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of the madeira and selvagens archipelagos - redmic

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plausível uma origem mais antiga dos endemismos da Madeira.<br />

Posteriormente alguns autores levantaram a hipótese<br />

de uma parte dos endemismos macaronésicos arborescentes<br />

não terem um carácter arcaico reliquial, mas representarem<br />

uma tendência evolutiva comum nas ilhas oceânicas,<br />

com um carácter derivado recente (neo -endémico). Estas<br />

plantas apresentam hábito arborescente caulirrosulado,<br />

arborescente em c<strong>and</strong>elabro ou de planta lenhosa monocárpica<br />

(Böhle et al. 1996), comum em representantes insulares<br />

das Boraginaceae (Echium), Asteraceae (Sonchus),<br />

Campanulaceae (Musschia), Scrophulariaceae (Isoplexis) e<br />

outros como Sideritis, Plantago e Euphorbia. Foi Carlquist<br />

(1974) quem designou esta tendência evolutiva insular por<br />

hábito lenhoso insular.<br />

A origem dos elementos da flora vascular da Madeira,<br />

descrita por Capelo et al. (2007), pode resumir -se do seguinte<br />

modo:<br />

1. paleo -endemismos, de origem subtropical terciária <strong>the</strong>tysiana,<br />

fanerófitos tal como os dos géneros Laurus, Ocotea,<br />

Apollonias, Persea, Clethra, Ilex, Picconia, Heberdenia,<br />

Myrica, Prunus e provavelmente Dracaena e Sideroxylon,<br />

bem como pteridófitos florestais e epifíticos;<br />

2. neo -endemismos, arbustos e plantas caulirrosuladas com<br />

origem continental (ou em outros arquipélagos) do final<br />

do Terciário e início do Quaternário e com forte radiação<br />

adaptativa (e.g. Aeonium, Sonchus, Echium, Sinapidendron,<br />

Euphorbia, Isoplexis, Musschia, Plantago, etc.);<br />

3. flora mediterrânica em que se podem distinguir elementos<br />

paleo -mediterrânicos (xerófitas, esclerófilas e termófilas,<br />

e.g., Euphorbia subsect. Pachycladae, Olea, Maytenus,<br />

Myrtus, Rhamnus, Asparagus, etc.), com origem<br />

subtropical que actualmente se distribui na área do Mediterrâneo,<br />

África tropical xérica/subdesértica, atingindo<br />

o Mar Vermelho; e os elementos neo -mediterrânicos (arbustos,<br />

e.g., Micromeria, Sideritis, Teline, Genista), com<br />

origem holártica continental;<br />

4. flora antrópica (plantas introduzidas pelo homem e naturalizadas<br />

a que correspondem arqueófitos, antes de<br />

1500 AD., e.g., Castanea, Pinus e Vitis; neófitos introduzidos<br />

depois de 1500 AD., e.g., Ageratina, Duchesnea,<br />

Ulex, Cytisus, Erigeron, Hedychium, etc.).<br />

Quanto à origem geográfica dos taxa ancestrais da flora<br />

vascular da Madeira (Panero et al. 1999; Barber et al. 2002;<br />

Molero et al. 2002; Capelo et al. 2004 e referências neles<br />

incluídas), sendo predominantemente mediterrânica, como<br />

os géneros Echium e Euphorbia, cujos endemismos arborescentes<br />

tiveram origem em taxa circum -mediterrânicos<br />

herbáceos, possui também elementos de origem eurossiberiana<br />

(géneros Silene e Saxifraga), norte -americana (géneros<br />

Pericallis e Sedum) e sul -americana (género Bystropogon).<br />

A vegetação potencial do arquipélago da Madeira inclui<br />

diferentes tipos de florestas – Mayteno umbellatae -Oleetum<br />

maderensis [zambujal]; Semele <strong>and</strong>rogynae -Apollonietum<br />

barbujanae [laurissilva do barbusano]; Clethro arboreae-<br />

-Ocoteetum foetentis [laurissilva do til]; Polysticho falcinelli-<br />

-Ericetum arboreae [urzal de altitude]. Costa et al. (2004)<br />

159<br />

more ancient origin <strong>of</strong> Madeira endemics was most plausible.<br />

Later, some authors raised <strong>the</strong> hypo<strong>the</strong>sis that a portion<br />

<strong>of</strong> <strong>the</strong> arborescent endemics <strong>of</strong> Macaronesia were not<br />

relictual, but instead represented a secondarily derived evolutionary<br />

trend (neo -endemics), common in oceanic isl<strong>and</strong>s.<br />

These plants present unusual habits such as caulirosulate<br />

shrub, c<strong>and</strong>elabra shrub or monocarpic rosette tree (Böhle<br />

et al. 1996) that can be found in insular members <strong>of</strong> <strong>the</strong> Boraginaceae<br />

(Echium), Asteraceae (Sonchus), Campanulaceae<br />

(Musschia), Scrophulariaceae (Isoplexis) <strong>and</strong> in o<strong>the</strong>rs<br />

like Sideritis, Plantago <strong>and</strong> Euphorbia. Carlquist (1974) named<br />

this insular evolutionary trend, insular woodiness.<br />

According to Capelo et al. (2007), <strong>the</strong> origin <strong>of</strong> <strong>the</strong> elements<br />

<strong>of</strong> <strong>the</strong> vascular Flora <strong>of</strong> Madeira can be summarised<br />

as following:<br />

1. Palaeo -endemics, phanerophytes (such as those <strong>of</strong> <strong>the</strong><br />

genera Laurus, Ocotea, Apollonias, Persea, Clethra, Ilex,<br />

Picconia, Heberdenia, Myrica, Prunus <strong>and</strong> probably also<br />

Dracaena <strong>and</strong> Sideroxylon), as well as forest -floor <strong>and</strong><br />

epiphytic pteridophytes with a subtropical Tertiary Tethysian<br />

origin;<br />

2. Neo -endemics, shrubs <strong>and</strong> caulirosulate plants with a<br />

continental origin (or from o<strong>the</strong>r <strong>archipelagos</strong>) by <strong>the</strong> end<br />

<strong>of</strong> <strong>the</strong> Tertiary <strong>and</strong> early Quaternary <strong>and</strong> exhibiting a wide<br />

adaptive radiation (e.g. Aeonium, Sonchus, Echium, Sinapidendron,<br />

Euphorbia, Isoplexis, Musschia, Plantago,<br />

etc.);<br />

3. Mediterranean Flora, including Palaeo -Mediterranean<br />

elements (xerophytes, sclerophylous <strong>and</strong> termophylous,<br />

e.g., Euphorbia subsect. Pachycladae, Olea, Maytenus,<br />

Myrtus, Rhamnus, Asparagus, etc.) with a subtropical origin<br />

<strong>and</strong> nowadays distributed throughout <strong>the</strong> Mediterranean<br />

<strong>and</strong> xeric/sub -desertic tropical Africa, reaching <strong>the</strong><br />

Red Sea; <strong>and</strong> <strong>the</strong> neo -mediterranean elements (shrubs,<br />

e.g., Micromeria, Sideritis, Teline, Genista), with a holartic<br />

continental origin;<br />

4. Anthropic Flora, human -introduced plants that became<br />

established. The archaeophytes (e.g. Castanea, Pinus<br />

<strong>and</strong> Vitis) were introduced before 1500 AD, while neophytes<br />

(e.g. Ageratina, Duchesnea, Ulex, Cytisus, Erigeron,<br />

Hedychium, etc.) were introduced after 1500 AD.<br />

The geographic origin <strong>of</strong> <strong>the</strong> ancestral taxa <strong>of</strong> <strong>the</strong> vascular<br />

plants <strong>of</strong> Madeira is predominantly Mediterranean (Panero<br />

et al. 1999; Barber et al. 2002; Molero et al. 2002; Capelo<br />

et al. 2004 <strong>and</strong> references <strong>the</strong>rein), like <strong>the</strong> genera Echium<br />

<strong>and</strong> Euphorbia where endemic woody species arose from<br />

circum -mediterranean herbaceous taxa; but <strong>the</strong>re are also<br />

elements with euro -siberian (genera Silene <strong>and</strong> Saxifraga),<br />

North -American (genera Pericallis <strong>and</strong> Sedum) <strong>and</strong> South-<br />

-American (genus Bystropogon) origins.<br />

The potential vegetation <strong>of</strong> <strong>the</strong> archipelago <strong>of</strong> Madeira includes<br />

different forest types – Mayteno umbellatae -Oleetum<br />

maderensis [Madeira Olea scrub]; Semele <strong>and</strong>rogynae-<br />

-Apollonietum barbujanae [mediterranean ‘barbusano laurisilva’];<br />

Clethro arboreae -Ocoteetum foetentis [temperate til laurisilva];<br />

Polysticho falcinelli -Ericetum arboreae [high altitude

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