratum, with: Ho/cus rigidus, Deschampsia fo/iosa;
and a low herbaceous stratum 01: Lysimachia azorica, Cardamine caldeirarum and Centaurium
scilloides. Besides these species others like the creepers Smilax diva rica ta and Rubia agostinhoi occur.
Rich lormation on epiphyte species Irom witch we detach Elaphoglossum semicylindricum and
Hymenophyl/um tunbrigense.

Slope (Taludes) - There are 2 types 01 slopes:

1. With a liUle evident vertical stratum , 01 Woodwardia radicans, that appears in this area lorming great
continuous stains, with larger predominance in lhe lop 01 lhe slope and inlermediale area. This lern
 Azores Centrallslands Vegetation and Flora
Dias, Eduardo et aI.
11 - VEGETATION ZONATION OF AN OLlGOTROPHIC LAKE (LAGOA DO NEGRO)

Lagoa do Negro is located at 540 m 01 altitude in the central plateau 01 Terceira island between Sta.
Bárbara Mountain and Pico Alto pick. It's a small lake (protected by Annex B-I 01 Habitats Directive
Decreto-Lei nº 49/2005) with 6 m2, low depth (1.8 m maximum) rich in load sediments, witch has
contributed to inlilling observed in the last decades, and has an accentuated seasonal variation 01 1 m.
It hasn't any direct Ilowing, being maintained by direct rain, water superficial movement Iram its hillsides
and water Irom the surrounding sphagnum peatbog. The lake's water is extremely poor in nutrients and
slightly acido This environmental condition aflects lhe density and the type 01 species 01 the lake. It's an
insular smalllake with a very limited and particular number 01 hygrophyte and hydrophytes species.

ln generic terms, vegetation distribution is similar to classic standards 01 lake vegetation zonation,
being possible to distinguish three areas: margins, benthic and unoccupied area.

Margins communities constitute the transition area Irom the margins to the lake, and Ihese
communities are at leasl once a year submerged. The lake margin has \WO morphological types lo witch
correspond to diflerenl vegetal settlements: (1) 50ft downhill margin, with Hydrocotyle vulgaris,
Polytrichum commune, Juncus effusus and Sphagnum palustre; (2) slope margin dominaled by
mosses, specially Polytrichum commune, but with vascular species like Osmunda regalis, Selaginella
kraussiana, Prunella vulgaris and Plantago coronopus.
Bentonic vegetation is very poor in diversity, only Potamogeton polygonifolius (introduced specie),
Litorella uniflora and Isoetes azorica occurs in this area. This is endemic and exlremely rare specie that
is protected by Annex B-II 01 HabitaIs Directive. II appears only in a narrow strip corresponding lo the
minimum levei reached by water, and restricted lo areas where sediments are loose.

A Ihird idenlilied zone, in the middle deeper part 01 lhe lake, is mentioned as unoccupied 01 vegelal
species due to the lact 01 sediments are loose and has a strong undulation caused by winter strong
winds.

These kinds 01 wetland lormation are also exlremely important for bird species lor nesting, breeding,
shelter, leeding as well lor crucial resting periods lor accidental aquatic birds, like Ardea cinerea.
p

Dias, Eduardo et aI. Azores Centrallslands Vegetation and Flora

12 - VEGETATION ZONATION OF SERRA DE SANTA BÁRBARA Campy/opus spp.). Beyond this limit we enter in the treeline (sensu stricto) where tree growth ceases
entlrely. II glves place to Deschampsia grassland, dominated by Deschampsia fo/iosa with cushion
The mountain Ilora 01 Azores consists 01 a relatively low number 01 vascular plants species, but ranks
growing habit that creates shelter microhabitats where other herbaceous species grow, like Potenti/la
among the regions with highest concentration 01 endemics lormations 01 Europe. ln the low altitudes
azonca, Centaurium scil/oides and Ho/cus rigidus. The hydrological net is very dense in the slopes. The
due to man impacts and the dominance 01 introduced species, most 01 natural vegetation communities
streams valleys are covered with gallery lorest that normally derive Irom structural evolution 01 the
were destroyed. Contrarily, mountain zones still maintain important natural communities. The "Serra de lormations on the streamside.
Santa Bárbara" is one 01 those examples, where it can be lound, nowadays, one 01 the most important
natural area 01 Europe, with a complex 01 lunctional natural habitats.

The "Serra de Santa Bárbara" (1021 m) is the youngest volcano lorm 01 Terceira Island and lies at the
NW end 01 the island. This active volcano has a truncated prolile, diversilied by trachyte domes and
coulés, both inside 01 the caldera and on the slopes 01 the "Serra". ln the past this area was used as
"Baldio", were people release their cattle and collect wood Ireely. Presently, part 01 the "Serra" was
converted to intensive pastures, an in the higher zones translormed in a natural reserve.

The "Serra de Santa Bárbara" slopes are covered with a continuum 01 vegetation lormations that
varies in structure and in floristic composition, rellecting the great variety 01 habitats, each one showing
dilferent combinations 01 environmental lactors. The most important environmental lactors are the wind
exposure, the excessive precipitation cloud, humidity and continuous ground wetness. The geology and
geomorphology are also important lactors, creating high number 01 microhabitats.

On the North lower altitude natural areas 01 "Serra Santa Bárbara", sheller places are covered by a
well developed Juniperus-/lex forest. This lorest is dominated by pense canopies 01 Juniperus breviflolia
and higher canopies 01 /lex pera do ssp. azorica. The structure is well dilferentiated with live layers
(emergent, canopy, canopy, scrub, high herbaceous, low herbaceous, bryophyte and epiphyte). The wet
arborescent scrubs lollow the lorest in the vegetation zonation prolile, and dominate most 01 the slope.
Due to the limiting stress lactors, this lormations result Iram lorest and wood dwarfness or structural and
floristic evolution 01 blanket lorest bogs. These lormations aren't unilorm and change depending on the
exposure. ln the North Slope they are richer in species showing "Iauroide" tendencies. They are
dominated by Juniperus brevifo/ia, but other species like /lex perado spp. azorica, Laurus azorica and
Vaccinium cy/indraceum occur with important cover values. The herbaceous layer develops at the same
levei 01 the scrub layer, due to the internal micro topography, and permits the occurrence 01 species
with dilferent ecological tendencies (e.g. Cu/cita macrocarpa, Dryopteris aemu/a, To/pis azorica).

The Juniperus lorest peatland precede the wet arborescent scrubs in the higher parts 01 the "Serra".
This vegetation type is arare and endemic peatbog with sparse canopies 01 Juniperus brevifo/ia and
important cover 01 Sphagnum sp.

The increasing climate severity with the increasing 01 altitude marks the timberline limit, beyond this
line the trees grow in small patches or as scattered individuais. They are dwarfed, semi-prostrate and
exhibit a krummho/z growth habit, lorming the helophytic mountain scrub. This vegetation lormation has
a complex horizontal structure, constitute by three types 01 patches ; shrub patches (dominate by
Cal/una vu/garis and Juniperus brevifo/ia); herbaceous patches (dominate by Deschampsia fo/iosa,
Festuca juba ta or E/eocharis multicau/is) and bryophyte patches (dominated by Sphagnum spp. and
Dias, Eduardo el aI. Azores Cenlrallslands Vegelalion and Flora

13 - JUNIPERUS FORESTED PEATLAND lN "SERRA DE STA BÁRBARA"

Sta Bárbara Mountain is integrally included in Terceira's Island Nalura 2000 area. It's extremely rich in
rare habilats and species, represenling one 01 lhe mosl important and well preserved biologic hol spol
nol only 01 Azores but also 01 Europe.

The ecological conditions 01 the highest mounlain 01 the island (enormous quantities 01 water
entrances and continuous placic horizon), is lavourable lo lhe development 01 wet vegetation
complexes, especially mires vegetation. These habilats occupy almost ali mounlain surfaces, but in lhe
hillsides, predominanl habitais are Juniperus brevifolia loresled pealland. This loresled pealland is one
01 lhe mosl inleresting communities 01 Azores, being, logether with Iheir associated ones - blanket bog,
on one side, and Juniperus loresl, lor the olher - lormations 01great ecological interest and endemics 01
Azores .

The sinlaxonomic posilion 01 Ihis lormalion can be seen as deriving 01 Blankel Bog wilch has evolved
in Iheir malurily increasing organic matter soil deplh, accentuating anoxic conditions, and soil f100d etc.
ar, lor olher hand, ii can be lhe result 01 an extreme ombrotrophic situation 01 Juniperus loresls
(Iormation Ihis belongs lo lhe Forests syslem 01 the archipelago). The greal singularily 01 Ihis lormation
is lhe lacl Ihat lhe dominant arborescenl specie is lhe endemic Juniperus brevifolia, uncommon as
dominant 01 these lormalions.

This peatland has shrub/tree coverage higher Ihan 50%. II extremely dense, peal will be lormed
entirely wilh lorest remains, because Sphagnum species are shade inlolerant. ln this dense lormalion,
inlerior vegelation layers are occupied with species like Oryopteris affinis, and Luzula purpureo-
splendens, . ln more open lormation Sphagnum carpet may reach an 'O ccupation levei 0180%, and olher
species like Juncus effusus, Cal/una vulgaris and I/ex perado ssp. azorica occur, even iI in a scattered
distribution.

ln some cases is possible lo walch a space competilion between Sphagnum and Juniperus in wilching
the moss carpet covers partially the Juniperus plant. This is a very interesting phenomenon because,
besides demonstrating an advanced evolulionary state 01 lhe peatland (witch may have several melers
deplh 01 peal), ii shows a parallel growing 01 both 01 these species. Juniperus is buried in lhe
Sphagnum peat bul It's roots are in the mineral subtracl 01 geological origin, some meters below, Irom
where II removes nutrienls.

172