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PALMS Luckhurst: Palms of Monserrate Vol. 55(1) 2011 The Palms of GERALD LUCKHURST Landscape Architect Monserrate, Avenida 25 de Abril, 56, Galamares, 2710-246 Sintra Sintra, Portugal Portugal [email protected]

1. Dome of Monserrate seen behind Trachycarpus fortunei and Phoenix canariensis.

The garden of Monserrate in Portugal contains a wealth of fine trees planted mostly in the second half of the nineteenth century including giant Araucarias, Kauri pines, Banyans and Metrosideros. The collection of palms is particularly rich and has great historical significance since the palms at Monserrate were among the first specimens of their kind planted in the open air in Europe. Today there are some seventy or more species of palm growing at Monserrate, twenty-four of them representing historic plantings (Fig. 1).

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Sintra, near Lisbon, Portugal, enjoys one of arches, Roman and Renaissance sculpture, the mildest climates in Europe, comparable Chinese urns and Iznik tiles. The house, built only to the southern-most coasts of Spain and on de Visme’s gothic castle walls, was Italy and some islands of the Mediterranean. decorated in “Moorish style” with an amalgam However, its position at the western-most of Indian and Venetian and Florentine point of continental Europe gives it a wholly Renaissance details – the palace of a Nabob in Atlantic outlook with abundant winter rains the words of one visitor. It contained works of and cool, misty summers. Winter minimum art in every room: paintings, sculpture, arms, tempera-tures on the northern slopes of the books, textiles, rugs, even stuffed birds and Serra de Sintra are warmer than those of tiger-skin rugs. Later he became Sir Francis surrounding areas, with frost-free conditions Cook, Bt. and Visconde de Monserrate, that allow classified for USDA Zone 10b renowned as the greatest art collector of his to be grown. The soils are composed of age, guided by Sir J. C. Robinson, keeper of decayed granite and abundant organic pictures to Queen Victoria, and founder of the material. Victoria & Albert Museum, London. Monserrate is an historic garden founded in Naturally he began to fill the garden with the late 18th century by Gerard de Visme, a exotic and expensive plants. This was the age wealthy English merchant who had made his of great Victorian glasshouses filled with the fortune in trade with Brazil. He built a neo- treasures of uncharted lands – Brazil, , gothic house that quickly fell to ruin during , South Africa, China and Japan. the Napoleonic invasions and ensuing civil Cook’s ambition was to make a garden at war in Portugal (1807–1834). Lord Byron saw Monserrate that would rival the greatest the ruined house and garden in 1809 and botanical collections of Europe. In England he wrote about them in Childe Harold’s Pilgrimage. lived on Richmond Hill, close to the epicenter Such was the effect of this poet’s fame that the of Victorian exploration, the Royal gardens instantly became an obligatory Botanic Gardens at Kew, where he maintained destination visited by all Grand Tourists of the a close relationship with the directors William nineteenth century. and Joseph Hooker. At Monserrate the garden was to contain a greater variety of plants than One such wealthy traveller was the young any other garden in Europe – except Kew! Francis Cook returning from a tour of Greece, Turkey, Egypt, Italy and the Iberian Peninsula. All of this was to be grown in the open air. Arriving in Lisbon he had letters of “Imagine,” said Dr. David Moore (director of introduction to his father’s trading partners Glasnevin botanic garden) in 1867, “imagine and was entertained in Sintra by English a piece of ground a thousand times the size of merchants who had their summer houses the palm house at Kew,” wholly dedicated to there. He met and fell in love with Emily Lucas, the culture of exotic plants. daughter of one of these families and tradition It was David Moore that gave us the first record has it that they discovered the ruins of of some of the successful plantations of palms Monserrate on their honeymoon in August from when the garden was first established. 1841. These were listed as Areca sapida, Chamaerops Cook was the second son of William Cook, martiana, Chamaerops fortunei, Seaforthia one of Britain’s largest traders in all kinds of elegans, Latania borbonica and Phoenix silk, linen, woolen and cotton goods. When his dactylifera. Here we are confronted with a elder brother Francis died in 1852 the family problem that dogs all those interested in the firm became Cook, Son & Co. He then decided history of Victorian horticulture – the it was time to begin making his garden in complexity of synonymy and inaccurate Portugal and planted the first Araucarias at botanical nomenclature, even when used by Monserrate in the same year. Re-building of the most eminent botanists. This is not the house was begun in 1856 and completed surprising when one considers the remote by 1860. origins of the palms and the difficulty that scientists had in procuring flowering In 1869 he inherited a fortune of two million specimens for taxonomic examination. The pounds. sheer size of the made even making herbarium sheets a daunting prospect. Francis Cook was a collector. All things rare and wonderful fascinated him. His garden Throughout this article the names of plants contained Etruscan sarcophagi and Mughal are referred to as cited by original authors.

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2. Palms growing between two giant Araucarias, including Livistona australis and L. chinensis, Washingtonia robusta, Archontophoenix cunninghamiana and Phoenix canariensis.

Please refer to Table 1 for modern accepted Latania borbonica Hort. as opposed to Latania names in accordance with the Kew World borbonica Lam. The first, as undoubtedly Check List of Selected Plant Families intended by Dr. Moore in 1867 is equivalent (http://apps.kew.org/wcsp/home.do). There are to Livistona chinensis (Jacq.) R.Br. ex Mart., however situations in which the treatment of whereas the latter is botanically considered horticultural names does not agree with synonymous with Latania lontaroides (Gaertn.) botanical synonymy. An example would be H.E. Moore.

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3. Monserrate: main entrance to the house with Archontophoenix cunninghamiana.

Garden making in Portugal in the mid- to be found out in the park. The greenhouse nineteenth century was dominated by two may be visited today; unfortunately it is totally aristocratic figures. Foremost was Ferdinand bereft of plants, and sadly the Jubaea no longer Saxe-Coburg and Gotha, king-consort of exists. Queen Maria II and regent during the infancy of his sons Pedro V and Luis I. He was a cousin The Duke of Palmella had in his service two of Queen Victoria’s husband Albert and exceedingly competent plantsmen, the nephew of Leopold of Belgium. In Portugal he botanist Friedrich Welwitsch and the gardener became known as Dom Fernando II. Secondly Jacob Weiss. Together they made a series of there was the Duke of Palmella, Dom Pedro de fantastic gardens on the Duke’s estates, the Sousa Holstein, an important figure in the long most important of which, at Lumiar in Lisbon, period of civil war, protector of Queen Maria, is today a public park. The first palms were and briefly prime minister of Portugal. brought to Lumiar in 1856 and were later planted in the open air. These were Jubaea Dom Fernando II made a number of gardens spectabilis, Chamaerops ghiesbreghtii, Chamae- in Portugal, the most important being that rops excelsa, Livistona sinensis, Phoenix leonensis surrounding his castellated palace of Pena on (P. spinosa), Copernicia sp. from Bahia, Brazil, top of the Serra de Sintra, but also others in Rhapis sp. aff. flabelliformis, Rhapis aspera, Lisbon at the Palace of Necessidades and in Chamaerops tomentosa; Seaforthia elegans; Sabal royal residences such as Alfeite on the southern umbraculifera, and of course, Phoenix dactylifera margin of the Tagus River and at Mafra, north and Chamaerops humilis. Livistona australis was of Sintra. He is famous amongst palm also planted in this garden, though somewhat enthusiasts for the fantastic Jubaea that he later. planted outside at Necessidades in 1858, but in fact the vast majority of his palms in this So it was against this aristocratic background garden were grown in his wonderful domed that Francis Cook began to plant his garden at palm house built in 1857. Before the Jubaea, Monserrate. His palms soon grew to impressive only Cocos flexuosa and Phoenix dactylifera were dimensions as recorded by another botanical

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4. Young Archontophoenix cunninghamiana growing next to Nolina longifolia.

visitor, Professor T. C. Archer from Edinburgh, Seaforthia Elegans, Chamærops Fortunei and who wrote up his Botanical Notes on the Garden Martiana, each about forty-five feet high, of Montserrate, Portugal in 1870. Archer Corypha australis, Sabal Blackburniana, and the considered that Monserrate surpassed even date palm.” Kew since “the whole of the plants at Forty-five feet high, or nearly fourteen meters Montserrat have no other covering than the is a tremendous height for any garden palm in azure sky above.” Europe at this time. The palm house at Kew is On the western side of the upper lawn he saw 66 feet (20 m) at its highest, but outside what “a large grove of palms, on the borders of was there to rival these dimensions? As a direct which are magnificent specimens of Cycas comparison the very first Trachycarpus fortunei revoluta and Cycas circinalis, the former with planted in the palm house at Kew had reached two immense cones of ripe fruit. The palms are only 28 feet by 1860. Prince Albert’s specimen chiefly Areca sapida, Latania borbonica, planted out at Osborne House (Isle of Wight)

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PALMS Luckhurst: Palms of Monserrate Vol. 55(1) 2011 Liebm. in Mart. (H. Wendl.) Toledo (H. Wendl.) (Wall. ex . Mart.) (Wall. H. Wendl (Cham.) Glassman (Hook.) H. Wendl. L. (Sol. ex G. Forst.) & H. Drude Wendl. (Gomes) Kuntze (Gaertn.) H.E. Moore (C. Moore & F. (C. Muell.) Moore Becc. & F. (Jacq.) R. Br. (R. Br.) Mart. (R. Br.) (F. Muell.) Becc. (F. (Molina) Baill. S.Watson (Mart.) Becc. L. Mart. Latania lontaroides Livistona chinensis Jubaea chilensis Livistona australis Allagoptera arenaria Euterpe edulis Howea belmoreana Lytocaryum Lytocaryum weddellianum Howea forsteriana Trachycarpus martianus Trachycarpus Syagrus romanzoffiana Chamaerops Chamaerops humilis Chamaedorea Chamaedorea pochutlensis fortunei Trachycarpus Caryota urens Brahea armata Butia capitata Currently Accepted Checklist, Name Kew) (World sapida Wall. Wall. Hook. hort. ex L. (Gaertn.) H. Wendl. C. Moore (F.Muell.) Becc. (F.Muell.) hort., non Lam. Kunth in HBK R. Br. Hook. f. (Mart.) Becc. L. Mart. Linden Sol. ex G. Forst. H. Wendl. Latania lontaroides H.E. Moore Latania borbonica (Gomes) Vasc. (Gomes) & Vasc. Franco Euterpe edulis Jubaea spectabilis Cocos Weddelliana Corypha australis Diplothemium arenarium Kentia belmoreana & F. Muell. & F. Howea Forsteriana ex Mart. Cocos plumosa Chamaerops Chamaerops martiana Chamaerops Chamaerops humilis Chamaerops Chamaerops fortunei Chamaedorea Chamaedorea elatior Caryota urens Brahea roezlii Butia capitata Areca Areca sapida 1923 1867 1885 1923 1885 1870 1946 1885 1885 1885 1867 1946 1867 1929 1885 1929 1946 Date Most likely match 1867 Latania lontaroides Latania borbonica Euterpe edulis Jubaea spectabilis Cocos Weddelliana Corypha australis Diplothemium arenarium Howea (Kentia) Belmoreana Howea Forsteriana Cocos plumosa Chamaerops Chamaerops martiana Chamaerops Chamaerops humilis Chamaerops Chamaerops Fortunei Chamaedorea Chamaedorea elatior Caryota urens Brahea Roezlii Butia capitata Species Areca Areca sapida Table 1. Historic Table Palm Collection at Monserrate. Dates refer to Literature Cited. 1867 = D. Moore. Archer 1870 = T.C.

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PALMS Luckhurst: Palms of Monserrate Vol. 55(1) 2011 H. Wendl. & H. Drude Wendl. (F.Muell.) H. Wendl. & H. Drude Wendl. (F.Muell.) H. Wendl. Mart. (R. Br.) Blume (R. Br.) (Linden ex André) ex H. de Wendl. Bary (Hook. f.) & H. Drude Wendl. Chabaud L. O’Brien (L.) Roxb. Jacq. (Lour.) Merr. ex A. Merr. Chev. (Lour.) T. Anderson T. (Walter) Lodd. ex (Walter) Schult. & Schult.f. (Thunb.) Henry Archontophoenix Archontophoenix cunninghamiana brasiliensis Trithrinax robusta Washingtonia Sabal palmetto elegans Rhapis excelsa Phoenix reclinata Phoenix roebelenii Phoenix rupicola Phoenix canariensis filifera Washingtonia Phoenix sylvestris Phoenix dactylifera Livistona saribus Teijsm. Teijsm. H. Wendl. Mart. Glazebrook L. Linden ex André Hook. R. Br. (L.) Roxb. Jacq. T. Anderson T. Verschaff. (Thunb.) Henry Hook.f. Seaforthia elegans Trithrinax brasilensis Trithrinax robusta Washingtonia Sabal blackburniana Seaforthia elegans Areca Areca baueri Rhapis excelsa (F. Muell.) & H. Drude Wendl. (F. Archontophoenix Archontophoenix Alexandrae Archontophoenix alexandrae Phoenix reclinata Phoenix rupicola Pritchardia filifera Phoenix tenuis Livistona Hoogendorpii Phoenix sylvestris Phoenix dactylifera & Binn. ex Miq. 1867 1929 1929 1870 1867 1885 1946 1885 19461885 (misidentification) 1885 1891 1891 1885 1891 1867 Seafortia elegans Trithrinax brasiliensis Trithrinax robusta Washingtonia Sabal Blackburniana Seaforthia elegans Rhopalostylis (Areca) Baueri (Archontophoenix (Archontophoenix Alexandræ) Rhapis excelsa Ptychosperma Alexandræ Phoenix loureirii Phœnix reclinata Phœnix rupicola Canariensis des P. Pritchardia fillifera Phœnix tenuis qui sont Table 1, continued . Table Livistona Hoogendorpii Phænix sylvestris Phœnix dactylifera

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and thought to be the largest of its species on the French Riviera in the nineteenth grown without shelter in Europe, was at this century it often succumbed to cold winters time only ten feet tall. without reaching impressive dimensions. Archer tells the story of how Cook’s gardener Corypha australis, in this context should be had transplanted a giant date palm. “It is interpreted as Livistona australis (R. Br.) Mart. supposed to be some centuries old, and Seeds placed casually as drainage material in formerly grew at Cascaes, a place twelve miles the bottom of Wardian cases sent by distant, whence it was removed by Mr Burt, the Cunningham were found to have germinated gardener at Montserrat, across the intervening on arrival at Kew in 1824. Conrad Loddiges Serra with great difficulty. Its were tied then used this technique to import large up, and its roots protected, and twenty-four numbers of seeds and the palm became quickly oxen worked for a whole week to bring it to established in the greenhouses of Europe. This the paradise in which it now so proudly palm was among the first contenders for flourishes ... The stem of this prince of palms acclimatisation since it is the southernmost is seven feet six inches in girth, and the height palm occurring on the Australian continent. not less than thirty feet, — the beautiful crown Small plants were growing in the open at the of leaves being at least ten feet more.” Naples Botanic garden at by 1867, and it was included in a list of palms that were The first Areca sapida (Rhopalostylis sapida) “reasonably common” at Nice in 1869. flowered in the palm house at Kew in 1859. The following year the French Revue Horticole Sabal blackburniana was first cultivated under was appealing to “la liberalité anglaise” so that glass in Europe by a Mr. Blackburn who had seeds might be supplied to southern gardens his plant from Lord Petre in 1737. It was for experimentation with outdoor culture. By probably grown from seeds collected in 1861 Messrs. Veitch & Son had the plant on Georgia (USA) by William Bartram. A sale in London and the plant was “tolerably multiplicity of other names makes it difficult well known in gardens.” On the French Riviera to determine when this palm was first it was not until 1882 that this palm reached cultivated outdoors. In fact, the earliest records flowering maturity. use a wholly erroneous designation: Corypha umbraculifera. Famously large specimens grown Latania borbonica is not what it seems. This under glass are frequently referred to by this name was used by Victorian gardeners for name. Messrs. Huber of Hyères on the French Livistona chinensis (Jacq.) R. Br. in the mistaken Riviera planted out a very young specimen of belief that this Chinese palm originated from Sabal blackburniana in the summer of 1864 and the French Island of Réunion (Île Bourbon). it was successfully over wintered. Perhaps the first open air planting of this species was at Hyères in 1846, on the French Encouraged by these early successes: “Mr. Cook Riviera some ten years before it was cultivated ... has lately experimented with Palms that he in Portugal. was chary of trying at first; a list of those that are healthy will, therefore, be valuable. They Seaforthia elegans is another problematic are Phœnix reclinata, Ptychosperma Alexandræ gardeners’ synonym, almost certainly what is (Archontophoenix alexandrae), Cocos plumosa intended is Archontophoenix cunninghamiana and Weddelliana, Howea (Kentia) Belmoreana, H. Wendl. & Drude. Some descriptions of and Rhopalostylis (Areca) Baueri.” This list Monserrate list both Seaforthia elegans and appeared in 1885 written by C.A.M. Ptychosperma cunninghamiana as growing side Carmichael in a long two-part article for The by side. So we should perhaps consider the Gardeners’ Chronicle. By then Seaforthia elegans, possibility that Ptychosperma elegans (R. Br.) Livistonia borbonica and Corypha australis were Blume was also cultivated at Monserrate. the tallest palms in the garden. However there is none growing there today and the Archontophoenix is still seen In another part of the garden known as throughout the gardens (Figs. 3 & 4). Not a “Mexico” Carmichael found that “Aloes and particularly hardy palm, what probably Yuccas of every kind abound, and monotony prompted the experimentation with is avoided by inserting Palms.” He gave the Archontophoenix at this early date was its ready following list: “Areca (Rhopalostylis) sapida, one availability by seed imported from Sydney. of which measured 12 feet; two wide- Archontophoenix cunninghamiana is very spread Date Palms, the circumference of the sensitive to cold and although widely planted branches of one was 39 paces; Phœnix rupicola,

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Caryota urens, Livistona Hoogendorpii, Howea forsteriana, with trunks 25 ft. high; Phoenix Forsteriana, and Euterpe edulis were among the canariensis, P. dactylifera, P. reclinata, with seven most prominent or worthy of notice for being well-furnished trunks from one root; Brahea grown in the open.” roezlii, Areca baueri, A. sapida, Jubaea spectabilis, Seaforthia elegans, Sabal blackburniana and Sabal Amongst this list are a number of palms that umbraculifera.” At other points in the article he even today’s enthusiasts of hardy palms would also mentions Trithrinax brasilensis, and be surprised to encounter. How many Chamaedorea elatior plants not mentioned in European gardens today are growing any previous description of the garden and Lytocaryum weddellianum (H.Wendl.) Toledo or presumably from his own introduction. Euterpe edulis Mart. in the open? The Euterpe and Cocos insignis were still thriving in 1910 Sir Herbert died in 1939. Following the Second as described by the French forester Léon Pardé. World War, the Monserrate estate was sold by The warm growing conditions found at the fourth baronet, who shared his name with Monserrate are indicated by the presence of his great grandfather, Sir Francis Cook. The such delicate palms as Howea belmoreana, rated garden eventually passed into the hands of zones 10 and 11 by Robert Lee Riffle and Paul the Portuguese state, managed by the Forest Craft in their Encyclopedia and Archontophoenix Service. The gardens continued as a great alexandrae rated 10b and 11. Both species are tourist attraction into the 1960s but were better growing today at Monserrate. Recently planted known for their rhododendrons and camellias Caryota urens (adapted only to zones 10b and than the exotic plants of the Victorian era. 11 and marginal in 10a) has come through the The palms continued to grow, but slowly many difficult winter of 2009/2010 and is making were lost either through old age or more new growth at the time of writing (March significantly through competition from 2010). invasive trees species such as Acacia melanoxylon and Pittosporum undulatum. Sir Francis Cook died in 1901, appropriately at the close of the Victorian era. He was Following the 1974 Portuguese Revolution succeeded by his son Frederick, who was little circumstances had changed so radically that it interested in horticultural affairs but who had the good sense to entrust the garden to Henri Visiting Gardens Navel, trained at the Ecole de Versailles and at Kew and who was later appointed head Monserrate gardener at the Lisbon botanic garden. The Parque de Monserrate gardens were visited during his tenure, as 2710-405 Sintra, Portugal mentioned above, by Léon Parde. His extensive Tel: 21 923 73 00 botanical descriptions indicate however that Fax: 21 923 73 50 the palm collection at this time was essentially that planted by Sir Francis. Parques de Sintra – Monte da Lua E-mail: [email protected] Only when the third baronet, Sir Herbert http://www.parquesdesintra.pt/ Cook, appointed Walter Oates as his head gardener was the collection to see new Palácio das Necessidades development. Walter Oates wrote Monserrate: Tapada das Necessidades A Short Guide to the Gardens in 1923, shortly Rua Capitão Afonso Pala after he had arrived as new head gardener He Lisboa 1350-215 had been working previously on the Italian http://www.mne.gov.pt/mne/en/ministerio Riviera and was well acquainted with hardy /palacio/ palms. Oates remained at Monserrate for the rest of Lumiar (Museu National do Trajo e da the decade and wrote a well-illustrated article Moda) for the Gardeners’ Chronicle in 1929 in which Parque Monteiro-Mor he lists many plants. He includes a list of Largo Júlio Castilho palms: “Palms are very numerous, and 1600-483 Lisboa mention of only a few of the best must suffice: Tel: (+351) 21 759 03 18 Cocos flexuosa. C. romanzoffianum, Washing- Fax: (+351) 21 759 12 24 tonia robusta, the tallest palm in the garden, http://www.museudotraje-ipmuseus.pt about 70 feet high; Kentia belmoreana, K.

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was impossible to maintain the garden. Staff COOK, H. AND W. OATES. 1923. Monserrate, were reduced to three people: one collecting Cintra, Portugal: a Short Guide to the tickets on the gate, one sweeping up around Gardens. Privately published. the house and the last of the old gardeners, José Luis, who faced with the forests of invasive CARMICHAEL, C. 1885. Montserrat. Gardeners’ plants that occupied the garden, retreated to Chronicle 23: 389–91, 426–427. the broken glasshouse and nursery. GOEZE, E. 1875. Jardins notaveis em Portugal, In 1987 a group of young Canadian landscape Lumiar. Jornal de Horticultura Practica 7: architects and horticulturists from the Royal 230–233. Botanical Gardens of Hamilton, Ontario began GOEZE, E. 1876. Jardins notaveis em Portugal, to work on the restoration of the gardens. This Palacio das Necessidades. Jornal de Horti- was the beginning of a long and difficult cultura Practica 6: 43. process that has yet to be completed. Today the gardens are managed by a state-owned, but GOMES, A.G. 1855. Da Aclimatação em Portugal self-funding organization called the Parques do Cycas revolutas, uma das árvores do Sagu. de Sintra – Monte da Lua. There is a Flora e Pomona 2: 108. comprehensive restoration project underway HARDY, A. 1864. Note sur la situation des of both house and gardens, co-ordinated by derniers plantations d’espèces ligneuses Prof. Antonio Lamas, the organization’s exotiques au Jardin d’Acclimatation a Alger. president. Bulletin de la Société d’horticulture du And the palms? Twenty-four species have département de Seine-et-Oise. p. 24. managed to survive from the original LEPELLETIER, M. 1860. Floraison de l’Areca sapida plantings, some of them centenarian and of dans la grande serre de Kew. Revue Horticole monumental dimensions. In addition, clearing 1860: 18, 19. work and reconstruction have enabled a program of replanting of the garden’s botanical LUCKHURST, G.L. 1988. Sintra: a Landscape with heritage. This includes the palm collection Villas. Edições Inapa, Lisbon. which today includes some 70 different taxa. A new head gardener has been appointed, Tim MOORE, D. 1870. Botanical and horticultural Stretton, who trained at Kew. One hopes the notes, made on a journey through parts of day will soon be here when Monserrate will the south of France, Spain and Portugal once again resemble the palm house of Kew. during the autumn of 1867. Journal of the But under the azure sky above! Royal Dublin Society 5: 241–255.

LITERATURE CITED OATES, W. 1929. Monserrate. Gardeners’ Chronicle 86: 90–93. ARCHER, T.C. 1870. Botanical notes on the garden of Montserrat, Portugal. Transactions PARDÉ, L. 1910. Excursion dendrologique au and Proceedings of the Botanical Society of Portugal et en Espagne. Bulletin de la Société Edinburgh 10: 414–420. Dendrologique de France 17: 118.

BEAN, W.J. 1925. Trees and Shrubs Hardy in SAINT VICTOR, L.G., 1891. Portugal: Souvenirs et the British Isles. John Murray, London. Impressions de Voyage. Librairie Blériot, Paris. BERGMAN, E. 1890. Palmiers en Portugal, Jubaea spectabilis. La Nature: revue des sciences et VASCONCELLOS, J. & J.A. FRANCO 1946. As de leurs applications aux arts et à l’industrie palmeiras de Lisboa e arredores. Portugaliae 888: 40–42. Acta Biologicae 2–4: 289–245.

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