access to collections. Dr Emilio Laguna (General- itat Valenciana, Spain) and Dr Daniel Guillot. (Jardín Botánico, Universidad de Valencia,. Spain), are thanked ...
Bradleya 33/2015 pages 58–81
Alien succulents naturalised and cultivated on the central west coast of Portugal Vasco Silva1, Estrela Figueiredo2,3 & Gideon F. Smith2,3 1. Centre for Applied Ecology Prof. Baeta Neves, Instituto Superior de Agronomia, Universidade de Lisboa, Tapada da Ajuda, 1349-017 Lisbon, Portugal 2. Department of Botany, P.O. Box 77000, Nelson Mandela Metropolitan University, Port Elizabeth, 6031 South Africa 3. Centre for Functional Ecology, Departamento de Ciências da Vida, Universidade de Coimbra, 3001-455 Coimbra, Portugal Summary: The mild climate of the central west coast of Portugal has made it predisposed to exotic plants becoming established and eventually invasive in the natural macchia vegetation. Here we record the occurrence of 34 succulents, representing eight families, as forming part of the alien flora of the region. Twenty of these are naturalised and 14 are casuals. Five taxa of succulent plants that are widely cultivated are also catalogued, given the propensity of these plants to become pests. Most of these exotic succulents are illustrated to ease identification.
Zusammenfassung: Das milde Klima der mittleren Westküste von Portugal führt dazu, dass sich hier exotische Pflanzen etablieren und mit der Zeit invasiv in die natürliche Macchia-Vegetation ausbreiten. Hier berichten wir über das Vorkommen von 34 Sukkulenten aus 8 Familien als Teil der Neophytenflora der Region. 20 dieser Arten sind naturalisiert, während 14 als gelegentliche Exemplare auftreten. Fünf weitere, weit verbreitet kultivierte Sukkulententaxa werden ebenfalls katalogisiert wegen der Leichtigkeit, mit der sie zu Neophyten werden. Die meisten dieser exotischen Sukkulenten werden abgebildet, um ihre Identifikation zu erleichtern.
Introduction Portugal has a coastline of about 1000km that is lapped by the Mediterranean Sea in the south and the Atlantic Ocean in the west. The climate of most of Portugal is decidedly Mediterranean with warm to hot summers and comparatively mild, wet winters. Over much of the country, rainfall, mostly recorded in the winter months, is high (Ribeiro, 1963). Given its mild
58
and pleasant climate, Portugal has over the years attracted garden-lovers who have brought vast numbers of exotic plants to the country (Aguiar, 2012; Attlee, 2007; Castel-Branco, 1999, 2014; Carita & Cardoso, 1990; Caixinhas, 2002; Liberato & Caixinhas, 2006; Segal, 1999). Several of these species have escaped from cultivation and subsequently became part of the naturalised alien flora of Portugal (see for example Almeida & Freitas, 2006, 2012; Smith & Figueiredo, 2007, 2009). Lisbon, the capital of Portugal, is situated on the mouth of the Tagus River more or less in the middle of country’s Atlantic coastline. Two coastal-resort towns, Cascais and Estoril, which face southward at the mouth of the Tagus, are situated just northwest of Lisbon, about 25km from the capital, and close to continental Europe’s most westerly point, Cabo da Roca (Ribeiro, 1940) (Figure 1). This area is a popular holiday destination because of the pleasant climate and has been referred to as the ‘French Riviera’ of Portugal (Baedeker, 1911). In addition, it is perfect for colonisation by introduced plants that have escaped from the gardens surrounding the magnificent houses and villas that dot the area (Goeze, 1876; Vasconcellos & Franco, 1958). These towns with their Bohemian flair are surrounded by verdant green metropolitan protected open spaces that, especially in the summer months, are frequented by ecotourists, cyclists, horse-riders, and holidaymakers.
Alien succulents in central coastal Portugal Central coastal Portugal is home to very few indigenous succulents, mostly in the family Crassulaceae, of which 37 species, including 15 natu-
Bradleya 33/2015
Figure 1. Map of the study area near Lisbon, Portugal. ralised ones, have been recorded in Portugal (Smith & Figueiredo, 2010, 2013; Smith et al., 2015). Of the indigenous succulents, Petrosedum sediforme (Jacq.) Grulich is one of the most common and widespread (Figure 2). Several exotic succulents have, however, become established in the area (Silva, 1989; Vasconcellos, 1974). Here we record the occurrence of 34 succulents representing eight families as forming part of the alien flora of central coastal Portugal. The list is based on personal observations and fieldwork, as well as a literature survey (Aedo et al., 2013; Almeida & Freitas, 2006; Almeida, 2012; Coutinho, 1939; Franco, 1971, 1984; Franco & Afonso, 1994; Silva, 1989; Silva, in press; Smith & Figueiredo, 2013; Smith et al., 2015). Five succulent taxa that are widely cultivated are also catalogued, given the predisposition of these plants to invade natural vegetation. The nomenclature of alien succulents is often confused and differently applied in different parts of the world, with various names used in hard-copy and web-based literature for the same plant (Pyšek et al., 2013). For example, Agave
Bradleya 33/2015
salmiana Otto ex Salm-Dyck has been persistently called Agave atrovirens Karw. ex SalmDyck over the years as well as in recent literature (e.g. Aedo et al., 2013), even though the correction has been made in the literature for the region (Smith & Figueiredo, 2007). The terminology for alien plants used in this paper follows Pyšek et al. (2004), where naturalised plants are defined as “alien plants that sustain self-replacing populations for at least 10 years without direct intervention by people (or in spite of human intervention) by recruitment from seed or ramets (tillers, tubers, bulbs, fragments, etc.) capable of independent growth”, casuals as “alien plants that may flourish and even reproduce occasionally outside cultivation in an area, but that eventually die out because they do not form self-replacing populations and rely on repeated introductions for their persistence”, and invasives as “a subset of naturalized plants that produce reproductive offspring, often in very large numbers, at considerable distances from the parent plants, and thus have the potential to spread over a large area”.
59
Figure 2. Petrosedum sediforme recorded from Cabo Raso, Cresmina Beach, near Lisbon. 28 September 2014. Photograph: Gideon F. Smith.
Figure 4a. Green-leaved form of Agave americana (subsp. americana) var. marginata. Photograph: Gideon F. Smith.
Figure 4b.Grey-leaved form of Agave americana (subsp. americana) var. marginata. Photograph: Gideon F. Smith.
60
Figure 3. Agave americana (subsp. americana) var. americana. Photograph: Gideon F. Smith.
Figure 5. Agave salmiana (subsp. salmiana) var. salmiana. Photograph: Vasco Silva.
Bradleya 33/2015
Naturalised succulents observed recorded for central coastal Portugal
and
Agavaceae / Asparagaceae
Agave americana L. (subsp. americana) var. americana (Figure 3) Place recorded in central coastal Portugal: Sintra (Praia Grande), Cascais, Estoril. Also widely naturalised elsewhere in Portugal. Country/region of origin: Mexico, southern USA. Status: Naturalised. References for the area: Coutinho, 1939; Franco & Afonso, 1994; Silva, 1989. Notes: This massive rosulate leaf succulent has uniformly blue-grey leaves and is widely naturalised in mild- and temperate climate regions of the world, including on the Iberian Peninsula. Also see Agave americana L. (subsp. americana) var. marginata, below.
Agave americana L. (subsp. americana) var. marginata Trel. (Figures 4a, b) Place recorded in central coastal Portugal: Cascais. Country/region of origin: Mexico. Status: Naturalised. References for the area: Goeze, 1876. Notes: This variety is a large to massive rosulate leaf succulent with blue-grey or bright green leaves that have yellow margins. Those forms of Agave americana with variegated leaves have a somewhat chequered and confusing nomenclatural history. For example, Bingre et al. (2007) regard a cultivar referred to as “cv. variegata” as applicable to plants with white-margined leaves, while they used “cv. marginata” for those with yellow-margined leaves. The varietal epithet marginata, as established by Trelease (1908), in fact includes plants with both white- and yellow-margined leaves. Although Gentry (1982) appeared to prefer the use of varietal epithets for these leafcolour variations, the use of cultivar epithets may indeed be more appropriate (http://www.plantdelights.com/Photo-Gallery/Agaves-Variegated-Cultivars/Variegated-Agave/). Two green-colour leaf forms of Agave americana L. (subsp. americana) var. marginata are found in central coastal Portugal. Both have yellow leaf margins, but one form has dark green leaves (Figure 4a) with bright yellow margins, while the other one is decidedly greyleaved with lighter yellow margins (Figure 4b). It is noteworthy though that in its native Mexico and in parts of the Iberian Peninsula the name Agave ingens A.Berger var. picta (Salm-Dyck) A.Berger is sometimes used for variegated-leaved plants
Bradleya 33/2015
that bear a very close resemblance to what is widely, but likely incorrectly, known as Agave americana (subsp. americana) var. variegata (González Elizondo et al., 2009; Guillot Ortiz & Van der Meer, 2006). Agave ingens A.Berger [Hort. Mortol.: 12, 360 (1912)] is a nom. illeg. though Berger (1912, 360) gave this name (as a ‘nom. nov.’) to the species that included this green form and the variegated form previously described by Salm-Dyck as A. picta. Thus A. ingens was a superfluous name, as the name A. picta was already published for the species.
Agave salmiana Otto ex Salm-Dyck subsp. salmiana var. salmiana (Figure 5) Locally used synonyms: Agave atrovirens sensu auct. non Karw. ex Salm-Dyck Place recorded in central coastal Portugal: Cabo Raso, Estoril, Parede. Country/region of origin: Mexico. Status: Naturalised. References for the area: Franco & Afonso, 1994. Notes: Agave salmiana Otto ex Salm-Dyck has been called Agave atrovirens Karw. ex Salm-Dyck in the literature (e.g. Aedo et al., 2013; Franco & Afonso, 1994). A correction has been made in the literature for the region (Smith & Figueiredo, 2007).
Furcraea selloi K.Koch (Figures 6a, b) Locally used synonyms: Furcraea foetida sensu auct. non (L.) Haw.; F. gigantea sensu auct. non Vent. Place recorded in central coastal Portugal: Sintra, Cascais, Lisbon (Tapada da Ajuda). Country/region of origin: Unknown. Status: Naturalised. References for the area: Aedo et al., 2013. Notes: The widespread Furcraea foetida is often recorded for the area, but it appears that such records are due to the name being misapplied (e.g. Almeida, 2012; Franco & Afonso, 1994). Unlike F. foetida, F. selloi has regularly spaced marginal teeth along the entire length of a leaf. Asphodelaceae / Xanthorroeaceae
Aloe arborescens Mill. (Figures 7a, b) Place recorded in central coastal Portugal: Nazaré. Cascais, Oeiras, Lisbon. Also in Arrábida (Setúbal). Country/region of origin: Southern Africa. Status: Naturalised. References for the area: Franco & Afonso, 1994; Smith & Figueiredo, 2009. Notes: Widely cultivated and it has escaped in some places (Smith & Figueiredo, 2009).
61
Figure 6a. Furcraea selloi. Photograph: Vasco Silva.
Figure 6b. Rooting bulbils of Furcraea selloi that start spreading into nearby vegetation. Photograph: Vasco Silva.
Figure 7a. Aloe arborescens. Photograph: Gideon F. Smith. Figure 8. Delairea odorata. Photograph: Vasco Silva.
Figure 7b. Inflorescences of Aloe arborescens. Photograph: Gideon F. Smith.
62
Figure 9. Senecio angulatus. Photograph: Vasco Silva.
Bradleya 33/2015
Figure 10a. Anredera cordifolia. Photograph: Vasco Silva.
Figure 10b. Close-up of an inflorescence of Anredera cordifolia. Photograph: Vasco Silva.
Figure 11b. Opuntia ficus-indica spreading into disturbed areas. Photograph: Estrela Figueiredo. Figure 11a. Opuntia ficus-indica. Photograph: Vasco Silva.
Figure 12a. Tradescantia fluminensis. Photograph: Neil R. Crouch.
Bradleya 33/2015
Figure 12b. Close-up of a flower of Tradescantia fluminensis. Photograph: Neil R. Crouch.
63
Asteraceae
Delairea odorata Lem. (Figure 8) Locally used synonyms: Senecio mikanioides Otto ex Walp. Place recorded in central coastal Portugal: Sintra, Cascais, Oeiras, Sesimbra. Country/region of origin: South Africa. Status: Naturalised. References for the area: Morais, 1938; Coutinho, 1939; Franco, 1984; Silva, 1962; Silva, 1989. Notes: The species is known to have been cultivated in Portugal since at least 1883 (Almeida, 2012). Although flattened and ivy-shaped in outline (like those of species of Hedera L., the ‘true’ ivy), the leaves of the species are slightly succulent. The species is included in catalogues of succulents (Smith et al., 1997: 39).
Senecio angulatus L.f. (Figure 9) Place recorded in central coastal Portugal: Sintra, Cascais, Oeiras. Also in Sesimbra and Azeitão (Setúbal). Country/region of origin: South Africa Status: Naturalised. References for the area: Almeida, 2012; Franco, 1984; Silva, 1989. Notes: It is recorded only for central Portugal, where it has escaped from cultivation, becoming naturalised in some areas. Although flattened and marginally roughly lobed, the leaves of the species are indeed thickened, and even the stems are slightly succulent. The species is included in catalogues of succulents (Smith et al., 1997: 40).
Basellaceae
Anredera cordifolia (Ten.) Steenis (Figures 10a, b) Locally used synonyms: Boussingaultia cordifolia Ten. Place recorded in central coastal Portugal: Ericeira, Sintra, Cascais, Oeiras, Sacavém. Also in Coimbra. Country/region of origin: Southern Brazil, Paraguay, northern Argentina. Status: Naturalised. References for the area: Fernandes, 1972d; Franco, 1971; Rainha & Silva, 1972; Silva (in press). Notes: It is known as naturalised in Portugal since 1961 (Rainha & Silva, 1972). This twining vine often produces enlarged aerial tubers, while the rootstock in time becomes irregularly thickened and lumpy, like that of a typical caudiciform succulent. The heart-shaped leaves are somewhat succulent. The species is included in catalogues of
64
succulents (Smith et al., 1997: 41; Walters et al., 2011). Cactaceae
Opuntia ficus-indica (L.) Mill. (Figures 11a, b) Locally used synonyms: Opuntia maxima sensu auct. non Mill. Place recorded in central coastal Portugal: Abano, Costa do Estoril, Lisbon. Also widely naturalised elsewhere in Portugal. Country/region of origin: Probably Mexico. Status: Naturalised. References for the area: Berthet, 1990; Coutinho, 1939. Notes: Cultivated in gardens for its fruit and as an ornamental. It has escaped where discarded plant remains are deposited, and seeds are spread by frugivorous birds. The name Opuntia maxima Mill. has at times been used for the plants we refer to Opuntia ficus-indica (e.g. Berthet, 1990).
Commelinaceae
Tradescantia fluminensis Vell. (Figure 12a, b) Place recorded in central coastal Portugal: Serra de Sintra. Also in Coimbra. Country/region of origin: South America. Status: Invasive (classified in legislation, Dec. Lei no. 565/99) (http://dre.pt/pdf1sdip/1999/12/295A 00/91009115.pdf). References for the area: Almeida, 2012; Franco & Afonso, 2003; Silva, 1956. Notes: Common in cultivation, possibly as a good luck charm, being known as erva-da-fortuna in Portuguese (English: fortune herb). It is now naturalised and invasive in large parts of Portugal (Silva, 1956), and becomes established especially through discarded stems that root easily. Although not many species of the family Commelinaceae are succulent, a few, such as some representatives of Aneilema R.Br. (Smith et al., 1997: 53), Cyanotis D.Don. (Smith & Crouch, 2009: 62), and Tradescantia L. (Walters et al., 2011: 223–231), have slightly fleshy leaves and stems. Crassulaceae
Aeonium arboreum (L.) Webb & Berthel. (Figure 13) Locally used synonyms: Sempervivum arboreum L. Place recorded in central coastal Portugal: Cabo Raso, Estoril, Queijas (Oeiras). Also in Setúbal and Palmela. Country/region of origin: Canary Islands.
Bradleya 33/2015
Figure 13. Aeonium arboreum. Photograph: Gideon F. Smith.
Figure 14a. Growth form of Aichryson laxum. Photograph: Miguel Porto.
Figure 14b. Leafy stalks of Aichryson laxum. Photograph: Miguel Porto.
Figure 14c. Close-up of the shiny leaves of Aichryson laxum Photograph: Miguel Porto.
Figure 15. Crassula multicava. Photograph: Gideon F. Smith.
Figure 16. Aptenia cordifolia. Photograph: Gideon F. Smith.
Bradleya 33/2015
65
Figure 17. Carpobrotus edulis. Photograph: Vasco Silva.
Figure 18b. Close-up of the flowers of Disphyma crassifolium. Photograph: Vasco Silva.
Figure 19b. Close-up of the thin, wiry stems and leaves of Drosanthemum floribundum. Photograph: Vasco Silva.
66
Figure 18a. Disphyma crassifolium. Photograph: Vasco Silva.
Figure 19a. Drosanthemum floribundum. Photograph: Vasco Silva.
Figure 20a. Mesembryanthemum crystallinum. Photograph: Neil R. Crouch.
Bradleya 33/2015
Figure 20b. Close-up of a flower of Mesembryanthemum crystallinum. Photograph: Neil R. Crouch.
Figure 21a. Tetragonia tetragonoides. Photograph: Vasco Silva.
Figure 21b. Close-up of the upper and lower surfaces of the leaves of Tetragonia tetragonoides. Photograph: Gideon F. Smith
Figure 23. Prominent teeth on a leaf margin of Agave salmiana (subsp. salmiana) var. ferox. Photograph: Gideon F. Smith.
Figure 22. Agave attenuata. Photograph: Gideon F. Smith.
Figure 24. Furcraea parmentieri. Photograph: Gideon F. Smith.
Bradleya 33/2015
67
Status: Naturalised. References for the area: Coutinho, 1939; Franco, 1971; Luisier, 1902. Notes: Widely cultivated in gardens from where it has escaped.
Aichryson laxum (Haw.) Bramwell (Figure 14a, b, c) Locally used synonyms: Sempervivum annum C. Sm. ex Link; Aichryson dichotomum (DC.) Webb Place recorded in central coastal Portugal: Colares, Sintra. Country/region of origin: Canary Islands. Status: Naturalised. References for the area: Coutinho, 1939; Silva, 1989. Notes: Occasionally encountered in woods and shaded, rocky places in the Serra de Sintra.
Crassula multicava Lem. (Figure 15) Place recorded in central coastal Portugal: Costa da Guia (Cascais) and Estoril. Country/region of origin: South Africa. Status: Naturalised. References for the area: Silva (in press). Notes: Cultivated in gardens from which it tends to escape. Mesembryanthemaceae / Aizoaceae
Aptenia cordifolia (L.f.) Schwantes (Figure 16) Locally used synonyms: Mesembryanthemum cordifolium L.f. Place recorded in central coastal Portugal: Praia das Maçãs, Cabo Raso, Praia da Parede, Paço de Arcos. Country/region of origin: Southern Africa. Status: Naturalised. References for the area: Coutinho, 1939; Fernandes, 1972b; Silva, 1989. Notes: Widely grown as an ornamental groundcover, it has escaped into natural vegetation.
Carpobrotus edulis (L.) N.E. Br. (Figure 17) Locally used synonyms: Mesembryanthemum edule L. Place recorded in central coastal Portugal: Widespread along the coast. Country/region of origin: South Africa. Status: Invasive (classified in legislation, Dec. Lei no. 565/99) (http://dre.pt/pdf1sdip/1999/12/295A 00/91009115.pdf). References for the area: Coutinho, 1939; Fernandes, 1972b; Franco, 1971; Silva, 1989. Notes: Widely established in many parts of the world on coastal dunes to, often unwisely, combat
68
wind and maritime erosion, the species has invaded natural vegetation. It tends to grow as a near-monoculture where it is firmly established.
Disphyma crassifolium (L.) L. Bolus (Figures 18a, b) Locally used synonyms: Mesembryanthemum crassifolium L. Place recorded in central coastal Portugal: Peniche, Praia Grande, Cabo da Roca, Cascais. Country/region of origin: South Africa, if D. crassifolium subsp. clavellatum (Haw.) Chinnock, from Australia, is considered as a separate species. Status: Naturalised. References for the area: Coutinho, 1939; Bacelar et al., 1987; Fernandes, 1972b; Franco, 1971; Silva, 1989. Notes: Widely grown as an ornamental groundcover, it has escaped into natural vegetation.
Drosanthemum candens (Haw.) Schwantes Place recorded in central coastal Portugal: Cabo da Roca. Country/region of origin: South Africa. Status: Naturalised. References for the area: Fernandes, 1972b; Franco, 1971; Silva, 1989. Notes: Widely grown as a white-flowered ornamental groundcover, it has escaped into natural vegetation and usually forms very thick mats where it is invading along the coast of central Atlantic Portugal.
Drosanthemum floribundum (Haw.) Schwantes (Figures 19a, b) Place recorded in central coastal Portugal: Guincho, Cascais. Country/region of origin: South Africa Status: Naturalised. References for the area: Fernandes, 1972b; Gonçalves, 1990a. Notes: Widely grown as an ornamental groundcover, it has escaped into natural vegetation. The flowers of D. floribundum are never pure white. D. floribundum crosses with D. hispidum (L.) Schwantes, another South African species, and probably D. candens as well (S. Hammer, pers. comm.).
Mesembryanthemum crystallinum L. (Figures 20a, b) Place recorded in central coastal Portugal: Peniche, Ericeira, Cascais. Also in Setúbal and Moita. Country/region of origin: Widespread along coasts in winter rainfall regions (Africa, Australia,
Bradleya 33/2015
Figure 25a. Yucca aloifolia. Photograph: Vasco Silva.
Figure 25b. Close-up of the flowers of Yucca aloifolia. Photograph: Gideon F. Smith.
Figure 26a. Yucca elephantipes. Photograph: Gideon F. Smith. Figure 27. Yucca gloriosa. Photograph: Gideon F. Smith.
Figure 26b. The inflorescences of Yucca elephantipes hardly overtop the leaf rosettes. Photograph: Gideon F. Smith.
Bradleya 33/2015
Figure 28a. Aloe maculata. Photograph: Estrela Figueiredo.
69
Figure 28b. Flat-topped inflorescences of Aloe maculata. Photograph: Gideon F. Smith.
Figure 28c. Aloe maculata. Note the presence of Aloe arborescens and the regular and yellow-variegated leaf forms of Agave americana in the background. Photograph: Vasco Silva.
Figure 29a. Aloiampelos ciliaris. Photograph: Vasco Silva.
Figure 29b. Close-up of an inflorescence of Aloiampelos ciliaris. Photograph: Gideon F. Smith.
Figure 30a. Senecio tamoides. Photograph: Vasco Silva.
70
Figure 30b. Close-up of inflorescences of Senecio tamoides. Photograph: Vasco Silva.
Bradleya 33/2015
Figure 32a. Opuntia leucotricha. Photograph: Vasco Silva. Figure 31. Austrocylindropuntia subulata. Photograph: Estrela Figueiredo.
Figure 32b. Close-up of a pad of Opuntia leucotricha. Photograph: Vasco Silva.
Figure 33b. Close-up of a pad of Opuntia monacantha. Photograph: Vasco Silva.
Bradleya 33/2015
Figure 33a. Opuntia monacantha. Photograph: Vasco Silva.
Figure 34. Opuntia dillenii (or Opuntia stricta var. dillenii, depending on the preferred classification system) showing the purple fruit. Photograph: Vasco Silva.
71
Figure 35. Aeonium arboreum subsp. holochrysum. Photograph: Vasco Silva.
Figure 36. Aeonium haworthii. Photograph: Gideon F. Smith.
Figure 37a. Bluish grey-leaved form of Cotyledon orbiculata. Photograph: Gideon F. Smith.
Figure 37b. Cotyledon orbiculata. Photograph: Vasco Silva.
Figure 38a. Graptopetalum paraguayense. Photograph: Gideon F. Smith.
72
Figure 38b. Graptopetalum paraguayense cultivated as a windowsill plant. Photograph: Vasco Silva.
Bradleya 33/2015
California, Macaronesia, Mediterranean basin). Status: Naturalised. References for the area: Fernandes, 1972b; Gonçalves, 1990b; Luisier, 1902; Vasconcellos, 1974. Notes: Another species of Mesembryanthemum, M. nodiflorum L., occurs in the area (Fernandes, 1972b; Luisier, 1902; Vasconcellos, 1974). It is native to the European Mediterranean area and considered to be indigenous to Portugal (Gonçalves, 1990b; Smith et al., 1998).
Tetragonia tetragonoides (Pall.) Kuntze (Figure 21a, b) Locally used synonyms: Tetragonia expansa Murray Place recorded in central coastal Portugal: From Praia das Maçãs to Adraga (Sintra), Almoínhas Velhas (Cascais). It has been recorded in all coastal provinces. Country/region of origin: It was first collected from New Zealand, and it spread all over the world as a food plant. It is widely and commonly known as New Zealand spinach. Status: Naturalised. References for the area: Coutinho, 1939; Fernandes, 1972c; Silva, 1989. Notes: Commonly cultivated in domestic gardens as a vegetable and frequently sold as plain spinach in fresh produce markets throughout the country. The leaves of the species are slightly succulent. The species is included in catalogues of succulents (Walters et al., 2011).
Notes on Mesembryanthemaceae/ Aizoaceae: One species included in the family, Sesuvium portulacastrum (L.) L., was previously recorded from maritime sands at the mouth of the Tagus River (Trafaria) (Coutinho, 1939; Fernandes, 1972a; Franco, 1971). However, it likely no longer occurs there (Gonçalves, 1990c; Silva, 1971). S. portulacastrum is widely distributed in the tropics and subtropics of both the Old and New Worlds; it has been reported from Mexico, the Antilles and Senegal (Coutinho, 1939). Interestingly, Carpobrotus acinaciformis (L.) L.Bolus and C. chilensis (Mol.) N.E.Br. were not recorded in the study area. Both species have been recorded as established in Spain but only Carpobrotus acinaciformis has been recorded as occurring in Portugal, in the province of Baixo Alentejo (Gonçalves, 1990d). When species of Carpobrotus N.E.Br. are not in flower, they can easily be misidentified as the more commonly encountered C. edulis.
Bradleya 33/2015
Casual and cultivated succulents observed and recorded for central coastal Portugal
Agavaceae / Asparagaceae
Agave attenuata Salm-Dyck (Figure 22) Place recorded in central coastal Portugal: Commonly cultivated in domestic and public gardens. Country/region of origin: Mexico. Status: Cultivated. References for the area: Almeida & Freitas, 2012; Bingre et al., 2007; Chodat, 1909. Notes: Medium-sized to large rosulate leaf succulent; leaves uniformly light green. This is one of very few agaves that form a trunk from which the dry leaves are shed early on. While it is very commonly cultivated in mild-climate parts of the world, it is rarely recorded as having become naturalised. It is naturalised on the island of Madeira (Vieira, 2002).
Agave salmiana subsp. salmiana var. ferox (K.Koch) Gentry (Figure 23) Locally used synonyms: Agave ferox K.Koch Place recorded in central coastal Portugal: Cultivated in gardens. Country/region of origin: Mexico. Status: Cultivated. References for the area: Almeida & Freitas, 2012; Bingre et al., 2007. Notes: This variety is distinguished from the typical variety by its very large leaf marginal spines arranged on prominent teats that give the margins a somewhat wavy appearance.
Furcraea parmentieri (Roezl ex Ortgies) GarcíaMend. (Figure 24) Place recorded in central coastal Portugal: Cultivated in historic parks and gardens, e.g. in Sintra, Cascais, Lisbon. Country/region of origin: Mexico. Status: Cultivated. References for the area: Goeze, 1876; Luckhurst, 2010. Notes: The species is widely cultivated in Mediterranean Europe. The leaves, carried on a thickened trunk, are fountain-like recurved, soft and pliable and the finely saw-toothed leaf margins are devoid of menacing teeth. In general appearance at reproductive maturity, the plant is reminiscent of that of a small palm tree.
Yucca aloifolia L. (Figures 25a, b) Place recorded in central coastal Portugal: Lisbon and surrounds. Also in Azeitão and Pegões
73
Figure 39b. An established stand of plants of Kalanchoe ×houghtonii. Photograph: Vasco Silva.
Figure 39a. Kalanchoe ×houghtonii. Photograph: Estrela Figueiredo.
Figure 40a. Kalanchoe delagoensis. Photograph: Neil R. Crouch. Figure 39c. Close-up of an inflorescence of Kalanchoe ×houghtonii. Photograph: Vasco Silva.
Figure 40b. Close-up of the flowers of Kalanchoe delagoensis. Photograph: Neil R. Crouch.
74
Figure 41a Sempervivum tectorum. Photograph: Gideon F. Smith.
Bradleya 33/2015
(Setúbal), and Coimbra. Country/region of origin: Southeastern USA, Mexico and Caribbean. Status: Casual. Cultivated, escaped. References for the area: Aedo et al., 2013; Almeida, 2012; Coutinho, 1896, 1939; Goeze, 1876. Notes: In 1876, Goeze noted that it was widespread on the coast of Cascais where it is subspontaneous in natural vegetation.
Yucca elephantipes Regel ex Trel. (Figures 26a, b) Place recorded in central coastal Portugal: Widely cultivated in gardens. Country/region of origin: Mexico and Central America. Status: Casual. Cultivated and escaped in places. References for the area: Chodat, 1909; Silva, 2013. Notes: The name Y. gigantea Lem. takes precedence over the more commonly used name Y. elephantipes. This species is widely cultivated as an ornamental, and discarded plant material is often found growing in abandoned lands (Guillot et al., 2013).
Yucca gloriosa L. (Figure 27) Place recorded in central coastal Portugal: Widely cultivated in gardens and parks, e.g. at Cascais, Lisbon. Country/region of origin: North America. Status: Cultivated. References for the area: Goeze, 1876; Silva, 2013; Vasconcellos, 1956. Notes: Yucca gloriosa is very cold- and droughthardy and is widely grown in continental Europe, as far afield as the Czech Republic and Slovakia with their severe winters. It has been suggested that this popular garden plant is a hybrid between Y. aloifolia and Y. filamentosa L. (Rentsch & Leebens-Mack, 2012). Asphodelaceae / Xanthorroeaceae
Aloe maculata All. (Figures 28a, b, c) Locally used synonyms: Aloe saponaria (Aiton) Haw. Place recorded in central coastal Portugal: Cascais, Estoril. Also at Figueira da Foz. Country/region of origin: Southern Africa. Status: Casual. Cultivated, escaped. References for the area: Almeida, 2012. Notes: Aloe maculata has been recorded as naturalised in the Iberian Peninsula, in Spain (Guillot Ortiz et al., 2008). In central coastal Portugal it has escaped from cultivation.
Bradleya 33/2015
Aloiampelos ciliaris (Haw.) Klopper & Gideon F.Sm. (Figures 29a, b) Locally used synonyms: Aloe ciliaris Haw. Place recorded in central coastal Portugal: Vale de Santa Rita (Estoril). Cultivated in historic gardens as an ornamental, e.g. in Cascais, Lisbon. Country/region of origin: South Africa. Status: Cultivated. References for the area: Liberato & Caixinhas, 2006. Notes: We follow the most recently proposed nomenclature for the scrambling aloes, which places this group in its own genus, Aloiampelos Klopper & Gideon F.Sm. (Grace et al., 2013).
Asteraceae
Senecio tamoides DC. (Figures 30a, b) Place recorded in central coastal Portugal: Sintra, Malveira da Serra and Costa da Guia (Cascais), Lisbon (Ajuda). Country/region of origin: Southern Africa Status: Casual. Cultivated, escaped. References for the area: Silva (in press). Notes: It is cultivated in gardens and tends to escape. It is casual in cool microhabitats and on the disturbed edges of watercourses at Serra de Sintra. Although flattened and shaped somewhat like those of the common ivy (species of Hedera L.), the leaves of the species are indeed slightly fleshy. The species is included in catalogues of succulents (Smith et al., 1997: 40). Cactaceae
Austrocylindropuntia subulata (Muehlenpf.) Backeb. (Figure 31) Place recorded in central coastal Portugal: Cascais, Paço de Arcos and Queijas (Oeiras). Also in Azeitão (Setúbal) and Almada. Country/region of origin: Bolivia to Peru. Status: Casual. Cultivated, escaped. References for the area: Bingre et al., 2007; Marchante et al., 2014. Notes: Planted as a hedge, barrier plant, and as an ornamental, from where it has escaped.
Opuntia leucotricha DC. (Figures 32a, b) Place recorded in central coastal Portugal: Cascais (Polima), Lisbon (Tapada da Ajuda). Country/region of origin: Mexico. Status: Casual. Cultivated, escaped. References for the area: Monteiro et al., 2002. Notes: It is cultivated around Observatório Astronómico de Lisboa, possibly as an ornamental.
75
Figure 41b. Sempervivum tectorum growing wedged among rocks sheets. Photograph: Gideon F. Smith. Figure 41d Close-up of flowers of Sempervivum tectorum. Photograph: Estrela Figueiredo.
Crassulaceae
Figure 41c. Inflorescence of Sempervivum tectorum. Photograph: Estrela Figueiredo.
Opuntia monacantha Haw. (Figures 33a, b) Place recorded in central coastal Portugal: Cascais, Lisbon (Tapada da Ajuda). Country/region of origin: South America. Status: Casual. Cultivated, escaped. References for the area: Monteiro et al., 2002; Silva et al. 2015; Vasconcellos, 1956. Notes: Cultivated in gardens from where it has escaped, and new populations become established where discarded plant remains are deposited.
Further notes on Cactaceae: Opuntia stricta (Haw.) Haw. [including Opuntia dillenii (Ker Gawl.) Haw. under var. dillenii (Ker-Gawl.) L.D.Benson] is also cultivated in the area (Monteiro et al., 2002, 2005) (Figure 34). The plant is often recorded in literature under the name Opuntia vulgaris (sensu auct. non Mill.) (e.g. Luisier, 1902; Vasconcellos, 1940).
76
Aeonium arboreum subsp. holochrysum (H.Y.Liu) Bañares (Figure 35) Place recorded in central coastal Portugal: Almoínhas Velhas (Cascais) and Portinho da Arrábida (Setúbal). Country/region of origin: Canary Islands. Status: Casual. Cultivated, escaped. References for the area: Silva et al., 2015. Notes: Plants with glabrous sepals are identified as Aeonium arboreum subsp. holochrysum (H.Y.Liu) Bañares, and have also been recorded as escaped in the area (Silva et al., 2015) and in Spain (Laguna Lumbreras & Mateo Sanz, 2001). According to Bañares Baudet et al. (2008), A. arboreum var. holochrysum H.Y.Liu is a new name (nom. nov.) established by Liu for plants with glabrous sepals. Plants with such sepals had been known under the epithet holochrysum at the species rank. This name was published as new because A. holochrysum Webb & Berth. is a synonym of A. arboreum var. arboreum (the plants described by Webb & Berth. have pubescent sepals) and was not the basionym for A. arboreum var. holochrysum. Therefore, a synonym that can be listed under the currently accepted name A. arboreum subsp. holochrysum (H.Y.Liu) Bañares is A. holochrysum sensu auct. non Webb & Berth.
Aeonium haworthii Webb & Berthel. (Figure 36) Place recorded in central coastal Portugal: Oeiras. Country/region of origin: Macaronesia. Status: Casual. Cultivated, escaped. References for the area: López González, 2001;
Bradleya 33/2015
Silva et al., 2015; Smith & Figueiredo, 2013. Notes: Recorded as a garden escape.
Cotyledon orbiculata L. (Figure 37a, b) Place recorded in central coastal Portugal: Almoínhas Velhas (Cascais), Estoril. Country/region of origin: Southern Africa. Status: Casual. Cultivated, escaped. References for the area: Silva et al., 2015. Notes: Cultivated in gardens. The specimens observed in natural vegetation probably became established from casual and deliberately discarded material.
Graptopetalum paraguayense (N.E.Br.) E.Walther (Figures 38a, b) Place recorded in central coastal Portugal: Paço de Arcos and Oeiras. Country/region of origin: Mexico. Status: Casual. Cultivated, escaped. References for the area: Almeida, 2012. Notes: This small Mexican species grows easily from cuttings and will strike root where material is discarded.
Kalanchoe ×houghtonii D.B.Ward (Figures 39a, b, c) Place recorded in central coastal Portugal: Almoínhas Velhas (Cascais), Oeiras, Lisbon (Tapada da Ajuda). Also cultivated further afield in Portugal, in the central inland, for example in the town of Alcanena. Country/region of origin: Both parents originate from Madagascar. Status: Casual. Cultivated, escaped. References for the area: Almeida, 2012; Monteiro et al., 2002; Smith et al., 2015. Notes: This hybrid between Kalanchoe delagoensis and Kalanchoe daigremontiana Raym.-Hamet & H.Perrier today occurs in many parts of the temperate world, and has been recorded as naturalised from as far afield as New Zealand (http://www.crassulaceae.ch/de/artikel?akID=68& aaID=3&aiID=H&aID=2044) and the Iberian Peninsula (Guillot Ortiz et al., 2014). Several cultivars have been selected from this hybrid (see for example Guillot Ortiz et al., 2014 on Kalanchoe ×houghtonii ‘Garbi’), but we here prefer to simply refer material found in Cascais as belonging to the hybrid proper. While K. ×houghtonii has the general appearance of K. daigremontiana, it tends to be a slightly smaller plant in all respects, and the leaves, while well-spotted with dark purplish brown blotches, in particular are much narrower.
Bradleya 33/2015
Kalanchoe delagoensis Eckl. & Zeyh. (Figure 40a, b) Place recorded in central coastal Portugal: Oeiras. Country/region of origin: Madagascar. Status: Casual. Cultivated, escaped. References for the area: Almeida, 2012. Notes: Kalanchoe delagoensis is one of the most widely naturalised succulents globally, and has become problematic in several countries, including South Africa (Walters et al., 2011). It spreads easily from plantlets carried on the pencil-shaped leaves, as well as through seed.
Sempervivum tectorum L. (Figures 41a, b, c, d) Place recorded in central coastal Portugal: Lisbon and surrounds. Country/region of origin: Continental Europe. Status: Casual. Cultivated, escaped. References for the area: Coutinho, 1939. Notes: It was probably introduced from further north (e.g. France) as a protective charm plant to ward off lightning, a use that has been known since antiquity. It was planted on thatched roofs in the north of the country but as these were replaced with slate roofs and, later, clay tiles, it became rarer (Silva et al., 1976).
Notes on Crassulaceae: Other species recorded for the provinces of Beira Litoral and/or Estremadura, but that were not observed in the study area are: Crassula aquatica (L.) Schönland [Franco, 1971; Fernandes, 1997]; Crassula muscosa L. [Almeida, 2012; Almeida & Freitas, 2006]; Crassula peduncularis (Sm.) Medigen [Almeida, 2012; Fernandes, 1997]; Sedum dendroideum Moçiño & Sessé [Almeida, 2012]; Sedum praealtum A.DC. [Almeida & Freitas, 2006; Tavares, 1960]; and Hylotelephium telephium (L.) H.Ohba [see Smith & Figueiredo, 2013]. Note that Sedum dendroideum and Sedum praealtum are often regarded as names applicable to the same species. However, they are regarded as distinct species by Hovarth (2014), for example. The South African Crassula spathulata Thunb. was reported from Sintra, but a specimen collected by Bento V. Rainha in Monserrate (LISE 80642) was probably of material cultivated in a garden.
Discussion The 39 naturalised, casual or cultivated succulent plant taxa treated in the present study belong to eight families, namely, (1) Agavaceae / Asparagaceae [10 taxa], (2) Asphodelaceae / Xanthorroeaceae [3 species], (3) Asteraceae [3 species], (4) Basellaceae [1 species], (5) Cactaceae
77
[4 species], (6) Commelinaceae [1 species], (7) Crassulaceae [10 taxa], and (8) Mesembryanthemaceae / Aizoaceae [7 species]. The growth forms covered by these taxa span the gamut of succulent plant growth forms, ranging from wiry to heavy soil-hugging creepers, through shrubs, scramblers, single rosette plants, and stem succulents, to climbers. Most of the taxa listed here originated from southern Africa (13), followed by Mexico and the southern USA (11). Four taxa originated from South America, four from Macaronesia, two from Madagascar, and one from each of continental Europe, elsewhere in North America, and New Zealand. The other two taxa are of widespread or undetermined origin. Twenty taxa are listed here as naturalised. Of these, only two species, Carpobrotus edulis and Tradescantia fluminensis, are classified as invasive in Portuguese legislation (Dec. Lei no. 565/99, http://dre.pt/pdf1sdip/1999/12/295A00/91009115.p df). It is noteworthy that, reportedly based on this legislation, Novoa et al. (2014) listed seven species of Cactaceae as invasive in Portugal. However, the legislation clearly includes only one species of cactus (Opuntia ficus-indica), which is not classified as invasive. The list of invasive taxa covered by this legislation has been under review for several years at the Institute for Nature Conservation and Forests (ICNF), which is the national institution responsible for nature conservation and for controlling the introduction of plant species. Since the early 2000s, the creation of environmental awareness, the dissemination of appropriate information and training, and the control and eradication of alien species have been undertaken by local and regional authorities and research institutions, e.g. universities, natural parks, municipalities, and non-governmental organisations. All of these have played an important role in controlling invasive species through applied research projects, and by engaging schools and civil society (http://www.icnf.pt/portal/naturaclas/patrinatur/resource/docs/exot/rel-exo-07-10). However, most of the action is limited in terms of time allocated, because of scarce technical resources, or the expiry of available funding. Critically important follow-up eradication activities are therefore often not conducted. Despite existing legislation, the lack of knowledge about the invasive potential of certain exotic plant species still remains a major drawback in Portugal. The introduction of some plants, including ornamental grasses and succulents, that
78
have proven to be problematic in other parts of the world after their inclusion in gardening and landscaping projects is of particular concern. As a result of their easier and less labour intensive (hence less costly) horticultural maintenance, these plants are becoming more popular in the country and often escape from cultivation, so spreading into vacant areas. Almeida & Freitas (2006, 2012) justifiably refer to the increase in the number of introduced plants in recent times in Portugal. The catalogues we present here prove this statement. It is therefore critically important to have up to date and nomenclaturally robust and accurate inventories of these available to facilitate control and education programmes.
Acknowledgements We thank Dr João D. Almeida (Universidade de Coimbra, Portugal) for information on plants naturalised in Portugal. Dr Jorge Capelo and Isabel Saraiva (LISE Herbarium, Instituto Nacional de Investigação Agrária e Veterinária, Oeiras, Portugal), and M. Dalila Espírito-Santo and A. Paula Paes (LISI Herbarium, Instituto Superior de Agronomia, Lisboa, Portugal) are thanked for access to collections. Dr Emilio Laguna (Generalitat Valenciana, Spain) and Dr Daniel Guillot (Jardín Botánico, Universidad de Valencia, Spain), are thanked for confirming the identity of some species. Alexandre Neto (Direção Geral de Recursos Naturais, Segurança e Serviços Marítimos, Algés, Portugal) is thanked for kindly helping with QGIS. Prof. Neil R. Crouch (South African National Biodiversity Institute, Durban, South Africa) and Dr Miguel Porto (Sociedade Portuguesa de Botânica, Portugal) kindly provided images of some species. Raul Puente Martinez (Desert Botanical Garden, Phoenix, Arizona, USA) is thanked for identifying some of the cactus species. Steven Hammer (San Diego, California, USA) confirmed the identity of some of the representatives of the Mesembryanthemaceae and provided useful comments on the species. An anonymous referee is thanked for useful comments on the paper.
References AEDO, C., HERRERO, A. & QUINTANAR, A. (2013). Agavaceae. In: RICO, E., CRESPO, M.B., QUINTANAR, A., HERRERO, A. & AEDO, C. (EDS), Flora iberica 20: 492–503. Real Jardín Botánico, CSIC, Madrid. AGUIAR, B. (2012). Dicas. Sobrevivem com pouca água. Suculentas ajudam a economizar no jardim. Jardins no. 122 (Ano 9): 40–42.
Bradleya 33/2015
ALMEIDA, J.D. (2012). Flora exótica subespontânea de Portugal Continental (plantas vasculares). Catálogo das plantas vasculares exóticas que ocorrem subespontâneas em Portugal Continental e compilação de informações sobre estas plantas. 5th revised edition. Coimbra [Thesis for Masters in Ecology, Faculty of Sciences and Technology, University of Coimbra, 1999]. ALMEIDA, J.D. & FREITAS, H. (2006). Exotic naturalized flora of continental Portugal. A reassessment. Bot. Complut. 30: 117–130. ALMEIDA, J.D. & FREITAS, H. (2012). Exotic flora of continental Portugal—a new assessment. Bocconea 24: 231–237. ATTLEE, H. (2007). The gardens of Portugal. Frances Lincoln Limited Publishers, London. BACELAR, J.J.A.H., CORREIA, A.I.D., ESCUDEIRO, A.C.S., SILVA, A.R.P. & RODRIGUES, C.M.A. (1987). Novidades da flora Sintrana. Bol. Soc. Brot., Ser. 2, 60: 147–162. BAEDEKER, K. (1911). The Mediterranean seaports and sea routes including Madeira, the Canary Islands, the coast of Morocco, Algeria, and Tunisia: Handbook for travelers. Karl Baedeker Publisher, Leipzig. BAÑARES BAUDET, A., MARRERO GÓMEZ, M.V. & SCHOLZ, S. (2008). Taxonomic and nomenclatural notes on Crassulaceae of the Canary Islands, Spain. Willdenowia 38: 475–489. BERGER, A. (1912). Hortus Mortolensis: enumeratio plantarum in Horto Mortolensi cultarum. Alphabetical catalogue of plants growing in the garden of the late Sir Thomas Hanbury, K.C.V.O., F.L.S. at La Mortola, Ventimiglia, Italy. West, Newman & Co., Hatton Garden, London. BERTHET, P. (1990). Opuntia Mill. In: CASTROVIEJO, S., LAÍNZ, M., LÓPEZ GONZÁLEZ, G., MONTSERRAT, P., MUÑOZ GARMENDIA, F., PAIVA, J. & VILLAR, L. (EDS), Flora iberica 2: 62–70. Real Jardín Botánico, CSIC, Madrid. BINGRE, P., AGUIAR, C., ESPÍRITO-SANTO, D., ARSÉNIO, P. & MONTEIRO-HENRIQUES, T. (coord. cient.) (2007). Guia de Campo – As árvores e os arbustos de Portugal continental. In: SANDE SILVA, J. (coord. ed.), Colecção Árvores e Florestas de Portugal IX. Jornal Público, Fundação Luso-Americana para o Desenvolvimento, Liga para a Protecção da Natureza, Lisboa. CAIXINHAS, M.L. (2002). Cactos e outras plantas suculentas na Estufa Doce de Lisboa. Editorial Verbo, Lisboa. CARITA, H. & CARDOSO, A.H. (1990). Portuguese gardens. Antique Collectors' Club, Woodbridge.
Bradleya 33/2015
CASTEL-BRANCO, C. (ED.) (1999). Botanic Gardens of Ajuda. Jardim Botânico da Ajuda, Lisboa. CASTEL-BRANCO, C. (2014). Jardins de Portugal. Clube do Colecionador dos Correios, CTT Correios de Portugal, Lisboa. CHODAT, R. (1909). Excursions botaniques en Espagne et au Portugal. Imprimerie Eugene Froreisen, Genève. COUTINHO, A.X.P. (1896). Contribuições para o estudo da flora portugueza. Bol. Soc. Brot. 13: 77–78. COUTINHO, A.X.P. (1939). Flora de Portugal, 2nd edn. Irmãos Bertrand, Lda., Lisboa. FERNANDES, R.B. (1972a). Aizoaceae. Anu. Soc. Brot. 38: 123–126. FERNANDES, R.B. (1972b). Mesembryanthemaceae. Anu. Soc. Brot. 38: 127–136. FERNANDES, R.B. (1972c). Tetragoniaceae. Anu. Soc. Brot. 38: 137–138. FERNANDES, R.B. (1972d). Basellaceae. Anu. Soc. Brot. 38: 139–140. FERNANDES, R.B. (1997). Crassula L. In: CASTROVIEJO, S., AEDO, C., LAÍNZ, M., MORALES, R., MUÑOZ GARMENDIA, F., NIETO FELINER, G. & PAIVA, J. (EDS), Flora iberica 5: 99–103. Real Jardín Botánico, CSIC, Madrid. FRANCO, J.A. (1971). Nova Flora de Portugal, vol. I Lycopodiaceae—Umbelliferae. Edição do Autor, Lisboa. FRANCO, J.A. (1984). Nova Flora de Portugal, vol. II Clethraceae—Compositae. Edição do Autor, Lisboa. FRANCO, J.A. & AFONSO, M.L.R. (1994). Nova Flora de Portugal (Continente e Açores), vol. III(I) Alismataceae—Iridaceae. Escolar Editora, Lisboa. FRANCO, J.A. & AFONSO, M.L.R. (2003). Nova Flora de Portugal (Continente e Açores), vol. III(III) Juncaceae—Orchidaceae. Escolar Editora, Lisboa. GENTRY, H.S. (1982). Agaves of continental North America. University of Arizona Press, Tucson. GOEZE, E. (1876). Jardins Notaveis de Portugal— Cascaes. Jornal de Horticultura Pratica VII: 8–13. GONÇALVES, M.L. (1990a). Drosanthemum Schwantes. In: CASTROVIEJO, S., LAÍNZ, M., LÓPEZ GONZÁLEZ, G., MONTSERRAT, P., MUÑOZ GARMENDIA, F., PAIVA, J. & VILLAR, L. (EDS), Flora iberica 2: 90–92. Real Jardín Botánico, CSIC, Madrid. GONÇALVES, M.L. (1990b). Mesembryanthemum L. In: CASTROVIEJO, S., LAÍNZ, M., LÓPEZ GONZÁLEZ, G., MONTSERRAT, P., MUÑOZ GARMENDIA, F., PAIVA, J. & VILLAR, L. (EDS), Flora
79
iberica 2: 79–82. Real Jardín Botánico, CSIC, Madrid. GONÇALVES, M.L. (1990c). Sesuvium L. In: CASTROVIEJO, S., LAÍNZ, M., LÓPEZ GONZÁLEZ, G., MONTSERRAT, P., MUÑOZ GARMENDIA, F., PAIVA, J. & VILLAR, L. (EDS), Flora iberica 2: 74–76. Real Jardín Botánico, CSIC, Madrid. GONÇALVES, M.L. (1990d). Carpobrotus N.E.Br. In: CASTROVIEJO, S., LAÍNZ, M., LÓPEZ GONZÁLEZ, G., MONTSERRAT, P., MUÑOZ GARMENDIA, F., PAIVA, J. & VILLAR, L. (EDS), Flora iberica 2: 83–86. Real Jardín Botánico, CSIC, Madrid. GONZÁLEZ ELIZONDO, M., GALVÁN VILLANUEVA, R., LÓPEZ ENRIQUEZ, I.L., RESÉNDIZ ROJAS, L. & GONZÁLEZ ELIZONDO, M.S. (2009). Agaves— magueyes, lechuguillas y noas del Estado de Durango y sus alrededores. CIIDIR Unidad, Durango, Instituto Politécnico Nacional, Durango, and CONABIO, Mexico, D.F. GRACE, O.M., KLOPPER, R.R., SMITH, G.F., CROUCH, N.R., FIGUEIREDO, E., RØNSTED, N., VAN WYK, A.E. (2013). A revised generic classification for Aloe (Xanthorrhoeaceae subfam. Asphodeloideae). Phytotaxa 76(1): 7–14. http://dx.doi.org/10.11646/phytotaxa.76.1.2 GUILLOT ORTIZ, D. & VAN DER MEER, P. (2006). Claves de las especies del género Agave L. cultivadas como ornamentales en la Península Ibérica e Islas Baleares. Acta Bot. Barc. 50: 441–457. GUILLOT ORTIZ, D., LAGUNA LUMBRERAS, E. & ROSSELLÓ PICORNELL, J.A. (2008). La familia Aloaceae en la flora alóctona valenciana. Bouteloua 6: 1–58. GUILLOT ORTIZ, D., LAGUNA LUMBRERAS, E., LÓPEZPUJOL, J., SÁEZ, L. & PUCHE, C. (2014). Kalanchoe ×houghtonii ‘Garbí’. Bouteloua 19: 99–128. GUILLOT, D., LAGUNA, E., ROSSELLÓ, J.A. & VAN DER MEER, P. (2013). El género Yucca L. en la flora alóctona valenciana. Bouteloua 14: 124– 149. HOVARTH, B. (2014). The plant lover’s guide to sedums. Timber Press, Portland, in association with the Royal Botanic Gardens, Kew, Richmond, Surrey. LAGUNA LUMBRERAS, E. & MATEO SANZ, G. (2001). Observaciones sobre la flora alóctona valenciana. Fl. Montib. 18: 40–44. LIBERATO M.C. & CAIXINHAS M.L. (2006). Cacti and other succulent plants—A vegetal patrimony in some Lisbon garden-museums. Rev. Ciênc. Agrár. 29: 356–364. LÓPEZ GONZÁLEZ, G.A. (2001). Los árboles y arbustos de la Península Ibérica e Islas Baleares.
80
Especies silvestres y las principales cultivadas, Tomo I, II. Ediciones Mundi-Prensa, Madrid. LUCKHURST, G. (2010). Jardim do México [online]. Associação Amigos de Monserrate. Available: http://amigosdemonserrate.com/artigo/jardimdo-mexico-por-gerald-luckhurst [Accessed November 2014]. LUISIER, A. (1902). Apontamentos sobre a flora da região de Setubal. Catalogo das plantas vasculares dos arredores de Setubal e da serra d’Arrabida. Bol. Soc. Brot. 19: 172–274. MARCHANTE, H., MORAIS, M., FREITAS, H. & MARCHANTE, E. (2014). Guia prático para a identificação de Plantas Invasoras em Portugal. Imprensa da Universidade de Coimbra, Coimbra. MONTEIRO, A., CHEIA, V.M., COSTA, J.C., VASCONCELOS, T., LOUSÃ, M., MOREIRA, I. & DINIZ, M.A. (2002). Reserva Natural "D. António Xavier Pereira Coutinho": Sugestão de atribuição de categoria e a problemática das invasoras. In: ESPÍRITO-SANTO, M.D, SOARES, A.L & COSTA, J.C. (EDS), Actas do VII Simpósio da Associação Ibero-Macaronésica de Jardins Botânicos “Jardins Botânicos. Que perspectiva para o futuro?”: 141–151. AIMJB, Lisboa. MONTEIRO, A., CHEIA, V.M., VASCONCELOS, T. & MOREIRA, I. (2005). Management of the invasive species Opuntia stricta in a Botanical Reserve in Portugal. Weed Res. 45: 193–201. MORAIS, A.T. (1938). Breves estudos na flora portuguesa. Anu. Soc. Brot. 4: 28–29. NOVOA, A., LE ROUX, J.J., ROBERTSON, M.P., WILSON, J.R.U. & RICHARDSON, D.M. (2014). Introduced and invasive cactus species: a global review. AoB PLANTS 7: plu078; doi:10.1093/ aobpla/plu078 PYŠEK, P., HULME, P.E., MEYERSON, L.A., SMITH, G.F., BOATWRIGHT, J.S., CROUCH, N.R., FIGUEIREDO, E., FOXCROFT, L.C., JAROŠÍK, V., RICHARDSON, D.M., SUDA, J. & WILSON, J.R.U. (2013). Hitting the right target: taxonomic challenges for, and of, plant invasions. AoB PLANTS 5: plt042; doi:10.1093/aobpla/042 PYŠEK, P., RICHARDSON, D.M., REJMÁNEK, M., WEBSTER, G., WILLIAMSON, M. &KIRSCHNER, J. (2004). Alien plants in checklists and floras: towards better communication between taxonomists and ecologists. Taxon 53: 131–143. RAINHA, B.V. & SILVA, M. (1972). Boussingaultia cordifolia Ten. Agron. Lusit. 34(3): 186. RENTSCH, J.D. & LEEBENS-MACK, J. (2012). Homoploid hybrid origin of Yucca gloriosa: intersectional hybrid speciation in Yucca (Agavoideae, Asparagaceae). Ecol. Evol. 2: 2213–2222.
Bradleya 33/2015
RIBEIRO, O. (1940). Remarques sur la morphologie de la région de Sintra et Cascais. Rev. Géogr. Pyrén. Sud-Ouest 11(3–4): 203–218. RIBEIRO, O. (1963). Portugal, o Mediterrâneo e o Atlântico, 2nd edn. Livraria Sá da Costa, Lisboa. SEGAL, B. (1999). The garden lover’s guide to Spain and Portugal. Princeton Architectural Press, New York. SEQUEIRA, M., ESPÍRITO-SANTO, M.D., AGUIAR, C. CAPELO, J. & HONRADO, J. (COORDS) (2011). Checklist da flora de Portugal (Continental, Açores e Madeira) [online]. Associação Lusitana de Fitossociologia (ALFA). Available: http://bibdigital.rjb.csic.es/PDF/Sequeira_y_al _Checklist_Flora_Portugal_2011.pdf [Accessed November 2014]. SILVA, A.R.P. (1956). Tradescantia fluminensis Vell. Agron. Lusit. 18(1): 19. SILVA, A.R.P. (1971). Les plantes synanthropiques au Portugal continental et aux Açores. Boissiera 19: 297–303. SILVA, A.R.P. (1989). A flora da Serra de Sintra. Catálogo. Portugaliae Acta Biol., Sér. B, Sist. 15: 5–258. SILVA, A.R.P., TELES, A.N. & LOPES, M.H.R. (1976). Sempervivum tectorum L. Agron. Lusit. 37(2): 174–179. SILVA, M. (1962). Senecio mikanioides Otto ex Walp. Agron. Lusit. 24(3): 201. SILVA, V. (2013). Vegetação de Parques e de outros Espaços Verdes Urbanos de Cascais. Internal report. 6 pp. + digital cartography (1:5000). Empresa Municipal de Ambiente de Cascais, E.M., S.A., Cascais. SILVA, V. (in press). Novas localidades na Estremadura para flora exótica estabelecida e outras novidades para Portugal. Silva Lusitana. SILVA, V., LAGUNA, E. & GUILLOT, D. (2015). Algunos datos sobre neófitos en Portugal. Bouteloua 20: 76–96. SMITH, G.F., CHESSELET, P., VAN JAARSVELD, E.J., HARTMANN, H., HAMMER, S., VAN WYK, B-E., BURGOYNE, P., KLAK, C. & KURZWEIL, H. (1998). Mesembs of the world. Illustrated guide to a remarkable succulent plant group. Briza Publications, Pretoria. SMITH, G.F. & CROUCH, N.R. (2009). Guide to succulents of southern Africa. Random House Struik Nature, Cape Town.
Bradleya 33/2015
SMITH, G.F. & FIGUEIREDO, E. (2007). Naturalized species of Agave L. (Agavaceae) on the southeastern coast of Portugal. Haseltonia 13: 52– 60. SMITH, G.F. & FIGUEIREDO, E. (2009). Aloe arborescens Mill. (Asphodelaceae) is spreading in Portugal. Bradleya 27: 165–167. SMITH, G.F. & FIGUEIREDO, E. (2010). Petrosedum sediforme, a beautiful species of Crassulaceae from Portugal. CactusWorld 28: 103–106. SMITH, G.F. & FIGUEIREDO, E. (2013). The family Crassulaceae in continental Portugal. Bradleya 31: 76–88. SMITH, G.F., FIGUEIREDO, E. & SILVA, V. (2015). Kalanchoe ×houghtonii (Crassulaceae) recorded near Lisbon, Portugal. Bouteloua 20: 97–99. SMITH, G.F., VAN JAARSVELD, E.J., ARNOLD, T.H., STEFFENS, F.E., DIXON, R.D. & RETIEF J.A. (EDS) (1997). List of southern African succulent plants. Umdaus Press, Pretoria. TAVARES, C.N. (1960). Perigo da introdução de vegetais exóticos. Protecção da Natureza. Bol. Inform. Liga Prot. Nat., n. sér., 5-6: 13–20. TRELEASE, W. (1908). Variegation in the Agaveae. In: VON LINSBAUER, K. (ED.), WiesnerFestschrift im Auftrage des Festkomitees. Pp. 332–356. C. Konegen, Wien. VASCONCELLOS, J.C. (1940). Anotações do Herbário. Anais Inst. Sup. Agron. (Lisboa) 11: 12. VASCONCELLOS, J.C. (1956). Reserva Botânica Natural “D. António Xavier Pereira Coutinho”. Agros 3: 127–136. VASCONCELLOS, J.C. (1974). Plantas das areias e rochedos litorais (Fanerogâmicas). Secretaria de Estado da Agricultura. Direcção-Geral dos Serviços Florestais e Aquícolas, Lisboa. VASCONCELLOS, J.C. & FRANCO, J.A. (1958). Novas plantas para a flora portuguesa. Bol. Soc. Brot., Ser. 2, 32: 237–251. VIEIRA, R. (2002). Flora da Madeira: plantas vasculares naturalizadas no arquipélago da Madeira. Boletim do Museu Municipal do Funchal (História Natural). Suplemento nº 8. 281 pp. WALTERS, M., FIGUEIREDO, E., CROUCH, N.R., WINTER, P.J.D., SMITH, G.F., ZIMMERMANN, H.G. & MASHOPE, B.K. (2011). Naturalised and invasive succulents of southern Africa. ABC Taxa 11. The Belgian Development Cooperation, Brussels.
81
