PALMS. BOTANICAL MONOGRAPHS by Jardí Botànic de la Universitat de València

Botanical monographs

PALMS

The Botanic Garden of the Universitat de ValĂ¨ncia

JosĂŠ Plumed Manuel Costa

Botanical monographs

PALMS JosĂŠ Plumed and Manuel Costa

Botanical Monographs. Jardín Botánico de la Universitat de València (Botanic Garden. University of Valencia) Volume 1: Palm trees © texts: José Plumed and Manuel Costa © foreword: Esteban Morcillo and Isabel Mateu © pictures and illustrations: José Plumed © edition: Universitat de València E. G. Director of the Collection: Isabel Mateu Technical Director: Martí Domínguez Graphic design and layout: José Luis Iniesta Translation: Alicia Donnellan, Interglobe Language Photographs: José Plumed Printing: Gráﬁcas Mare Nostrum, S. L. Legal Deposit: V-2262-2013 ISBN: 978-84-370-9132-7

Index

Foreword ..............................................................................................................................................................7 Introduction .................................................................................................................................................. 11 The Arecaceae family .........................................................................................................................15 – General description.............................................................................................................................. 17 – Stem................................................................................................................................................................18 – Crown ............................................................................................................................................................19 – Terminal bud .......................................................................................................................................21 – Leaves .....................................................................................................................................................23 – Root system...............................................................................................................................................26 – Flowers..........................................................................................................................................................27 – Inﬂorescences..........................................................................................................................................29 – Fruits and seeds .....................................................................................................................................30 – Distribution and ecology...................................................................................................................31 – Systematics................................................................................................................................................32 – Systematic position of the palms in the Botanic Garden ...........................................33 – Living together: fauna and ﬂora associated to palms in the Botanic Garden ..34 – Uses and exploitation of palms ....................................................................................................35 – Palms and climatic conditions ......................................................................................................37 – Wind .........................................................................................................................................................37 – Sun ............................................................................................................................................................37 – Storms .....................................................................................................................................................38 – Fire.............................................................................................................................................................38 – Cold ..........................................................................................................................................................39 Genus data sheets..................................................................................................................................41 Glossary of terms ................................................................................................................................123 Bibliography...............................................................................................................................................129 List and plan of species ...............................................................................................................131

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Foreword

he Botanic Garden of the Universitat de València was founded in 1567, initially linked to medical studies, today this superb place is devoted to teaching, research and culture. A landmark in the city of Valencia, it is a true living museum housing a diversity of plants in its various collections, exhibiting their characteristics and uses. Here plants from exotic countries are grown that would hardly be known otherwise. One of the most exotic and highly valuable plant collections is the palm garden with over 80 species and 335 specimens, many of which are truly monumental. Palms include nearly 3,000 species, grouped within the botanical family Arecaceae which, despite its variety, form an easily identifiable monocot group. Palms are instantly recognizable, borne upon woody stems, which are usually unbranched and crowned with a fan of leaves, and exhibiting other hallmark floral traits. Palms are typical of tropical and subtropical climates, with outdoor cultivation being limited to areas with a mild climate and without hard frosts. For this reason, despite the great wealth in variety of palms, only two species are to be found in Europe, Phoenix teophrastii, distributed over a small area in the eastern Mediterranean, with small populations on the Greek island of Crete, as well as in Turkey, and the Mediterranean fan palm (Chamaerops humilis), the only Iberian palm, inhabiting the north and southern slopes of the western Mediterranean basin. To these we should add the Canary palm (Phoenix canariensis), native and endemic to the fortunate islands that gave it its name. Many palms or their derivatives, such as the fan palm, date palm, oil palm, betel nut, rattan, carnauba wax or raffia, make them economically important and of commercial interest, both for the products they provide and for their ornamental value. The mild climate of Valencia, and much of this region, enables a wide variety of palms to be grown, among which is the date palm (Phoenix dactylifera), traditionally cultivated near farmsteads and in urban gardens. Thus species such as the Canary palm or species of Washingtonia, Trachycarpus and Livistona

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Aerial View of the Jardín Botánico, Valencia.

all form part of our landscape. In recent years the cultivation of palm trees has rocketed due to the increase in landscape gardening. Indeed, a consequence of this uncontrolled trade in palms has been the introduction of the red palm weevil, the pest that is on its way to leaving us a landscape devoid of our more traditional palms. This superbly documented book compiled by Manuel Costa and José Plumed includes the 39 genera represented in our Botanic Garden. Here, the reader will ﬁnd a great deal of information presented in summary form. Next to the common and Latin names, the book provides the etymology, taxonomic and ecological data of interest, distribution maps, as well as traditional uses and exploitation. The high quality images and texts, carefully and impeccably reviewed by José Manuel Alcañiz, are highlighted by the excellent design and layout, for which

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we are grateful to the prestigious and award-winning popular science journal Mètode. To be fair, we should make it clear that this collection, and therefore this book, would not exist without the efﬁcient management and care of the garden, diligently run by the gardening staff and the various curators of our Botanic Garden, whose dedication has safeguarded these ancient trees for us to enjoy. One of the missions of the Botanic Garden is to maintain and improve plant collections, and one way of contributing to this is to make them known. This book is the beginning of a series of monographs on the plant collections hosted by the Botanic Garden of the Universitat de València. The publications mirror the scientiﬁc rigor of a botanical garden itself, but also target the non-specialist, trying to awaken interest in the world of plants, which is all too often largely unknown. Thus, this series bridges a gap and is useful both for our visitors and for anyone who, like us, feels a real fascination for plants and all that surrounds them. Esteban Morcillo Vice-Chancellor of the Universitat de València

Isabel Mateu Director of the Botanic Garden of the Universitat de València

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Introduction

he Botanic Garden of the Universitat de València dates back to the midsixteenth century, when the medical school saw the need for a simple garden to study medicinal plants. The garden was placed in different locations across the city until the year 1802, when it found its final and definitive location outside the fortress walls of the city. It was placed in the so-called Garden of Tramoyeres, the street now known as Quart. The Botanic Garden is part of the Universitat de València and is a centre for research, education and conservation. Today the Botanic Garden is one of the oldest and most important botanical gardens in Spain, not only because of the richness and number of its species and collections, but also because of its age, monumentality, originality and quality of the plants it hosts. The palm collection is amongst the most outstanding hosted at The Botanic Garden, as evident by the slender and subtle silhouettes of some of the larger specimens, towering above even the most majestic monumental tree tops. The collection is distributed throughout many different areas of The Botanic Garden: in the “Escuela Botánica” (Botany School Section), lining the main walkway, next to the ponds and buildings and inside the shade house or greenhouse. The palms have been present in the Garden almost since its creation. The first available data on record are in the first catalogue of cultivated plants published in 1856 by its director at the time José Pizcueta, listing more than 6,000 species, of which 35 were palm trees planted outside. Some of them, those most resistant to cold, have survived to this day. In 1930, Francisco Beltrán, then director of the garden, made a new list of plants of which 35 palm species remain, where the most sensitive to cold are missing, and those most resistant to it have been maintained and increased in number. These palms are now adult specimens. Restoration work was carried out in the garden in 1987, allowing the revision and enrichment of the collections. In 1989 numerous palm trees were introduced in an effort to acclimatize them; consequentially, today we can admire interesting species, varieties and hybrids. The greenhouse called La Balsa built in 1888 was restored and renovated in 1990, for growing the species most sensitive to cold.

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The Arecaceae Family

alms are identiﬁed with tropical environments, be it the rainforest or the broad savannah, the high mountains of the tropics or the deserts’ oasis. At present, they constitute the world’s second most economically important plant group, and millions of people in tropical and subtropical areas of the planet depend on them, directly or indirectly, to survive. However, there are many species of palms cultivated and planted outside the tropics. This occurs especially in areas with a Mediterranean climate, where a great variety of them can survive outdoors. In contrast, in temperate and boreal countries they are usually grown in greenhouses.

On the previous page, aerial view, tropical forest in Rio Napo, Ecuador. Above, Chamaerops humilis. Monsul Beach. Cabo de Gata (Almería).

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To the left, dead specimens of the canary palm, attacked by the red weevil, Almenara (Castellón). On the right, Rhynchophorus ferrugineus, adult specimens.

Some palms can even survive naturally outside the present tropical range (in Paleo and Neotropical regions) and these areas constitute their primary habitat. This applies to the European Chamaerops humilis and Phoenix theophrasti in the Mediterranean basin. This is also the case with Phoenix canariensis, taking refuge in the ravines of the islands that give it its name, the Canaries. Extensive palm “groves” of Phoenix dactylifera, in this case cultivated but of ancient origin, can be seen in places like Elche, Orihuela and Crevillente. The climatic diversity of Spain would allow viable plantations of various species of these beautiful plants. However, the reality is different. Plantations are scarce and monotonous, usually with the repetitive use of the four or ﬁve most common and popular species. The importation of palm trees without proper phytosanitary control has facilitated the introduction of various pests and diseases into Europe. Notable among them is the outbreak of the red palm weevil (Rynchophorus ferrugineus), which is endangering the entire palm populations of the Mediterranean basin.

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Brahea armata.

General Description The Arecaceae family includes many genera and species. The total figure is constantly changing, due to the continuing discovery of new species and classification changes experienced as the family is studied in greater depth. At present, the number of genera comprised is about 180, which represent 2,400 species. In this guide we have taken as reference the classification published in Genera Palmarum. The evolution and classification of palms (2008), written by John Dransfield, Natalie W. Uhl, Conny B. Amussen, William J. Baker, Madeline M. Harley and Carl E. Lewis. The family Arecaceae (Arecaceas) is the only in the order Arecales (old order Príncipes) within the Monocotyledons, and it includes all palm trees, in fact it was called Palmaceae before receiving its current name. It is a family of great originality and represents quite a natural and homogeneous group, since the various species combine: arboreal stand, wide leaf blades and a well-developed vascular system in all vegetative organs, consisting of numerous small separated vascular bundles, each surrounded by a fibrous sheath. On their stems the

Botanical Monographs 17

Stem cross section. Phoenix dactylifera.

scars left by falling leaves are frequently found. They are usually plants with unbranched stems (monopodial), although in some species like Nypha and Hyphaene dichotomous branching may occur. Also, the stems are usually capped by a characteristic crown of leaves, each endowed with a wide clasping base. The leaves are quite complex and can come in various shapes and sizes. The inflorescences are also very diverse in form. The flowers are unisexual or bisexual and plants monoecious or dioecious, although in some cases they may be polygamous. Pollination can be anemophilous (by wind), but more generally it is entomophilous (by insects) and some species are pollinated only by beetles (coleopterophilous). The fruit is fleshy, a monospermic berry or a drupe.

Stem The trunk of the palm trees, called stem or stipe, lacks secondary growth in thickness. It consists of ligniﬁed elements, it is solid and has several functions: to support and raise the crown, to act as a link between the roots and the aerial structures, and accumulate reserve substances, especially water and starch. During the ﬁrst years of the palm’s life, the stem grows thick and tall, but once it has reached the size typical of its species, it only grows in height.

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From left to right, Phoenix dactylifera, Phoenix canariensis and Archontophoenix alexandrae.

The stems have a varied array of shapes, dimensions, surfaces and textures, being elements of diagnostic value for determining the species. They can be straight, solitary or ringed throughout their length; they may have a swollen or bell-shaped base. They may also grow prostrate, creep along the ground or on other plants, or are subterranean. In some species, the stems are very thick, cylindrical and smooth while in others various swellings or “bellies” may be observed. Stems may be spindled, elongated and tapered at the ends. There are few genera with dichotomous branching, but others may branch anomalously if the terminal bud is damaged. There are species with very short stems, some are even acaulescent, while others may become very large, raising their crown to more than 60 m in the air. In a cross section the stems present three distinct areas: the core, the pericycle and the cortex.

Crown The crown is one of palm’s most characteristic elements, in general and for some species in particular. There are palm trees with very leafy tops, others however, are quite translucent and have few leaves. Their shape may be globose,

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Livistona australis.

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subglobose, hemispheric, or obconical. In general, each individual species presents a certain number of leaves, which are usually constant during the life of the specimen. New leaves, situated at the apex, sprout from the centre of the crown in upright position and are shaped like a duster; eventually the young replace more adult leaves as they move towards the base, taking each-otherâ&#x20AC;&#x2122;s place. From the apex downward ďŹ rst the new adult leaves appear, then the mature and ďŹ nally the dead leaves, brown and located at the bottom of the crown. There are species in which the leaves wither but do not fall for many years (marcescent), remaining dry and adhering to the stem. In others, however, they are shed soon after drying. The crown of the palm is designed to capture the maximum possible solar energy and to offer minimum wind resistance. In the case of strong winds, some palms shed their leaves to reduce the pressure on the stem and avoid collapse. In the crown there are three distinct parts: the bud, the leaves and the reproductive system.

Terminal Bud The terminal bud is a key element in the life of the plant. It is a growth point; the only generating point (meristem) the palm trees posses on the top of the stem. It is the most vulnerable point of the plant, because if it dies so does the palm. For this reason, it is usually protected inside the stem, buried at a depth that varies with the species. Palms protect the bud from the weather and animals through

From left to right, apical bud of Phoenix canariensis, Phoenix reclinata and Archontophoenix cunninghamiana.

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From left to right, leaves of Chambeyronia macrocarpa and Chamaerops humilis.

physical and chemical barriers: the sheaths and petioles of the leaves may be heavily armed with thorns, contain poisonous substances or a high concentration of tannins. The rest of the organs that make up the palmsâ&#x20AC;&#x2122; canopy are formed from its spiral twisting growth. In branched palms (clustered) terminal buds are to be found at the extreme tip of each stem. Sometimes, and in some species, basal buds are observed, even attaining a certain height, which can generate new stems.

Leaves Palm leaves are similar to those of other plants and have the same basic structure, but with intrinsic characteristics particular to each species Young leaves sprout in the central part of the apex, upright, folded, shaped like a sword or spade and protected by thick cuticles, waxy coverings, tomentum and scales

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From left to right, leaves of Sabal domingensis and Caryota maxima.

to preserve them from external environmental agents. Some of these protections are deciduous and therefore are shed when the leaf matures. The basic function of leaves is to produce sap sugars through photosynthesis. The two most common leaf types are: palmate leaves (fan-shaped) and pinnate or feather-shaped leaves, but we can also ďŹ nd costapalmate and bipinnate leaves. The size varies, but generally they tend to be large. The largest leaf known, that of the African palm Raphia regalis, can reach more than 25 m in length. The leaves, though usually green, can have different variations and shades of colour, usually associated with different environmental conditions.

Pinnate leaves Of the various leaf forms, we will limit the description to pinnate ones, being so common that they are almost ďŹ ve times higher in number than palmate-leaved

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species. They are formed by the sheath, the petiole and the limbus, which in turn is composed of the rachis and the pinnae. The sheath or leaf base is the portion of the leaf’s blade which connects to the stem, clasping it to a greater or lesser degree depending on the different species. It is also the place where the vascular bundles penetrate from the stem. Species like Phoenix roebelenii have a very small sheath, but clasp the stem and are often accompanied by various fibres woven into a kind of burlap. Others are broad and slightly clasping, such as Phoenix canariensis, in which the remains of the sheath usually stay attached for a long time, leaving scars with the imprint of their shape on the stem once they fall. The leaves may also have clasping sheaths which completely surround the stem and are spaced apart, as in Chamaedorea, or sprout close together and remain clustered as in Hyophorbe or Archontophoenix. In this case, they form a bulge at the end of the stem known as crownshaft. The crownshaft is distinctive in some palm trees and facilitates identification. Usually it presents many shades of green or is red, orange or yellow. It can also be covered by various hairs (floccose), tomentum and scales of different colours. The petiole is the main axis of the blade, situated between the sheath and the first pinna. It has a widened base that tapers towards the rachis; it is strong and resistant, but flexible. It may be covered with thorns or spikes along its entire cuticle

epidermis subepidermis sclerenchyma ﬁbres

vein

chlorophyll parenchyma

vascular bundle xylem

stoma Leaf cross section. Chamaerops humilis.

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stoma

Photo: Ana Ibars

phloem

Pinnate

Bipinnate

Palmate

Pinnae

Costapalmate

segments

Lamina

Phoenix

Rachis

Ligule

Petiole Seath

Caryota

Chamaerops

costa

Livistona

Sabal

surface, only at the margins, or be unarmed. It may also be partially covered by various fibres, tomentum, scales or hairs of various colours. The rachis is the continuation of the main leaf stalk along which the pinnae are inserted. It is wider at the base, gradually decreasing towards the apex, cuneiform, rounded or grooved. Usually it is more or less straight, but sometimes it can become curved, as in Butia capitata, giving the leaf a characteristic appearance. Like the petiole, it can also present various colourings and be partially covered with wax, scales, hairs or tomentum. The pinnae are highly varied in shape and size. They can be narrow or wide, rigid or pendulous, linear, lanceolate, cuneate, with a termination that may be acute, obtuse, toothed, premorse, etc. The ones nearest the petiole can be modified and present thorn-like shape, as in Phoenix, and in this case they are called acanthophylls. Pinnae form a row arranged on each side of the spine, either in one plane, as in Ravenea rivularis, or forming a V, as in Howea belmoreana. They may be disposed regularly in a single plane and with opposite pinnae as Chamaedorea radicalis, alternate as in Pinanga diksonii or grouped as in Aiphanes horrida. They can also be positioned forming several planes, such as Syagrus romanzoffiana or Dypsis madagascariensis. Most species have reduplicated pinnae, i.e., shaped like a tent or a Greek lambda crownshaft Λ, such as Jubaea chilensis, but there are genera that have unduplicated pinnae, channel or V shaped, such as Phoenix or Arenga. Some species have the pinnae at the apex joined together, giving them the appearance of a fish tail, as in Chamaedorea microspadix. The pinnae usually have their central vein visible or prominent in one of their sides (rib), and sometimes with the underside covered with ramenta (scale-like hairs), as in Archontophoenix cunninghamiana; these central veins are often accompanied by several secondary veins and in some species smaller ones can be present forming a kind of transversal grid, as in Jubaea chilensis. Some species, such as Chambeyronia macrocarpa, have the particularity of

Botanical Monographs 25

Root system. Phoenix canariensis.

Root cross section. Phoenix canariensis.

presenting new leaves of a striking red colour. This occurs just when they open and lasts for a short time, later turning green. It is unusual to ﬁnd variegated leaves in this family, but there are always exceptions. On the island of La Palma one can see a Phoenix canariensis with tinted leaves of spectacular beauty.

Root System In palm trees, the roots spring from the basal bud. From this generating point, located at the lower end of the stem, issue large amounts of fascicled roots, fibrous, fleshy and rigid, that extend both radially and superficially, covering large areas in search of water, oxygen and nutrients. Branched (clustered) palms have a basal bud on each of its component stems. The roots, depending on their thickness and position can be of primary, secondary, tertiary or fourth order. The primary roots are longer and thicker, and are responsible for anchoring the plant to the ground. The primary give rise to the secondary and from these arise those of a lower order.

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Palm roots do not present absorbent root hairs, but in the vicinity of the root apex there is a lighter coloured area, not suberized, which is the main region for the absorption of water and nutrients. These roots can also establish symbiotic relationships with other organisms in the soil, particularly endomycorrhizae. The internal structure of the roots has two distinct parts: the cortex and medulla. From a biomechanical point of view, the root system of palms is very effective because it is very dense, plentiful with a great capacity for regeneration. It is unusual to see palm trees killed or uprooted by the wind.

Flowers The flowers of palm trees are generally unisexual, small, although they can also be bisexual. They are arranged on bracts or bracteoles and are pedicellate or sessile. Palms can be monoecious, in which case flowers have both sexes on the same plant or bisexual flowers. They can also be dioecious, when the flowers are unisexual and are in different individuals. Some species are polygamous, with unisexual and bisexual flowers. Pollen grains are contained in the anthers of the stamens and are characteristic of each group. Pollen is an important character for botanists, as it allows comparative study of the current and fossilized pollen. The flowers are often inconspicuous, creamy, white or yellow, although some are more visible and are coloured purple or orange. In some species they are very fragrant and perfumed.

From left to right, Sabal mauritiformis bisexual ﬂower, pistillate and staminate ﬂowers of Arenga englery.

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On the previous page, infrafoliar inﬂorescences and infructescences of Archontophoenix cunninghamiana. On this page, left to right, the fruits of Butia capitata, infructescences of Livistona chinensis and Rhapidophyllum hyxtrix.

At present, due to mass planting of different palm species in nearby locations, hybrids occur between species and genera which would have never occurred naturally, as has happened between the date palm and the Canary Islands palm.

Inflorescences Palm flowers are usually grouped in inflorescences, very ornamental and characteristic. They develop in the leaf axils and provide the specialist with numerous distinctive features useful for classification of the different species. When placed under the leaves they are referred to as infrafoliar; suprafoliar inflorescences bloom at the top of the crown. But most commonly they sprout between the leaves, and then they are called interfoliar.

Fruits and Seeds Fruits are usually berries, as in the case of dates, or drupes, as in the case of coconuts. Their function is to spread the seed they harbour, ensuring the

Botanical Monographs 29

On the left, Barranco de Fataga Gran Canaria, Phoenix canariensis. On the right, sierra Segaria in Beniarbeig (Alicante). Chamaerops humilis.

continuity of the species. In palms, dispersal can occur when the fruit falls down freely by gravity; also through endozoochory, when animals are in charge of dispersion by ingesting the fruit and later excreting the seed. Dispersion can also occur through hydrochory, when the vector is water. After fertilization of the flowers, the inflorescences will become infructescences. The fruits are much more visible than the flowers, with a variety of shapes, sizes and colours. Typically, each fruit contains a single seed, although some species can have three. Many of palm trees’ fruits are edible, but some are toxic as pulp contains calcium oxalate crystals. Seeds come in a great diversity of shapes, sizes and colours. They are generally wrapped in a tough and leathery layer occupied by the albumen. This stores the large amount of nutrient reserves required for the germination and nutrition of the seedling it holds. In general, seeds do not maintain their viability for a long time; in addition, many insects and other organisms feed on them, so it is advisable to plant them as soon as possible after ripening.

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The palm groves of Elche (Alicante). Phoenix dactylifera.

Distribution and Ecology The current distribution of the family is tropical, both Paleo- and Neotropical, a range with uneven distribution. Palm trees occupy a variety of ecosystems, from hyper-humid rain forests to deserts. They can be part of the climatophilous vegetation communities, of permanent edaphoxerophilous or edaphohygrophilous communities on waterlogged soils, with temporal or permanent humidity, and xeric ridges. Some species live outside the Tropics, such as Chamaerops humilis, found on the thermo-Mediterranean ďŹ&#x201A;oor of the Mediterranean basin. Others have adapted to cold climates thanks to mechanisms protecting the apical bud, like Trachycarpus, which can survive altitudes of up to 2,400 m in the Himalayas. Other extra-tropical palms are Phoenix theophrasti, on the island of Crete, and Phoenix canariensis, native to the canyons of the Canary Islands.

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Systematics Palms belong to the Kingdom Plantae, Division Magnoliophyta, Class Liliopsida, Subclass Arecidae, Order Arecales, Family Arecaceae. It is a family with complex systematics; however, and according to the updated classification in Genera Palmarum. The evolution and classification of palms (2008), five subfamilies are recognised, each containing different tribes: SF. Calamoideae Tr. Eugeissoneae Tr. Lepidocaryeae Tr. Calameae SF. Nypoideae SF. Coryphoideae Tr. Sabaleae Tr. Cryosophileae Tr. Phoeniceae Tr. Trachycarpeae Tr. Chuniophoeniceae Tr. Caryoteae Tr. Corypheae Tr. Borasseae SF. Ceroxyloideae Tr. Cyclospatheae Tr. Ceroxyleae Tr. Phytelepheae SF. Arecoideae Tr. Iriarteeae Tr. Chamaedoreeae Tr. Podococceae Tr. Oronieae Tr. Sclerospermeae Tr. Roystoneeae Tr. Reinhardtieae Tr. Cocoseae Tr. Manicarieae Tr. Euterpeae Tr. Geonomateae Tr. Leopoldinieae Tr. Pelagodoxeae Tr. Areceae

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Systematic Position of the Palms in the Botanic Garden SUBFAMILIA

TRIBU

GÃ&#x2030;NERO

Arecoideae

Areceae

Adonidia Archontophoenix Areca Chambeyronia Dypsis Howea Ptychosperma Chamaedorea Gaussia Hyophorbe Aiphanes Allagoptera Butia Jubaea Syagrus

Chamaedoreeae

Cocoseae

Ceroxyloideae Ceroxyleae Coryphoideae Borasseae Caryoteae Chuniophoeniceae Cryosophileae

Phoeniceae Sabaleae Trachycarpeae

Ravenea Bismarkia Latania Arenga Caryota Nannorrhops Coccothrinax Thrinax Trithrinax Phoenix Sabal Acoelorrhaphe Brahea Copernicia Chamaerops Licuala Livistona Pritchardia Rhapidophyllum Rhapis Serenoa Trachycarpus Washingtonia

Botanical Monographs 33

On the left, Kramer parrot (Psittacula krameri) eating the fruit of Livistona chinensis. On the right, bee on the staminate ﬂower of Arenga englery.

Living Together, Flora and Fauna Associated to the Palms in in the Botanic Garden The Botanic Garden of the Universitat de València is a mature garden, a large green oasis in the city, brimming with life. Its palms and large trees give food and shelter to many animal species, providing ideal living conditions. When the palm trees bloom, they attract many animals with their fragrance and nectar: bees, hornets, wasps, ﬂies, beetles, weevils and many others that act as pollinators. Birds do not miss out, and attracted by the colours of the fruit and fructiﬁcations, they are excellent dispersal agents for palm trees. The garden is visited daily by parrots and parakeets, thrushes, doves and many other birds. Some mammals, such as squirrels, rats or mice, often choose the crown of the palm to live in and reproduce there. Living on the palm trees there are also lichens, ferns, mosses, vines and other plants that colonize their stems and crowns.

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The Phoenix dactylifera crown is tied to prevent the new leaves from synthesizing. The white palm leaves, Palma blanca, are used for religious processions on Palm Sunday. Daya Vieja (Alicante)

Uses and Exploitation Palms are very important for humans living in a large part of the planet. They provide sustenance and various raw materials of utmost necessity to millions of people. Their uses and applications are countless. The palm trees that have successfully spread across the globe for their economic and commercial interests are the coconut palm (Cocos nucifera), the date palm (Phoenix dactylifera) and the Palmyra or sugar palm (Borassus flabellifer). The whole of the coconut palm is used, but its fruit, the coconut, is the product that generates most benefits. The date palm gives us dates, but the rest of the plant is also used in one form or another. From the Palmyra palm or black palm all components are also exploited to varying degrees; notably the fruits, with edible and rich pulp, but even more important is the sago flour extracted from the stem. There are some palm trees that are less known and widespread, which also have significant economic value. For instance Areca catechu, from whose seeds we obtain the areca nut, which wrapped in betel leaf (Piper betle) together with

Botanical Monographs 35

To the left, date collection of Phoenix dactylifera, Ghardaia (Algeria). On the right, shelter roofed with palm leaves, Rio Napo (Ecuador).

lime and spices, serves as a masticatory pill with medicinal and stimulating properties. The so-called oil palm, Elaeis guineensis, originating from Africa, and Elaeis oleifera, from Central America, provide oil from the fruit’s pulp, used for domestic consumption. Some species have a more restricted use. The roots of Bactris gasipaes are useful as antiparasitic agents, while Borassus have diuretic and vermifuge applications. The stem of many species is used for construction, in the form of posts, beams, planks and ﬂooring. Other palms, such as Calamus, have a thin and ﬂexible stems that are used to manufacture furniture known as rattan. There are palms, such as Ceroxylon quindiuense, with waxy stems, used to manufacture candles and make various cosmetic preparations. The terminal bud or “palmito” of many species is edible and is usually eaten fresh in salads or cooked as an accompaniment to many dishes.

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The sap produced by palms contains many sugars, which have traditionally been extracted for human consumption. This happens for example with Arenga, Borassus, Caryota, Jubaea or Phoenix, whose terminal buds and inflorescences are cut, letting the sap to flow out. This sap, when properly fermented, produces palm wine. The leaves also have multiple uses. The very tender ones of some species like Coccothrinax argentea can be consumed as vegetables. In Spain, the leaves of Chamaerops humilis and Phoenix canariensis have been used to make a variety of handicrafts, such as hats, baskets, brooms and even ties. Young leaves of Phoenix dactylifera which have been deprived of light for a few months (a practice known as “encaperuzado” or “hooding”) turn white and artisans in the town of Elche twist them for the Easter processions. In general the leaves can be used to construct roofs, as writing surfaces, to make hats and mats, to fashion traps for fish and small mammals or construct border fencing, among many other uses. A lot of palms produce edible fruit and seeds, which can be eaten fresh or made into jams and liqueurs. Serenoa repens seeds have chemical properties used to combat prostate cancer, benign prostatic adenoma.

Palms and Climatic Conditions Wind Palm trees in general, due to their root system and stem structure, can withstand high winds, hurricanes and tornadoes much better than other trees. But the wind can exacerbate dryness in plants during hot weather, causing them to dehydrate and even burn the tips of some pinnae. The same thing occurs with cold: the wind causes higher penetration of low temperatures, increasing frost damage. In locations near the sea, the salty air damages the leaves of palm trees and plants in general.

Sun The sun, as for all plants, is essential for the palm’s life. However, the extent, intensity and duration of exposure are either life-giving, conversely, lifethreatening, depending on the kind of species and the stage in the life cycle. Some palms grow perfectly from seedlings to adults in full sun, others need partial shade in order to grow and develop during the juvenile stages and their demand for more sun increases as they become adults. Some species grow in the understory of tropical forests, where little light reaches them.

Botanical Monographs 37

On the left, wind and salt cause damage. Phoenix dactylifera. On the right, damage caused by lightning to the stem of Washingtonia robusta.

Storms and lightning Some species such as Washingtonia robusta, sometimes exceeding 35 meters in height, are vulnerable to storms and can be damaged in heavy gales. Lightning can cause severe damage in some tall palm trees. In areas with frequent thunderstorms, it is important to install lightning rods protruding above the canopy of large palm trees, in nearby buildings or on the tree itself.

Fire Most palms, but especially species that live in open and exposed forest thickets or savannas, are well adapted to fire. Their vascular bundles are well protected, and their terminal bud is hidden inside the stem. However, they are not invulnerable to intense and prolonged flames. Palm trees near fields suffer damage from agricultural burning, and those planted in cities do not escape fires caused by accidents or acts of vandalism.

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On the left, damage caused by low temperatures on the leaves of Pritchardia hillebrandii. On the right, vandalism, ﬁre in the stem and crownshaft of Washingtonia robusta.

Cold Outside the tropics, the cold and frost are a concern for gardeners and palm growers. The resistance of each species is very diverse, depending on both genetic and environmental factors. There are many species that support moderate frost, down to -5 °C, such as Livistona australis. There are few species able to withstand lower temperatures, including Butia capitata, which can survive temperatures of -9 °C. But some can resist even lower temperatures: in extremely cold climates Trachycarpus fortunei or Rhapidophyllum hystrix can be planted, as they withstand freezing conditions of -20 °C. The altitude at which the various species grow in the wild gives us clues about the hardiness of the species to withstand cold. Some palm trees in the tropics live in mountainous areas at considerable altitudes, withstanding temperatures much lower than their counterparts near the sea. For a proper selection of palms, one must choose the specimen carefully, since within the same genus each species can have marked differences in their tolerance to cold. Generally, the warmest and most sheltered areas of the garden should be

Botanical Monographs 39

Excessive pruning, Phoenix canariensis.

kept for those species most sensitive to low temperatures, which must also be protected adequately during the winter. When palm trees have been damaged by cold, the bud may have survived despite all the leaves having died. In this case, the palm will sprout again when conditions are favourable and generate new leaves. Branched species may also suffer severe damage during a freeze. Consequently, one must always wait a while after frost to see whether the palm possibly recovers. The parts of the palm tree most sensitive to cold are the roots, leaves and the terminal bud. In addition to direct cold resistance, other growth conditions should be controlled. Specimens that show deďŹ ciencies or suffer from pests and diseases are less resistant to low temperatures than healthy individuals. Pruning is another aspect that can negatively affect palmsâ&#x20AC;&#x2122; resistance in the case of a frost. Dried leaves adhering to the stem are a very effective system for maintaining a constant temperature inside. Pruning, the way it is currently performed (removing several crowns of green leaves, inďŹ&#x201A;orescences and infructescences) adversely affects palms, as eliminating functional leaves means reducing overall protection as well as the photosynthetic capacity of the plant. These plants will become more vulnerable, weak and sensitive to frost.

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Genus Data Sheets

Genus Data Sheets 1. Acoelorrhaphe H. Wendl. 2. Adonidia Becc. 3. Aiphanes Willd. 4. Allagoptera Ness 5. Archontophoenix H. Wendl. & Drude 6. Areca L. 7. Arenga Labill. 8. Bismarckia Hildebrandt & H. Wendl. 9. Brahea Mart. ex Endl. 10. Butia (Becc.) Becc. 11. Caryota L. 12. Chamaedorea Willd. 13. Chamaerops L. 14. Chambeyronia Vieill. 15. Coccothrinax Sarg. 16. Copernicia Mart. ex Endl. 17. Dypsis Noronha ex Mart. 18. Gaussia H. Wendl. 19. Howea Becc. 20. Hyophorbe Gaertn. 21. Jubaea Kunth 22. Latania Comm. ex Juss. 23. Licuala Thunb. 24. Livistona R. Br. 25. Nannorrhops H. Wendl. 26. Phoenix L. 27. Pritchardia Seem. & H. Wendl. 28. Ptychosperma Labill. 29. Ravenea BouchĂŠ ex H. Wendl. 30. Rhapidophyllum H. Wendl. & Drude 31. Rhapis L. f. ex Aiton 32. Sabal Adans. 33. Serenoa Hook. f. 34. Syagrus Mart. 35. Thrinax L. f. ex Sw. 36. Trachycarpus H. Wendl. 37. Trithrinax Mart. 38. Washingtonia H. Wendl. 39. x Butyagrus nabonnandii (Prosch.) Voster.

Botanical Monographs 43

Acoelorrhaphe H. Wendl. Its name comes from the Greek a-koilos-raphĂŠ and indicates that the seeds have no groove, seam, or raphe. Genus contains a single species, Acoelorrhaphe wrightii.

Distribution and ecology It is native to the southern United States (South Florida Everglades). Optimum habitat in the CaribeoMesoamerican Region, as it extends through Bahamas, Cuba and northwest of Central America. It lives in areas close to the coast, scrub savannah and coastal forests of Pinus elliottii and Pinus palustris.

Description Small palm, with multiple stems forming large groups or clumps, occasionally solitary. Stems slender, straight or inclined. Crown small, globose, with leaves remaining on stem for quite some time after drying. Leaves small, webbed or slightly costapalmate, with bright green upper leaf and pale green or glaucous undersides. The margins of the petiole are yellowish with red border and are covered with small triangular and curved spines. The genus is monoecious and polycarpic. The inďŹ&#x201A;orescences sprout between the leaves and are longer than them, they are yellow or orange, branching to up to four orders. Flowers small, bisexual, sessile and creamy-white, solitary or in groups. The fruit is a globose berry, glossy black when ripe.

Uses and exploitation The stems are used to make hedges and fences. Home roofs and brooms are made with the leaves. It can be widely used in gardening.

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Adonidia Becc. Its name refers to the red colour the fruit acquires when ripening, like the drops of blood shed by dying Adonis. A single species: Adonidia merrillii.

Distribution and ecology Paleotropical and Indo-Malaysian (Philippines and Malaysia) genus. It grows on various soil types and extends from the coast to the foothills. Palm classiﬁed at low level of extinction risk, threatened by deforestation of its habitat.

Description Middle-sized palm, single stem and unarmed. The stem has a flared base, is straight, slender, smooth, ringed, banded, striated longitudinally and an ornate crownshaft in its distal area. The crown is small, hemispherical. Leaves pinnate, arching, green on both sides, and shed quickly once dry. Clasping sheath, bright green, with grey hues from the wax and tomentum covering it. Pinnae linearlanceolate, reduplicated, apex pendulous with sharp termination or obliquely truncated and toothed. The genus is monoecious and polycarpic. Inflorescences sprout below the last sheath and branch up to three or four orders. The flowers are arranged forming triads at the base, paired or solitary towards the apex. They are small, unisexual and sessile. The staminate ones are green, and the pistillate yellow-green and melliferous. The fruits are ovoid berries, bright red when ripe.

Uses and exploitation Sensitive to cold, palm widely used in gardening in warm tropical and subtropical areas.

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Aiphanes horrida

Aiphanes Willd. The name comes from the Greek aeiphanes, which means always visible because of its bright red fruits. Other authors indicate that the name means jagged, from the uneven shape of the pinnae apex. Twenty-four species are recognized, of which Aiphanes horrida grows in the Botanic Garden.

Distribution and ecology Neotropical genus with a wide distribution in the Caribbean and in northern and central South America. It inhabits rainforests, forming part of the undergrowth and lives under the protection of the canopy. It ranges from sea level up to 2,800 m altitude.

Description Small to medium-sized palms, with numerous spines on stems, leaves and inďŹ&#x201A;orescences. Stems slender and ringed, which may be solitary, multiple or acaulescent. Crown irregular, sparse, with leaves that are shed when dry. Leaves pinnate except Aiphanes macroloba, with whole leaves. Broad pinnae, green on both sides, reduplicated, with truncated and premorse apex. Genus is monoecious and polycarpic. InďŹ&#x201A;orescences are interfoliar and shorter than the leaves, usually branched up to one order, but there are unbranched species or branched up to two orders. Flowers unisexual, small, generally forming triads towards the base of the rachilla, with only staminate male ďŹ&#x201A;owers, paired or solitary at the apex. The fruits are globose drupes bright red when ripe, with three characteristic pores located in the central area.

Uses and exploitation The apical bud and fruit of some species are edible. Aiphanes horrida and Aiphanes minima seeds are used in the preparation of sweets and pralines. Palms are seldom used in gardening because of the many thorns that protect them.

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Allagoptera arenaria

Allagoptera Neess. Its name comes from the Greek allage, meaning different, and pteron meaning plume, referring to the arrangement of the pinnae on the leaf. The genus is comprised of ďŹ ve species, of which Allagoptera arenaria grows in the Botanic Garden.

Distribution and ecology This genus is Neotropical and extends through east and central South America. It inhabits areas near the Atlantic coast, on beaches and sand dunes, in dry forests or near the shore. Part of the cerrado vegetation up to 1,500 m altitude.

Description Small-sized palm trees, with or without spines on the petiole, with single or multiple stem. Stem subterranean, acaulescent, somewhat prostrate or creeping, less often straight and slender, sometimes double branched. Initially covered by residual sheaths, stalks and ďŹ bres, and later becomes cylindrical, rough and ringed. Crown small, globose with marcescent leaves. Very characteristic pinnate leaves, upper blade green or bluish-green, underside waxy, bluish, greyish or silvery; pinnae long, linear-lanceolate, which are arranged along the rachis regularly or in small groups and forming different planes. The genus is polycarpic and monoecious. InďŹ&#x201A;orescences sprout solitary between the leaves, are shorter and are not branched. Flowers unisexual, the pistillate are sessile and globose, the staminate, ovoid-obovoid and pedicellate. The fruits are an obovoid drupe, yellowish-green or brown, with three pores near the base.

Uses and exploitation The fruits and seeds of some species are edible. They are little known ornamental palms in Spain, despite their vast potential use in gardening and landscaping in warm coastal areas.

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Archontophoenix alexandrae

Archontophoenix H. Wendl. & Drude The name comes from the Greek word archon, meaning chief or ruler, and phoenix, meaning palm, in allusion to its majesty. Six species are recognized, of which Archontophoenix alexandrae and Archontophoenix cunninghamiana grow in the Botanic Garden.

Distribution and ecology This genus is endemic to eastern Australia. It grows in wetlands from sea level up to 1,200 m altitude. It also grows in valleys and ravines, near streams.

Description Palms of medium to large size, single stem, unarmed. Stem with swollen base, cylindrical, slender, smooth, ringed or stepped, cracked lengthwise and with the apex crowned by an ornate crownshaft. Crown hemispheric, with leaves that are shed when dry. Leaves pinnate, with long sheaths, clasping, tubular, cylindrical and swollen at the base; green and sometimes covered by tomentum of different colours. Pinnae lanceolate, apex reduplicated and irregularly sharp, green on both sides, such as Archontophoenix cunninghamiana; with grey-silvery scales on both sides, like in Archontophoenix purpurea, or only on the undersides in the other species. It is a monoecious and polycarpic genus. The inďŹ&#x201A;orescences are infrafoliar, composite and generally branched up to four orders. Flowers unisexual, sessile, creamy white to purple. The fruits are elliptic-globose berries, pink to deep red when ripe. The species of this genus hybridize easily.

Uses and exploitation The apical bud of some species is edible. Due to their beauty and elegance, they are widely used in landscaping, adding a touch of class wherever planted.

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Areca triandra

Areca L. The generic name is a Portuguese adaptation of the term areec, name used in the Malabar Coast (India) to name these palms. Some forty-seven species are recognized, of which Areca catechu and Areca triandra grow in the Botanic Garden.

Distribution and ecology Paleotropical genus, Indo-Malayan. Most species live in the understory of rainforests. Some live in open areas (Areca catechu); others thrive along waterbeds (Areca rheophytica) as they are able to withstand water currents.

Description Small to medium-sized palms, unarmed, acaulescentscent, or with single, multiple stem. Some species show aerial roots, as Areca triandra; sometimes stilt-rooted, as Areca vestiaria. Stem cylindrical, slender, ringed, greenish, at least in young parts; stem has an ornate crownshaft, more or less attractive, especially in Areca vestiaria due to its orange sheaths. Crown small, sparse, with leaves that are a shed when dry. Leaves pinnate, or whole with pinnate venation, pinnae highly variable. Genus monoecious and polycarpic. The inﬂorescence usually sprouts in the leaf axil, but does not develop until the latter dies and falls, giving the appearance of being infrafoliar, branching up to three orders. The ﬂowers are unisexual, sessile and fragrant. The pistillate are globose and larger than the staminate. The fruits are ovoid-globose berries, orange, or bright red when ripe.

Uses and exploitation The palmetto (palmito) or terminal bud of certain species and the fruits are edible. The leaves are used for making baskets, and inﬂorescences as decoration in religious ceremonies. Areca catechu is used in traditional medicine, rituals and ceremonies. The fruits have great economic value for their parasympathomimetic and anthelmintic properties. In Southeast Asia it is chewed as mastic for its antiseptic and anti-halitosis beneﬁts.

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Arenga engleri

Arenga Labill. in DC. The name comes from aren, the native name this genus is known by in Java. It encompasses some twenty species, of which only Arenga engleri grows in the Botanic Garden.

Distribution and ecology Genus originally native of tropical and subtropical Asia, reaching as far as New Guinea and northern Australia. Its current distribution range is Southeast Asia, ranging from India to northern Australia. Many species are part of the primary or secondary forests in lowlands or in mountains, some may grow at 2,150 m altitude.

Description Palm trees of varying size, unarmed or a little spiny. Stem often multiple, solitary, or acaulescent, which usually remains covered by residual sheaths and petioles. Crown small, with pinnate leaves, occasionally whole and marcescent. The sheath and petioles often have very ornamental tomentum and scales. Pinnae induplicate, very variable in shape and size, with the apex irregularly toothed; green on both sides such as Arenga australasica or green upper blade and grey hues on the underside, such as Arenga obtusifolia. Monocarpic and mostly monoecious genus, but with dioecious species; others with multiple stems, such as Arenga englery, may fructify several times before dying. InďŹ&#x201A;orescences are interfoliar, solitary, unisexual, pendulous, branching to one or two orders, occasionally infrafoliar, multiple, bisexual and unbranched. Staminate ďŹ&#x201A;owers large, fragrant, with free sepals and numerous stamens. The fruits are berries, globose-ellipsoid, yellowish, reddish or purplish.

Uses and exploitation The terminal bud is edible in some species and trunks are used as beams and supports in construction. The leaves are used for weaving various objects. The stem of Arenga pinnata is used to extract starch. Wine is obtained by distilling the sap of the terminal bud, and sugar is made from the inďŹ&#x201A;orescences (sugar palm).

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Bismarckia Hildebr. & H. Wendl. Name dedicated to Otto Von Bismarck, First Chancellor of Germany. It comprises a single species: Bismarckia nobilis.

Distribution and ecology It is endemic to the island of Madagascar (Malagasy Region), where it is distributed along pluviseasonal tropical climates. It characterizes the dry savannah vegetation, and may occasionally be found in wetlands and ﬂooded plains.

Description Medium to large-sized palm tree, with single and unarmed stem. Stem with swollen base, straight, brownish-grey, rough, ringed and striate with longitudinal grooves. Crown globose, dense, leaves marcescent in juveniles and in adults deciduous. Leaves large, costapalmate, leathery, waxy, green, blue or grey on both sides. Lamina orbicular, divided halfway or slightly less of its length in various leaf segments, which are lanceolate, induplicate, leathery, apex cleft and acute terminations. In the division point of the segments bright fibres are observed, long, thin and curved. It is a dioecious and polycarpic species. Inflorescences similar in both sexes, interfoliar, solitary, shorter than the leaves and branched up to two orders. Male flowers ovoid in bud, purplish; the pistillate globose, shortly pedicellate and green. The fruits are drupes, ovoid-globose, dark brown when ripe.

Uses and exploitation The stems are used as columns and beams in construction, also in pens and fences. Leaves are used for roof thatching and to produce several types of baskets. Species widely used in gardening.

Botanical Monographs 59

Brahea armata

Brahea Mart. ex Endl. Genus dedicated to the Danish astronomer Tycho Brahe. Some ten species are recorded, of which Brahea aculeata, Brahea armata, Brahea brandegeei, Brahea edulis and Brahea sarukhanii grow in the Botanic Garden.

Distribution and ecology Neotropical genus of Mexican optimum, spreading to Central America (Caribeo-Mesoamerican). It grows from sea level to over 2,000 m altitude and inhabits mountain slopes, canyons and ravines, on limestone soils or soils of igneous origin.

Description Small to medium-sized palms, with slow growth and armed petioles. Stem solitary, except Brahea dulcis and Brahea decumbens, in which it can also be multiple; usually remains protected by residual sheaths, petioles and fibres, which are eventually shed, leaving rough, slightly ringed and striate surface. Leaves palmate or slightly costapalmate and marcescent. Lamina almost circular, divided halfway or more in various segments, with the division point bearing various filaments. Segments induplicate, more or less rigid, with the apex deeply bifid and acute terminations. It is a monoecious and polycarpic genus. Inflorescences are interfoliar, solitary, equal or longer than the leaves and branched up to four orders. Flowers small, solitary or in small groups, bisexual, sessile, yellowish-white, often tomentose. The fruits are berries, globose or ovoid, brown, dark blue, or black when ripe.

Uses and exploitation The stems are used as columns, beams or supports. Leaves are used in rooﬁng, construction and braided to make rope, hats, mats and toys. The apical bud of some species is edible and alcohol is obtained from processed sap. The fruits of some species are edible and fresh ground seeds are also consumed as food.

Botanical Monographs 61

Butia capitata

Butia (Becc.) Becc. The name originates from the Portuguese adaptation of the indigenous Brazilian term mbo-tiá, curved tooth. About nine species are recognized, of which Butia capitata grows in the Botanic Garden.

Distribution and ecology This Neotropical genus of BrazilianParanaense and Pampean origin, has a distribution that spans from southern Brazil to north-eastern Argentina and northern Uruguay. It constitutes the communities of the cerrados. Populations are being depleted due to crop encroachment.

Description Small to medium-sized palm trees, slow growing, armed or unarmed. Stem solitary, multiple, or subterranean, usually covered by the bases of the leaves, which are eventually shed, leaving characteristic scars. Leaves pinnate, arching, and in some species blue-green or grey. Pinnae numerous, linear, reduplicate and rigid, with the apex acute, acuminate or asymmetrical. It is a monoecious and polycarpic genus. The inﬂorescences sprout between the leaves, are protected by a prophyll and a woody, tubular bract, smooth or slightly striate, glabrous, or, as in Butia eryospatha, tomentose; they are shorter than the leaves and are divided up to one order. Flowers unisexual: the staminate ﬂowers have six stamens, the pistillate are ovoid-globose and larger. The fruits are drupes, ovoidglobose, yellowish to brown-purple when ripe, with three characteristic pores.

Uses and exploitation The leaves are used for rooﬁng, or are braided into baskets, nets and hats. The fruits are eaten fresh or they are made into jellies, jams, ice cream, liquor and oil. The endosperm of the seeds (“coconuts”) is eaten fresh or dried. From the sap honey and palm wine are made. Inﬂorescences are cut to obtain sugar from their sap. Also palm wine is obtained by macerating the fruits in alcohol.

Botanical Monographs 63

Caryota urens

Caryota L. The name comes from the Greek word karyon, meaning nut. There are thirteen species in this genus, two of which are found in the Botanic Garden: Caryota mitis and Caryota urens.

Distribution and ecology The genus’ distribution is Paleotropical, mainly Indo-Malayan, from India to northern Australia. They are part of the primary forest, growing both at sea level and in mountainous areas, 2,000 m maximum altitude.

Description Fast growing palm, unarmed. Abundant and shallow roots, adventitious occasionally. Stem solitary or multiple, columnar; remains covered by sheaths and fibres, eventually shed leaving visible rings. Leaves are large, very characteristic, bipinnate and green on both sides. Pinnae with various shapes like a fish tail, induplicate, slightly pendulous, with the upper margin undulated, deeply praemorse. It is a monoecious and monocarpic genus. Inflorescences solitary, spring between the upper leaves, are shorter than these, generally pendulous, branching up to an order. Usually produce several inflorescences for several years, sprouting beneath the leaves and when final fruiting occurs the palm or stem dies. Flowers grouped in triads, sessile and unisexual. The fruits are globose berries that turn orange, red or purple when ripe.

Uses and exploitation The terminal bud is edible and stem pith is extracted to make a food like sago. The wood is very hard and resistant, and thus used widely. The leaves have very durable ﬁbres, the “Kitul”, which the indigenous people use to make ropes, baskets and other household utensils. Before opening, the inﬂorescences are bled to extract the sap due to its high sugar content. The fruit contains abundant calcium oxalate crystals, toxic if swallowed and irritant if touched.

Botanical Monographs 65

Chamaedorea microspadix

Chamaedorea Willd. The name of this genus comes from the Greek word khamaí, which means ground or creeper, and doreá, meaning gift, in allusion to the ease in collecting their fruits. One hundred and ten species are recognized, of which Chamaedorea cataractarum, Chamaedorea costarricana, Chamaedorea elegans, Chamaedorea metallica, Chamaedorea microspadix, Chamaedorea oblongata, Chamaedorea potchutlensis, Chamaedorea radicalis and Chamaedorea seifrizii grow in the Botanic Garden.

Distribution and ecology Its distribution is Neotropical, from central Mexico to Peru and Bolivia. It grows both in the Atlantic and in the Paciﬁc, from sea level to 2,000 m altitude. It is part of the tropical rain forest understory or lives in periodically ﬂooded areas. Some species of this genus have spread worldwide as houseplants.

Description Small-sized, fast-growing palm, unarmed. The root system is usually broad and widespread, in some species epigeic roots are observed. Stem is multiple, solitary or acaulescent, very thin, green, covered with clasping sheaths, or ornately ringed; may grow straight, inclined or prostrate, be a creeper as Chamaedorea elatior or have dichotomous branching such as Chamaedorea cataractarum. Leaves are usually pinnate, biﬁd, or occasionally whole with pinnate venation. It is a dioecious and polycarpic genus. Inﬂorescences spring between or below the leaves, solitary or multiple, unbranched or branched one or even two orders. Flowers similar in both sexes, small, unisexual and sessile. The fruits are oblong-globose berries, which hold a single seed.

Uses and exploitation Decorative uses such as gardening and landscaping, with many applications especially in small spaces such as gardens or patios and terraces. Also used as a potted houseplant.

Botanical Monographs 67

Chamaerops L. The name comes from the two Greek words: khamaí, which means low, dwarf or “that touches the soil”, and rhops, meaning bush, coppice. The genus comprises just one species Chamaerops humilis, although numerous varieties are grown at the Botanic Garden such as C. humilis and C. argentea, with some specimens exceeding 100 years in age.

Distribution and ecology Chamaerops humilis is a relict species of the western thermomediterranean ﬂoor. It cannot withstand frost, but exceptionally it can grow at the 2,400 m altitude in the Moroccan High Atlas mountains. Part of the maquis, garrigue and xeric shrublands. Traditional uses of the plant have reduced its natural populations, so it is now protected.

Description Palm highly variable and high morphological plasticity, tufted, with multiple stems. Stems slender, straight, slightly inclined or even creeping. Palmate leaves, leathery and waxy, with margins of petiole armed strong spines. Lamina semicircular to circular, dark green, glaucous or grey, but can also present various tomentum, especially on the underside; is deeply divided into several segments that are linear, induplicate, rigid and deeply forked. It is a dioecious and polycarpic species, occasionally bisexual ﬂowers. The inﬂorescences sprout between the leaves, are erect and very short, branching up to two orders. Flowers small, yellow. The fruits are berries, ovoid-ellipsoid, yellowish-red, becoming brown when mature.

Uses and exploitation Its rich and nutritious apicial bud has been consumed since ancient times. A drink similar to sarsaparilla is made from its roots. With the ﬁbres of its leaves baskets, brooms, mats or strings can be made; they have also been used for rooﬁng buildings. The fruit, called palmiches or Raposa dates, are small and sweet when ripe. The pulp is rich in tannins, being used in traditional medicine as an astringent.

Botanical Monographs 69

Chambeyronia macrocarpa

Chambeyronia Vieill. Genus dedicated to Charles Chambeyron, captain of the French navy. It comprises two species, of which Chambeyronia macrocarpa grows in the Botanic Garden.

Distribution and ecology This genus is endemic to New Caledonia, in the Australian region, with pluviseasonal tropical climate. It lives in tropical rainforests and gallery forests from sea level to about 1,600 m above sea level, on various soil types.

Description Medium-sized palm, solitary stem, unarmed. Stem base slightly thickened, straight, slender, ringed; at its upper end, the sheaths form a colourful crownshaft, green or greenish-blue, covered with indumentum of different colours. Crown hemispheric with leaves that are shed when dry. Pinnate leaves, slightly arched at the time of opening and for a short period they are a striking red colour, becoming green later. Pinnae large, lanceolate, reduplicated, leathery, green on both sides and with the apex acuminate or acute. It is a monoecious and polycarpic genus. The inďŹ&#x201A;orescences sprouting beneath the leaves are composite, with pendulous branches and branched up to two or three orders at the base and only one towards the apex. Flowers unisexual, sessile, pink or creamy. The fruits are berries, subglobose or ovoid, which turn reddish when ripe.

Uses and exploitation They are very ornamental palms, with ample opportunities in gardening. In Spain they are virtually unknown and are usually found only in botanical collections.

Botanical Monographs 71

Phoenix dactylifera Coccothrinax crinita

Coccothrinax Sarg. The name originates from the Greek kókkos which means seed, grain or berry, and thrînax, trident or trifurcated. Some ﬁfty species are known, of which only Coccothrinax crinita grows in the Botanic Garden.

Distribution and ecology The species of this genus are mostly Neotropical, CaribeoMesoamerican. They live in open areas near the coast, seasonally ﬂooded savannas, high or low mountain forests, both as shrubs and tree species. They thrive on limestone soils, rocky or sandy, or magnesiumrich serpentine soils.

Description Slow-growing palm, stem solitary, unarmed or lightly armed in their sheaths. The stem is usually covered by sheaths and ﬁbres, which can be long and soft like a mane, woven like a mesh, or spiky. When they are shed they reveal a cylindrical stem, thin, smooth and striate; however, some species show a thickening at the base or at a certain height, such as Coccothrinax spissa. Leaves palmate and marcescent. Leaf is obconical to circular, divided halfway into several segments, green and glabrous on the upper blade, with the underside usually silver-grey and sometimes tomentose. It is a monoecious and polycarpic genus. Inﬂorescences sprout between the leaves and are generally shorter than these and branching up to two orders. Flowers bisexual, solitary and pedicellate. Fruit small, globose or subglobose, ranging from red to purple, becoming black when ripe.

Uses and exploitation In its countries of origin, the stem is used as timber. Brooms and ropes are made with the leaves, and they are also used for rooﬁng due to their strength and durability. In Spain, these palms are known only in some botanical collections, usually grown in greenhouses.

Botanical Monographs 73

Phoenix dactylifera Copernicia alba

Copernicia Mart. ex Endl. Genus dedicated to the Polish astronomer Nicolas Copernicus. Twenty-one species are recognized, of which only Copernicia alba grows in the Botanic Garden.

Distribution and ecology The genus is primarily located in Cuba, Dominican Republic and Haiti (Caribeo-Mesoamerican Region), but extends throughout much of South America. Generally inhabits xeric tropical climates. It grows in open areas near the coast, in the caatinga, seasonally ﬂooded savannah and open woodlands.

Description Stem solitary, occasionally multiple, with petioles armed with strong curved spines. Stem straight, slender or thick, and in some species with very characteristic bumps or swellings. Crown usually globose, marcescent leaves in young individuals and deciduous when mature. Leaves costapalmate or slightly palmate, green or with blue-grey and waxy undersides. Lamina obconical to circular, split up to a quarter or a third of its length into various segments, which are induplicate and rigid; they may have small spines on the margins and have the apex biﬁd with sharp ends. It is a monoecious and polycarpic genus. The inﬂorescences sprout between the leaves, are usually longer than them and branched up to six orders. Flowers bisexual, sessile and ovoid in bud. The fruits are ovoid-globose berries, black when ripe.

Uses and exploitation The stem of some species have dense and resilient wood, and therefore used in rural construction and for making poles, posts and fences. The young leaves are used as fodder for livestock, while from the adult wax and ﬁbres are extracted to make rope, baskets or hats. The fruits are an important source of food for wildlife, especially birds, which disperse the seeds.

Botanical Monographs 75

Phoenixlutescens Dypsis dactylifera

Dypsis Noronha ex Mart. The name derives from the Greek word dýptein, which means to descend or go down, normally referring to the head. Some hundred and forty species are known, of which only Dypsis lutescens grows in the Botanic Garden.

Distribution and ecology This genus is Paleotropical, mainly Malagasy. It grows in different types of habitats, such as rainforests, open plains or dunes, from sea level to about 2,000 m altitude. Some species are endangered and others seriously threatened due to disturbance of the regions they inhabit.

Description Palm of single, multiple or subterranean stem, always unarmed. The stem is cylindrical, or tapering towards the apex, grows straight, inclined or prostrate; some species have curious bumps, others have ramifications, are creepers or rheophytic. Leaves pinnate, entire or bifid with pinnate venation, usually green on both sides; cylindrical sheaths that form an ornate crownshaft at the top of the stem, which is glabrous, sometimes waxy, or with indumentum of different colours; there are also species with bulky sheaths, arranged in three rows, forming noticeably triangular crowns such as Dypsis decaryi. Pinnae usually linear-lanceolate, reduplicate, with acute termination, bifid, or occasionally truncated and praemorse. It is a monoecious and polycarpic genus. Inflorescences sprout between the leaves or below these, such as Dypsis decipiens. Flowers arranged in triads, sessile and unisexual. The fruits are globose-ellipsoid berries, usually brightly coloured, brown or black.

Uses and exploitation In some species the apical bud is edible, in others it is bitter or toxic. The stems are used in the construction of houses; those of some species, hollowed out, serve as a blowgun. The leaves are used for rooﬁng buildings, making boxes, brooms and ropes. The fruits of some species are sweet and eaten fresh, but can also be fermented to obtain alcohol.

Botanical Monographs 77

Gaussia maya

Gaussia H. Wendl. Genus dedicated to the astronomer and mathematician Carl Friedrich Gauss. Five species are known, two of which, Gaussia maya and Gaussia spirituana, ďŹ&#x201A;ourish in the Botanic Garden.

Distribution and ecology Neotropical pluviseasonal genus with Caribeo-Mesoamerican distribution. Gaussia maya grows in Mexico, Guatemala and Belize, while Gaussia spirituana is typical of Cuba. These palm trees live in forests on steep ridges, rocky areas at low altitude, on limestone soils. Because of its edapho-xerophilous nature it often grows among the Mayan ruins, especially in Yucatan and Guatemala. Some species are included in the red list of endangered extinction-threatened species, due to deforestation of the habitats they occupy.

Description Small to medium-sized palms, stem single and unarmed. The stem base can be bulky and covered in roots, or show swellings at a certain height; grows straight or tilted, is sleek, smooth and ringed, cylindrical or tapering towards the apex. Crown hemispherical or irregular. Leaves pinnate, deciduous when dry, from which only the pinnae are shed with the rachis remaining. Sheath embracing, broad at the base, not forming a crownshaft. Pinnae linear-lanceolate, base swollen, reduplicate and apex acute, with upper blade green, with lighter and waxy undersides. Monoecious and polycarpic genus. The inďŹ&#x201A;orescences sprout between the leaves, are shorter than these and branch up to 1-3 orders. Flowers small, unisexual, sessile, green when buds, yellow when open. The fruits are globose berries or kidney-shaped, ranging from orange to bright red when ripe.

Uses and exploitation The stems are used in construction as pillars and posts as they are sturdy and durable. The fruit pulp is caustic and an irritant. They have no outstanding ornamental characteristics and are virtually unknown in Spain.

Botanical Monographs 79

Phoenixforsteriana Howea dactylifera

Howea Becc. The name refers to the island of Lord Howe, baptised in honour of the British admiral Richard Howe. Beccari proposed the name Howeia for this beautiful palm, but it was ﬁnally named Howea. Two species are known, Howea belmoreana and Howea forsteriana, both of which grow in the Botanic Garden.

Distribution and ecology This genus is endemic to the small island of Lord Howe, located in the Paciﬁc Ocean, east of Australia (Tropical-Australian Region). The island is a special enclave with a high rate of endemic palms. Howea belmoreana inhabits humid forests in mountainous enclaves, and may grow at the 400m altitude, on basalt soils. Howea forsteriana grows in the forests of the lowlands, especially in developing coral and calcareous soils. These palms are included in the red list of endangered species, classiﬁed as vulnerable, due to overexploitation of their seeds in natural populations.

Description Medium to large-sized palms, stem single and unarmed. Stem with thickened or flared base, straight, slender, cylindrical, smooth and ringed; initially dark green then becoming grey. Crown hemispheric, with pinnate leaves of dark green and lighter undersides, reduplicate in Howea forsteriana, induplicate en Howea belmoreana. Sheaths clasping, swollen at the base, not forming a crownshaft. The pinnae are linear-lanceolate, reduplicate, apex acuminate or acute and the central nerve of the underside covered in dark ramenta. It is a monoecious and polycarpic genus. The inflorescences sprout between the leaves and are somewhat shorter than these; Howea belmoreana has a solitary spathe, while in Howea forsteriana eight spathes can sprout up from the same point. Flowers unisexual and sessile, the staminate bloom first, which are ovoid in bud, and later the pistillate, which are globose. The fruit is an ovoid-ellipsoid berry, which turns brown-red when ripe.

Uses and exploitation The inhabitants of the island of Lord Howe live mostly from collection, production and sale of this palm, exporting thousands of them a year. When juvenile, it is an elegant and valued houseplant.

Botanical Monographs 81

Phoenix dactylifera Hyophorbe lagenicaulis

Hyophorbe Gaertn. The name derives from the Greek words hy^s, pig, and phorbe, food, in reference to its fruits used as fodder for pigs. Five species are recognized, of which Hyophorbe lagenicaulis and Hyophorbe verschaffeltii grow in the Botanic Garden.

Distribution and ecology Paleotropical genus endemic to the Mascarene Islands archipelago, located in the south-west Indian Ocean (Madagascar Region). Hyophorbe lagenicaulis is endemic to the island of Round, while Hyophorbe verschaffeltii is native to the island of Rodriguez. In a not too distant past they were very abundant in the mountains and valleys of these islands, but now their colonies have been drastically reduced by human occupation of their habitat.

Description Small palms, stem single and unarmed. Stem thickened at the base and bottlelike, spindle-shaped or cylindrical; ringed and ďŹ nely striated, showing at the distal part an ornate crownshaft. Crown hemispheric, with leaves that are shed when dry. Leaves pinnate, induplicate and slightly arched. Pinnae linear-lanceolate, reduplicate, arched towards the apex, termination acuminate to acute; the central nerve forms a rib on both sides, in some species of a lighter colour, with the underside covered in ramenta. It is a monoecious and polycarpic genus. InďŹ&#x201A;orescences are solitary and sprout beneath the leaves, erect and fusiform bud, later horizontal, compound and branched up to three or four orders. Flowers small, unisexual, sessile and globose. The fruits are ellipsoid-globose berries, with orange to reddish colour, or black when ripe.

Uses and exploitation Attention should be paid to the handling of the fruits, as they can cause allergic skin reactions. The seeds are edible. The genus is a very ornamental species, little known in Spain, usually only seen in private collections and botanical gardens.

Botanical Monographs 83

Jubaea Kunth. Genus dedicated to King Juba II of Numidia, ancient Berber kingdom in North Africa. Comprising a single species, Jubaea chilensis.

Distribution and ecology Genus endemic to Chile and one of the southernmost palms. It inhabits areas with Mediterranean climate Mesochilenan-Patagonian Region (Central Chile), in valleys, slopes and gullies from sea level to 2,000 m altitude. This palm is protected and classiﬁed as vulnerable in the red list of threatened species.

Description Large palm, stem long-lived and solitary. Stem with bulky base, columnar, tapering gradually towards the apex; straight, very thick, dark grey, with narrow rhomboidal brands. Crown globose and dense. Leaves large, pinnate, shed when dry. Pinnae linear, reduplicate, straight in young leaves, somewhat curved in the mature ones, apex cleft, asymmetric and with acute terminations; dark green blade, with a waxy underside, bluish-green or grey, and the central nerve covered with dark ramenta. It is a monoecious and polycarpic genus. The inﬂorescences sprout between the leaves, being shorter than these, and branch up to one order. Flowers are unisexual, dark purple, the staminate slightly pedicellate and ovoid, the pistillate slightly larger, sessile and globose. The fruits are an ovoid-globose drupe, yellow-orange when ripe.

Uses and exploitation The leaves are used for roof building, and for making mats, baskets, curtains and brooms. They are also used in religious celebrations. The fruits are eaten by livestock and the seed or “Coquito” eaten fresh or its oils extracted to make cosmetics. The sap is used to produce honey and palm wine, requiring the felling of palm trees for its extraction. They are very ornamental palms, which were introduced into European gardens in the middle of the eighteenth century.

Botanical Monographs 85

Latania lontaroides

Latania Comm. ex Juss. The name of the genus appears to come from the latinization of the word that the people of the Mascarene Islands gave to one of these palms. Three species are recognized, of which Latania lontaroides grows in the Botanic Garden.

Distribution and ecology Paleotropical genus, endemic to the Mascarene Islands archipelago (Malagasy Region). Latania lontaroides is originally from Reunion island. In the past its presence was common in the cliffs, mountains and valleys of these islands. Today, as a result of environmental degradation and the introduction of exotic plants and animals, natural populations are on the brink of extinction. Hybridization occurs easily between species of the genus.

Description Medium-sized palm, stem solitary, usually without spines. Stem with bulky base, straight, cylindrical, ringed, rough, grey or dark brown. Crown globose with marcescent leaves when juveniles and deciduous in adults. The three species show distinctive colours or various indumentum in sheaths, petioles, laminae and nerves. Leaves costapalmate, with the base of the sheath wide and Λ –like split; the lamina is divided up to a third or half of its length in various segments, which are lanceolate, induplicate, rigid, apex acute or shortly biﬁd. The genus is dioecious and polycarpic. The inﬂorescences sprout between the leaves, are shorter than these and branch up to two orders. Flowers unisexual, sessile and globose in bud, with the staminate being smaller. The fruits are drupes, large, obovoid-globose, dark when ripe; they contain three seeds, with an endocarp which is characteristic to each species.

Uses and exploitation The stems are hardwood and resistant, and consequently they are used in construction. The leaves are used for thatching houses and for making baskets and other containers. Latania lontaroides seeds are edible.

Botanical Monographs 87

Phoenixgrandis Licuala dactylifera

Licuala Thunb. It would seem that the name comes from the local expression leko wala, with which the inhabitants of the Moluccas refer to some of the species of this genus. One hundred and thirty-four species are known, of which only Licuala grandis is present in the Botanic Garden.

Distribution and ecology Paleotropical genus, widely distributed throughout the IndoMalayan region. It seems to have its genetic centre in Borneo. They are small palms that inhabit different types of rainforests. They generally live under other plants as part of the undergrowth in forests near the sea, on the edges of the mangroves or swampy bogs.

Description Small to medium-sized palms, stem solitary, multiple, or subterranean. Stem usually cylindrical, thin, occasionally with bulging base and covered in roots; can grow straight, inclined or creeping, be covered by residual sheaths and ďŹ bres, or naked with marked rings. Crown globose or irregular, with leaves marcescent. Leaves palmate in very different ways: the lamina may be entire, more or less circular, pleated, with undulated margin and shortly biďŹ d termination; or divided into deep ribs up to the base, forming various segments, more or less wide and wedge shaped. The genus is monoecious and polycarpic, occasionally dioecious. InďŹ&#x201A;orescences between the leaves, spike-shaped, or compound and branch out to three orders. Flowers small, usually bisexual. The fruits are globose berries, ovoid or ellipsoid, brightly coloured when ripe.

Uses and exploitation The thin strong stems can be used to make walking sticks, while the thicker and more resistant ones are used as posts or to build fences. The leaves are used for constructing roofs or braided to make ropes, mats and other items. They also serve to wrap up food or as a dish. The apical bud and marrow of some species are edible and highly prized. They are very widely used in ornamental gardening inside the tropics and as houseplants.

Botanical Monographs 89

Livistona australis

Livistona R. Br Genus dedicated to Patrick Murray, Baron of Livingston. Some thirty-ﬁve species are recognized of which Livistona australis, Livistona chinensis, Livistona decora, Livistona fulva, Livistona saribus and Livistona muellery are grown in the Botanic Garden

Distribution and ecology Paleotropical genus of wide distribution, although most species are concentrated in the Indo-Malayan region, reaching the Australian region. An African species, Livistona carinensis, is in Somalian-Ethiopian and Sahelian-Sudanese regions, reaching the SaharanArabic. Its ecology is very diverse: they inhabit rainforests, swamps and peatlands, as well as living in the vicinity of water beds or in places where the water table is closely available. Usually grouped together, generating more or less extensive palm groves of great beauty.

Description Palms of solitary stem, with or without spines at the base of the petiole. The stem can have a bulky or flared base, is straight, cylindrical, or tapering towards the apex; remains covered by residual sheaths and petioles and is ringed or striate, from scars left after shedding the pods. Crown generally globose with marcescent leaves. Leaves costapalmate, except palmate in Livistona exigua; the laminae is highly variable, almost circular and shortly divided, as in Livistona rotundifolia, or deeply divided, as in Livistona decora; the blade’s underside may also display colourful tomentum as in Livistona fulva. The genus is monoecious and polycarpic, occasionally dioecious. Inflorescences sprout between the leaves, are shorter than the latter and generally branched up to five orders. Flowers bisexual or functionally unisexual, of cream-yellow colour. The fruits are berries with a great variety of colours and shapes.

Uses and exploitation In some species, the apical bud and leaves are edible if unopened. The stems are used as pillars and when emptied, can serve as containers. The outer wood is very hard, dark in colour, and is used to make walking sticks or spear points. The leaves are used for roofs and can also be twisted to make rope, hats and mats.

Botanical Monographs 91

Nannorrhops H. Wendl. The name comes from the Greek nannos, meaning dwarf and from rhops, meaning shrub, due to its small size and growth form. Monotypic genus: Nannorrhops ritchieana is its only known representative.

Distribution and ecology The genus is distributed throughout the Holarctic of the Middle East: Arabian Peninsula, Iran, Afghanistan and Pakistan (Irano-Turanian region). It lives in mountainous areas, where it can reach altitudes of up to 1800 m; inhabiting areas where the climate varies from xeric Mediterranean and desert to dry tropical with marked continentality. This palm forms colonies on mountain slopes, by seasonal water beds and where the water table is not very deep.

Description Small palm trees with multiple stems, unarmed. Stems slender, straight, creeping or subterranean; the straight ones are dichotomously divided at the apex; the decumbent too, but often divided at an axillary bud near the base of the stem. The crown is globose, with marcescent leaves. Leaves slightly costapalmate, waxy and grey-bluish; sheath divided forming a Λ, covered with tomentum, thick, woolly and brownish-orange; lamina semicircular or wedge-shaped, divided over half of its length in various segments, which are lanceolate, induplicate, rigid and deeply forked. It is a monoecious and monocarpic genus. Flowering occurs once the stem has divided; the stem that holds the inﬂorescence dies. Inﬂorescence terminal, longer than the leaves and branched up to four orders. Flowers bisexual, ovoid in bud, creamy-white in colour. The fruits are subglobose-ellipsoid berries, brownish-orange when ripe.

Uses and exploitation The apical bud, the newly emerged leaves and the fruits are edible. All parts of the plant are used as fuel. The stems are used as columns, beams or supports. The leaves are used to cover roofs and plated to manufacture ropes, mats and baskets. This species is little known in Spain.

Botanical Monographs 93

Phoenix dactylifera

Phoenix L. The name comes from the Greek phoînix, meaning purple, dark red; it also refers to the mythological phoenix, as the date palm rises from its own ashes. Of the fourteen species in this genus, the Botanic Garden hosts Phoenix canariensis, Phoenix dactylifera, Phoenix reclinata, Phoenix roebelenii, Phoenix rupicola, Phoenix sylvestris and Phoenix theophrasti.

Distribution and ecology They extend from the Holartic region to Paleotropical African and Indo-Malayan regions. Phoenix canariensis and Phoenix theophrasti are Mediterranean. Its ecology is very diverse: they can live in semiarid areas, where their edaphohygrophilous behaviour indicates the existence of groundwater. They also thrive in monsoon climate zones, on the edges of mangroves and wetlands. They can reach up to 1,500 m altitude. Some species are often grouped together, producing more or less extensive palm groves.

Description Palms of single stem, multiple, or subterranean. Some species may have a bulky base covered in roots, or may branch out at different heights. The stem grows straight, inclined or prostrate; it is columnar or tapering towards the apex; covered with ﬁbrous persistent sheaths ellipsoid in shape that when shed leave a lattice drawing on the stem. Crown of semicircular-globose shape, leaves pinnate and marcescent. Pinnae usually lanceolate, induplicate and apex acute; some species have the central vein covered in whitish ramenta, and the pinnae closest to the base of the rachis are transformed into hard, sharp spines. It is a dioecious and polycarpic genus. Inﬂorescences sprout between the leaves, are shorter than these and branch up to one order. Flowers unisexual and sessile. The fruits are oblong-ellipsoid berries, reddish-yellow, or brown-blackish when ripe.

Uses and exploitation The apical bud, the newly emerged leaves and dates of some species are edible. From the prepared sap sugar, palm honey can be extracted, and alcohol distilled. Leaves can be used for rooﬁng constructions and also serve to make rope, fabric and mats.

Botanical Monographs 95

Phoenix dactylifera Pritchardia hildebrandii

Pritchardia Seem. & H. Wendl. Genus dedicated to William Thomas Pritchard, British ofﬁcer and consul of the Fiji Islands. About twenty-seven species are recognized, of which Pritchardia munroi and Pritchardia hildebrandii are grown in the Botanic Garden.

Distribution and ecology Most species are Polynesian (Paleotropical) and distributed in various Pacific Ocean archipelagos and islands. For the most part they are located in the Hawaiian Islands (Hawaiian Region), which include among their flora five endemic species of this genre, some of them on the brink of extinction. There are several species in other Pacific islands, mainly in the Polynesian and the Fijian-Papuana regions (Indo-Malayan). They live on soils of coral or volcanic origin, in steep valleys, slopes and ridges from sea level to 1,400 m altitude.

Description Small-medium sized palms, with a single stem, and unarmed. The stem may have a flared base, be cylindrical or taper towards the apex; grows straight, is covered by residual sheaths and petioles or is naked, ringed with longitudinal fissures. Crown is hemispheric-globose. Lamina is semicircular-circular, costapalmate, the same colour on both sides or with the underside slightly bluish or greyish; is divided up to a third or more of its length in various segments, which are lanceolate, induplicate, straight or pendulate, with indented apex and sharp terminations. It is a monoecious and polycarpic genus. The inflorescences sprout among the leaves, solitary or in groups of two to four protected by the same prophyll. Flowers bisexual, ellipsoid in bud, white, yellow, or orange; when opened, they lose the three petals that form the corolla. The fruits are ovoidglobose berries, reddish-brown or black when ripe.

Uses and exploitation The stems are hard, dense and resistant, traditionally used as supports in buildings. Their large leaves are used to cover roofs and manufacture baskets, mats or fans. The fruits are edible.

Botanical Monographs 97

Ptychosperma macarthurii

Ptychosperma Labill. The name comes from the Greek terms ptyché, meaning creased, crumpled, and spérma, meaning seed, in reference to its grooved seed. Twenty-nine species are recognized, of which the Botanic Garden hosts Ptychosperma macarthurii.

Distribution and ecology Paleotropical asiatic genus with a wide distribution in the IndoMalayan region, reaching the Australian region. New Guinea features the largest number of species and endemisms. They occupy diverse habitats, ranging from mountain rainforests to gallery forests by the river beds and in areas close to the coast with periodic ﬂooding, such as wetlands or mangroves.

Description Small to medium sized palms, without spines, with single or multiple stem. The stem may have a widened base; grows straight, prostrate or slightly curved, is cylindrical, thin, grey-green, ringed and its distal part it has an ornate crownshaft, glabrous or covered with indumentum. Crown is hemispheric, occasionally irregular with marcescent leaves. Leaves pinnate, with pinnae linear, lanceolate or cuneate, reduplicate, biﬁd apex, obliquely truncated, concave, and serrated; the pinnae terminals are joined at the base and are generally not discernible from the rest. It is a monoecious and polycarpic genus. Inﬂorescences are solitary and sprout beneath the leaves, are shorter than these, and branch generally up to two to four orders. Flowers unisexual, sessile and ovoid in bud. The fruits are small ellipsoid-globose berries, orange, reddish, or blackish.

Uses and exploitation The apical bud of some species is edible. The stems - hard, dense and resistant - are used to make fences, bows and spears. The fruits of some species contain crystal irritants. They are very ornamental palms virtually unknown in Spain, where their cultivation is limited and intended for botanical collections.

Botanical Monographs 99

Phoenix dactylifera Ravenea rivularis

Ravenea Bouché ex H. Wendl. Genus dedicated to Louis Ravené, French ofﬁcial at the Berlin Botanical Garden. About eighteen species are recognized, two of which, Ravenea glauca and Ravenea rivularis, grow in the Botanic Garden.

Distribution and ecology Paleotropical palms of the Malagasy Region. Most are endemic to the island of Madagascar. Many species live in rain forests, from sea level up to 2,000 m altitude. Others inhabit the woods that grow along water courses or in ravines and gorges, usually on rocky or sandy soils. Several of these species are on the Red List of threatened species due to the destruction of their natural habitat.

Description Palm trees with a single stem, exceptionally multiple and branched; may have a bulky base, straight, cylindrical or tapering towards the apex, usually ringed without a crownshaft. The crown is obconical to hemispheric; leaves pinnate, unarmed, with numerous pinnae regularly arranged, linear-lanceolate, reduplicate, rigid with acute or acuminate apex. It is a dioecious and polycarpic genus. The inﬂorescences sprout among the leaves, and generally are shorter than these, the staminate ﬂowers are solitary or multiple, but protected by a single prophyll, branching up to one or two orders; the pistillate are solitary and branched up to an order, with their rachilla longer and thicker. Flowers are alike, unisexual, shortly pedicellate, ovoid-globose in bud. The fruits are berries, ellipsoid, globose or lobed, and turn yellow, reddish or blackish brown.

Uses and exploitation The apical bud of some species is edible. The stems are usually hard, dense, durable, and are used for support in construction or for building canoes after hollowing them out. From others the pith is removed as a source of sago. Leaves are used to cover and thatch buildings and make baskets, mats, and hats. They are very ornamental plants that can be widely used in gardening.

Botanical Monographs 101

Rhapidophyllum H. Wendl. & Drude. The name comes from the Greek term phylon, leaf, and Raphis, a genus of the palm family, indicating that it is a palm with leaves like that of Raphis. Genus with only one species, Rhapidophyllum hystrix.

Distribution and ecology This species has its optimum distribution in the central plains of the peninsula of Florida (CaribeoMesoamerican Region, Florida Province, Los Llanos Florida), reaching northward to Georgia, Mississippi and North Carolina, to the south of the American Atlantic Region. It grows in moist, low-lying, often on sandy soils, calcareous or rich in humus. It forms colonies in scrub-covered plains near the coast, and is also present in pine forest understory, deciduous forests and swamp forests. It is one of the more cold-tolerant palms and can stand up to -20Â° C. It is endangered.

Description Small palm trees, stem multiple, sheaths armed with long thin black spines. Stems slender, straight, inclined or prostrate; the latter, upon contact with the ground take root and produce a new plant. The crown is subglobose, with marcescent leaves. Lamina is semicircular, divided up to three quarters of its length into several segments, each with several nerve ribs and folds; the upper sheath is dark green and the undersides grey-green or silver. Segments narrowly lanceolate, with apices truncate and irregularly lobed. It is a genus which is usually dioecious and polycarpic, occasionally monoecious, even polygamous. The inďŹ&#x201A;orescences sprout among the leaves, straight and very short, branching up to one or two orders. Flowers globose in bud, sessile, fragrant, with calyx and corolla of a burgundy hue. Fruits are globose-ovoid, small and covered in brown tomentum, deciduous.

Uses and exploitation During the last century their stems were cut for use in interior decoration or transplanted to be used in ornamental horticulture.

Botanical Monographs 103

Phoenixexcelsa Rhapis dactylifera

Rhapis L. f. ex Aiton. The name derives from the Greek rhapis, which means rod or cane, probably alluding to its thin stem. About eleven species are recognized and Rhapis excelsa, Rhapis humilis and Rhapis multiﬁda grow in the Botanic Garden

Distribution and ecology This genus is native to South-east Asia (Paleotropical, Indo-Malayan) introduced and naturalized in southern Japan. They live at low to moderate altitudes, in tropical pluviseasonal climates. They grow under the tree canopy in deciduous tropophilous tropical forests, usually as part of the dense undergrowth and often on limestone.

Description Small palm trees, stem multiple and unarmed. Stems very thin, straight or inclined, initially covered by residual sheaths and fibres, shed over time, leading to a cylindrical stem, chlorophyllic and ringed. Crown globose-subglobose, with palmate leaves, small and marcescent. Laminae semicircular-circular divided up to three quarters of its length or sometimes almost to the base into several segments, which have several ribs and folds. Segments linear-lanceolate, finely serrated margins, apex acute, irregularly toothed or truncated. Genus dioecious, occasionally polygamous, and polycarpic. The inflorescences sprout among the leaves, are solitary and shorter than these; the pistillate are branched to two orders and the staminate to three, and longer. Pistillate and bisexual flowers are globose, and the staminate more elongated or obovate. The fruits are ovoidglobose berries, creamy-white, brown, or purple-black.

Uses and exploitation With the thin stems, ﬂexible and sturdy, decorative light walking sticks and staves are manufactured. They are highly ornamental palm trees, which can be widely used in gardening. They were introduced in Europe in the eighteenth century, mainly as a houseplant.

Botanical Monographs 105

Phoenix Sabal domingensis dactylifera

Sabal Adans. The genus was described by the French botanist Adanson, and the name granted to this genus is of unknown origin. About sixteen species are recognized, well represented in the Botanic Garden by Sabal causiarum, Sabal, bermudana, Sabal domingensis, Sabal mauritiiformis, Sabal mexicana, Sabal minor and Sabal palmetto.

Distribution and ecology This genus extends over a wide area, comprising the southeast U.S. (Atlantic-American region) much of the islands of the Caribbean (Caribbean-Mesoamerican region), reaching the northern South-American region (New Granada). They live in low to moderate altitudes, usually not exceeding 1,500 m. They populate diverse and varied habitats, but most inhabit open spaces such as dry savannas that undergo periodic ﬁres. In areas close to the sea they can live on dunes, and even colonize swampy or periodically inundated habitats.

Description Palm with a single unarmed stem, which is subterranean in three species. Stem straight, cylindrical, initially covered by residual sheaths and petioles, later naked, smooth or rough, brown or grey, ringed and sometimes slightly cracked. Crown globose, marcescent leaves on young plants and deciduous in mature plants. Leaves costapalmate, with the base of the sheath wide with Λ –like split. Lamina almost circular, divided halfway or almost to its base in many segments, usually solitary, occasionally together in small groups; segments lanceolate, induplicate, apex acute or biﬁd. It is a monoecious and polycarpic genus. The inﬂorescences sprout between the leaves, are compound and can be divided up to four orders. Flowers bisexual, sessile, white or greenish, fragrant. The fruits are berries, small, globose to pyriform, blackish when ripe.

Uses and exploitation The apical bud and fruit of some species are edible. The leaves are used for thatching and for making many utensils such as baskets, mats, brooms and hats. They are highly ornamental palms.

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Serenoa Hook. f. Genus endemic to south-eastern United States, dedicated to the famous American botanist Sereno Watson. Genus with a single species, Serenoa repens.

Distribution and ecology This genus extends over a wide area of the south-eastern United States, in the American-Atlantic Region (Province Llanera de la Costa); it also extends across the peninsula of Florida (Neotropical-Austroamerican region, Caribeo-Mesoamerican region). In general, these palms inhabit littoral areas, with temperate and humid tropical climate, with mild winters and warm summers. They form large colonies of continuous open shrub-lands near the coast, often on sandy soils. It can also be found in the shrub layer of forests of Pinus elliottii.

Description Small palm trees, armed, with multiple or subterranean stem, forming large bushy shrubs, dense and impenetrable. Stem cylindrical, thin, straight, bowed, or creeping; they possess axillary buds that in rare cases can produce new inďŹ&#x201A;orescences or stems. The crown is globose, with marcescent leaves. Leaves small, palmate, light green, glaucous or grey on both sides. Lamina almost circular, divided up to over half its length in various segments, which are lanceolate, induplicate, apex shortly cleft with acute terminations. Genus is monoecious and polycarpic. InďŹ&#x201A;orescences sprout between the leaves, are shorter than these and branch up to three orders occasionally four. Flowers bisexual, sessile, white, fragrant and melliferous. The fruits are ellipsoid berries, black or blue-black when mature, possessing a strong odour.

Uses and exploitation The fruits were used by the natives of Florida for centuries, both as food and for treating diseases of the urinary tract. At the end of last century the effectiveness of fruit extract for the treatment of benign prostate hypertrophy was demonstrated. It is an invasive plant growing in areas and treated as such. This is a very ornamental palm especially the glaucous varieties used in gardening.

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Phoenix dactylifera Syagrus romanzofďŹ ana

Syagrus. Mart. The name comes from the Greek sýagros, which means boar. Some ﬁfty-three species are recognized, as are numerous hybrids. Syagrus coronata, Syagrus schizophylla, and Syagrus romanzofﬁana all grow in the Botanic Garden.

Distribution and ecology These neotropical palms occupy a large area of South America, with Brazil (Brazilian-Paranaense region) being the country harbouring the largest number of species. Populations reach the Amazon Region, Guyana, Orinoco, Chaco and Pampas. Some species live in Amazonian rain forests, or forests of the Atlantic coast, with high rainfall. But most are part of diverse xeric vegetation types, such as the caatinga and the cerrado. They can also be found in coastal shoals, often on sandy or stony soil.

Description Palms with a single stem, multiple or subterranean, which may have ﬂared or bulky base and covered in roots. Stem straight, thin to moderately thick, partially covered by residual sheaths and petioles, or smooth with varying striate and ringed patters depending on the species. Crown hemispheric-globose, with pinnate leaves, occasionally entire, the same colour on both sides. Pinnae arranged on a single plane and regular, or in groups arranged in various planes; they are linear, reduplicate, with the apex acute, rounded, or shortly cleft. Genus monoecious and polycarpic. Inﬂorescences sprout between the leaves, are shorter than these and branch up to one order. Flowers are unisexual, sessile and usually fragrant. The fruits are drupes, ovoid, ellipsoid or globose, with bright colours, brown, or green, with three characteristic pores in the endocarp.

Uses and exploitation The terminal bud and the pith are edible. The stems are used in construction as poles, posts and fences. The leaves are harvested for thatching and to fashion diverse utensils. Oil is extracted from them to make soap or they are harvested for fuel. The ﬂesh of the fruit and seeds is edible, the latter is also a source of oil.

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Phoenix radiata Thrinax dactylifera

Thrinax L. f. ex Sw. The name derives from the Greek word thrînax, which means trident, probably referring to the leaf shape when newly opened. Genus consists of three species, of which Thrinax radiata grows in the Botanic Garden.

Distribution and ecology This genus is mainly Neotropical (Caribeo-Mesoamerican). It inhabits low altitude sites (up to 800 m.) and generally grows on limestone. Thrinax radiata favours areas that are close to the sea and often forms dense stands along dunes or nearby calcareous soils. Cultivated in some South American countries.

Description: Small-medium sized palms, with a single stem, unarmed. The stem base can be bulky and covered in roots; is straight, cylindrical, thin, initially covered by residual sheaths and petioles but eventually smooth, ringed and grey-brown. Crown is globular with marcescent leaves. Leaves palmate, with tomentose sheath and split in Λ shape. Lamina almost circular, green on both sides, or with waxy and grey undersides; its divided up to more than half its length in many segments, which are lanceolate, slightly pendulous, induplicate and with long apices, acute or biﬁd. Genus is monoecious and polycarpic. The inﬂorescences sprout between the leaves and branch up to two orders. Flowers are bisexual, pedicellate, and the calyx and corolla are fused together forming a fragrant cup. The fruits are small globose berries that turn white when mature.

Uses and exploitation The stems are used in construction as pillars and posts. The bud of some species is edible. The leaves are harvested for thatching and for making baskets, hats and other utensils. It is a very ornamental species, which can be widely used in gardening.

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Phoenix dactylifera Trachycarpus fortunei

Trachycarpus H. Wendl. The name consists of two Greek words, Trakhýs, meaning rough or coarse, and karpós, fruit. The term refers to the shape and appearance of the fruit. Usually eight species are recognized, of which Trachycarpus fortunei grows in the Botanic Garden.

Distribution and ecology Paleotropical genus, with a wide Indo-Malayan distribution. These palms grow solitary or in small groups, in mountainous areas up to 2,400 m in altitude, usually on limestone. They are part of the vegetation in mountain forests, moist oak forests, tropophilous forests and grasslands. They withstand cold temperatures and some species remain covered with snow for several months of the year.

Description Small-medium sized palms, solitary stem, occasionally multiple, or subterranean. Stem is straight, cylindrical, thin, covered with persistent sheaths and petioles; these leave have a smooth stem when shed, closely ringed, ﬁnely striate with oblique scars. Crown globular with marcescent leaves. Leaves palmate green on both sides or with greyish waxy undersides. Lamina hemispheric to circular, divided into numerous segments, solitary units or grouped, which are linear-lanceolate, induplicate, with apex cleft and terminations obtuse or acute. This genus is polycarpic, dioecious and occasionally polygamous. Inﬂorescences are solitary, sprout between the leaves, are shorter than these and divided up to four orders. Flowers similar in both sexes, unisexual or bisexual, sessile and globose in bud. The fruits are small kidney-shaped or a coffee bean-shaped berries, from brown to blueblack in colour.

Uses and exploitation The stems are used in construction for poles and pillars. The ﬁbres of the sheaths are used to make brooms and brushes, or twisted to make rope or woven to make coarse clothes for protection against the rain. The plaited leaf segments are used to make many household items such as baskets, hats, fans or mats. The inﬂorescences are eaten as a vegetable. The fruits are food for cattle and the wax layer that covers them is also put to several uses. The seeds have medicinal properties, as they are hemostatic.

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Phoenix dactylifera Trithrinax campestris

Trithrinax Mart. The name comes from the Latin word tri, three, and the Greek thrînax, trident (three forks), probably referring to the division of its leaves. Three species are recognized, and Trithrinax campestris is cultivated in the Botanic Garden.

Distribution and ecology This small genus is NeotropicalAustroamerican, spreading across the Brazilian-Paranaense regions, the Chaco and the Pampas. It inhabits tropical pluviseasonal climates, with pronounced seasonal droughts and moderately cold winters. It lives in open, mountainous or rocky places and in high altitude forest clearings. It is edaphohygrophilous, preferably inhabiting areas by dry river beds where the water table is shallow. It generally grows on arid, sandy, rocky and dry soils, singly or in small groups.

Description Small-medium sized palms, armed, with solitary or multiple stem. The stem base can be bulky and covered in roots; it grows straight, inclined or decumbent, is cylindrical, thin, initially covered by residual prickly sheaths and petioles; eventually rough, grey, ringed and striate. The crown is globose, with marcescent leaves. Leaves palmate or slightly costapalmate, green, blue or grey. Tubular sheath, with reticular and thorny fibres. Lamina is hemispheric-circular split up to more than half its length in many segments, which are linear-lanceolate, induplicate, apex bifid, with acute terminations, lignified and very sharp. Genus is monoecious and polycarpic. Inflorescences sprout between the leaves, are shorter than these and divided up to two orders. Flowers are bisexual, ovoid in bud, pale yellow, fragrant. The fruits are small, globular and yellowish-white, darkening as they mature.

Uses and exploitation The stems are used as posts, pillars and torches. The leaves are used for thatching and to make hats, brooms and baskets. The fibres of the sheaths are used as filters and as filling material. The fruits are edible and fermented produce an alcoholic beverage. The seed is rich in oil. Highly ornamental palm trees, which can be widely used in gardening.

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Phoenix dactylifera Washingtonia robusta

Washingtonia H. Wendl. Genus name dedicated to George Washington, ﬁrst president of the United States. Two species are recognized, Washingtonia ﬁlifera and Washingtonia robusta, both grown in the Botanic Garden.

Distribution and ecology This small genus is native to North America. It is located in the deserts of the south-western United States (Mohave) in the Great Basin region, and in north-western Mexico (Sonora) in the xerophytic Mexican region. They live in semi-arid regions, including deserts, stony and calcareous soils. They grow solitary or in colonies along valleys, plains, places near the coast, canyons and ravines, but always associated with depressions and streams with accessible groundwater.

Description Large palm trees, single stem, which grows straight, base usually bulky and sometimes covered in roots. In W. filifera it is cylindrical and thick, while in W.robusta it is more conical and slender; it remains covered with dry leaves, or is smooth, greyish, slightly ringed with longitudinal fissures. Crown globose. Leaves costapalmate, green, or grey or silvered; the sheath is split like a Λ and petioles armed with curved spines. The lamina is circular, divided up to a third or half its length in many segments, which are linear-lanceolate, apex deeply bifid and acute terminations. It is a monoecious and polycarpic genus. The inflorescences sprout among the leaves, are equal to or longer than these and divide up to three or four orders. Flowers bisexual, creamy-white and fragrant. The fruits are small berries, ovoid-globose, brown-black when ripe.

Uses and exploitation Cahuilla Indians extracted numerous resources from these plants. In the desert they marked places where to ﬁnd shelter and shade. With the leaves they made their homes and various utensils. But above all, they ate its fruits as food: sun toasted and ground, they were turned into ﬂour, used in jams, fermented to produce alcoholic beverages or used in herbal teas and infusions.

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Phoenix X Butyagrus dactylifera nabonnandii

X Butyagrus nabonnandii (Prosch.) Voster. Distribution and ecology This Neotropical genus is located in the Brazilian-Paranaense Regions, the Chaco and the Pampas. In their natural habitat, the genus Butia generates hybrids both intraspeciﬁc and intergenerically. Naturally, Butia capitata crosses with Syagrus romanzofﬁana, giving rise to hybrid X Butyagrus. However, most of these hybridizations have been carried out experimentally since the early twentieth century.

Description Small palm tree, with highly variable morphological characteristics. Stem has slightly ﬂared base, is thick and columnar; initially covered by residual sheaths and petioles; when shed it is rough, grey and ringed. Crown globose, usually with large leaves, pinnate, armed, arched and deep green or bluish-green. The pinnae are arranged on both sides of the rachis, sub-opposite or alternate, forming a single plane, are linear-lanceolate, reduplicate, arched and with acute termination. It is a monoecious and polycarpic species. Inﬂorescences sprout between the leaves, are shorter than these and branch up to an order. Flowers are unisexual, sessile and creamy-white, the staminate ellipsoid in bud, and the pistillate ovoid. The fruits are drupes, globose or ovoid, yellow-red when ripe. Seeds are sterile.

Uses and exploitation French horticulturist Paul Nabonnand created the first artificial hybrid between Butia capitata and Syagrus romanzoffiana in early 1900. It is a very ornamental palm, which can be widely used in gardening, although in Spain it is practically unknown.

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Glossary

Acaulescent. Lacking a visible stem, stem so short leaves appear to sprout from the ground. Aculeus. Prickle or spine of epidermic origin, lacking vascular bundle. Acuminate. Organ, generally a leaf, tapering to a point. Acute. Organ that is distinctly and sharply pointed but not drawn out. Apical. Relating to the apex of the organ. Terminal. Apex. Top or furthest extreme from the point of insertion of an organ. Armed. Describes plants with thorns or spines. A xillary. Concerning the leaf axils. Originating or located between the stem and the leaf.

Belly. Bloated, ventricose. Berry. Fleshy fruit in which the two inner layers (mesocarp and endocarp) are fleshy, although sometimes the endocarp may be fibrous, membranous, massive or papery, but in no case woody. It is the fruit of Phoenix and Chamaerops, among others. Bifid. Body divided into two parts, so that the indentation does not exceed half the length of the organ. Bipinnate. Doubly pinnate. Pinnate leaf in which the leaflets are themselves pinnate. Bisexual. Carrying the two sexes in the same plant or the same flower. Blade. Adaxial (upper) side of any laminar body, which normally faces towards the sun. Bract. Transformed leaf that develops at the base of the inflorescence or, more frequently, of each flower. Branching order. In the inflorescence, position of a branch depending on the number of times the inflorescence has branched from the rachis.

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Canopy. Overstory foliage in jungles and rainforests, formed by the tops of the large broadleaf trees. Carpel. Each of the transformed leaves composing the gynoecium of ﬂowering plants, inside of which are the ovules. Clustered. Plant that from the same foot produces several stems. Manystemmed Crown. The cluster of leaves borne at the tip of the stem. Crownshaft. Roughly cylindrical thickening that occurs in the apex of the stem of some species of palm trees, caused by overlapping tubular leaf sheaths. Costapalmate. Palmate leaf, shaped like the palm of a hand where the petiole is prolonged towards the interior of the lamina, like a short midrib or costa. Cuneiform. Wedge-shaped, triangular, tapered towards the base. Cluster. Stems arising from a single base on a clustered palm.

Deciduous. Describes a short-lived organ, shed periodically. Decumbent. Stem that tends to grows reclining, prostrate on the ground, but with the apex or tip ascending. Dichotomous. Describes a ramiﬁcation in which the growing point is forked into two equal branches. Dioecious. Plant with unisexual ﬂowers arranged on different individuals, so as to have male and female individuals. Drupe. Fleshy fruit in which the innermost layer (endocarp) is stiff, so that the seed is surrounded by a woody layer. It is the type of fruit of Cocos, Butia or Jubaea, among others.

Edaphic. Relative to soil. Endocarp. Inside of the pericarp of the fruit, which corresponds to the epidermis of the carpelar leaf. Elliptical. Ellipse shaped. Epicarp. Outermost layer of the fruit, the external part of the pericarp. Epigeous. Is said of the organ that, being usually subterranean, is developed aboveground.

Fibrous. Formed by ﬁbres or thin like a ﬁbre. Fusiform. Spindle-shaped, narrowed towards the ends.

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Gineceum. Set of carpels of a ﬂower. Glabrous. Said of a surface devoid of hairs, scales or tomentum; hairless. Glaucous. Light green, blue.

Hybrid. Offspring resulting from the cross between two individuals of different species of the same or different genus.

Indument. Set of hairs, glands and scales that cover the surface of plant organs. Induplicate. Said of any laminar body (leaves, pinnae segments ...) having upturned margins so that the section is V-shaped Inflorescence. Part of the branching system capable of forming flowers, including all its bracts and branches Infrafoliar. Positioned below the leaves. Infructescence. Mature inflorescence, where the flowers are replaced by fruits. Interfoliar. Situated between the leaves.

Lattice. Surface ornamentation of any body that forms a mesh-shaped pattern or grid. Lamina. Widest part of the leaf, generally planar. Also called blade. Lanceolate. Laminar body (leaf, bract, petal ...) shaped like a spearhead. Leathery. With a hard and ﬂexible consistence, leather-like. Ligule. Laminar appendix or tongue in insertion area of the sheath and the petiole of some species. It is also used as a synonym for hastula. Linear. Body, generally laminar, much longer than wide, sides roughly parallel throughout its development.

Marcescent. Describes the organs that, once dry, remain attached to the plant without shedding. Melliferous. Plants or ﬂowers that produce much nectar, and consequently attract honey bees. Mesocarp. Middle part of the pericarp of the fruit. Monocarpic. Plant that blooms only once during its life. Monocaule. Plant that develops a single stem.

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Monoic. Plant with unisexual flowers arranged on the same individual, so that it presents male flowers and female flowers on the same stem. Monotypic. Genus that consists of a single species.

Obconical. Of inverted conical shape, with the apex wider than the base. Oblong. Wider than long. Obovate. Of inverted ovate shape, with the apex wider than the base. Obtuse. Rounded apex. Orbicular. Circular, round. Ovate. Describes lamellar bodies (leaves, bracts, petals ...) egg-shaped. Ovoid. Describes solid organs (fruits, seeds ...) egg-shaped.

Palmate. Leaf type shaped like an open hand. Palmetum. Collection of palm trees. Petiole. Stalk that joins the leaf blade to the sheath or stem. Pedicel. Stalk that holds each flower in a compound inflorescence. Peduncle. Stalk holding each of the flowers in a simple inflorescence, or linking the inflorescence, simple or compound, to the stem. Perianth. Structure that protects reproductive parts of the flower, usually formed by petals and sepals. Pericarp. Wall of the fruits formed from the carpelar leaf. Pinnae. Each of the parts of a compound leaf; leaflet. Pinnate. Type of compound leaf, with pinnae in two rows one on each side of the stem. Piriformis. Pear-Shaped. Pistil. Carpel or group of carpels fused or free which are arranged in a flower, gynoecium. Pistillate flower. Flower with pistil; in the case of unisexual flowers, the female. Polycarpic. Plant that blooms several times throughout its life. Premorse. Describes irregularly truncated bodies, nibbled. Prophyll. The first leaf that arises from any lateral bud, regardless of whether it becomes a branch or an inflorescence. Pubescent. Said of any organ covered with fine soft hairs.

Rachilla. Any of the lateral branches of the rachis in a branched inflorescence on which flowers are arranged. Rachis. Main axis of the inflorescence. Midrib of compound leaves on which are arranged the pinnae.

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Ramentum. Set of little hairs or elongated scales, arranged in the base or along the underside of some pinnae. Raphe. Embossed or depressed line observed on the surface of the seed. Reduplicated. Said of any laminar body (leaves, pinnae segments ...) having margins bent downwards, so that the section has an inverted V shape. Rhizome. Underground stem. Ringed Describes a stem with circular marks, more or less regular, left after shedding the leaves.

Segment. Refers to each of the elements that reach the median vein or the base of the leaf blade, in palmatisect or pinnatisect leaves. By extension, each of the elements that a palmate leaf is divided into. Sessile. Body that is directly attached to the stem without petiole, peduncle or pedicel. Sheath. Leaf base, more or less enlarged, which at least partly embraces the stem; may be more or less succulent to clearly fibrous. The sheath is tapered until it extends to the petiole. Sinus. Each incision in a palmate leaf. Spathe. Large bract, often very hard, that encloses the spadix before opening. Spike. A racemose inflorescence, unbranched with sessile flowers. Spine. Hardened organ, pointed, with vascular bundle. Staminate flower. Flower with stamens; in the case of unisexual flowers, the male. Stamen. Flower organ where pollen is produced. Staminode. Non-functional sterile stamen. Stem. Main axis. In palms long, usually unbranched, trunk, sometimes called stipe. Sub-. Prefix that attenuates the meaning of the word it accompanies: without reaching, almost near.

Terminal. Said of any organ that sits at the end of a stem, leaf or flower; apical. Tomentum. Dense coating of short hairs more or less intertwined. Transverse venation. Set of veins that arise from the main vein. Triad. Set of three things closely interlinked. In palms, the term is applied to each set of three flowers, a central female flower and two lateral male flowers, which are arranged on some inflorescences. Trunk. In palms, the stem or stipe. Truncated. As though cut off nearly straight across.

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Unisexual. Describes a ďŹ&#x201A;ower or plant that only has one sex. Unarmed. Plant or body carries no thorns or aculeus. Underside. Abaxial side (bottom) of any laminar organ, which normally faces towards the ground.

Variegated. With various colours. Vascular. Relative to conducting tissues. Vein. Each of the vascular bundles developed in lamellar bodies (leaves, bracts, sepals, petals ...) of a plant. Whorl. Set of similar organs, in number greater than two, born at the same level on the axis that carries them.

Xerophyte. Plant adapted to live in dry regions or with long periods of drought.

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Bibliography

BALICK, M. J. 1988. The palm, tree of life. Biology, utilization and conservation, Volumen 6. The New York Botanical Garden Press, New York, USA. BALICK, M. J. y H. T. BECK. 1990. Useful palms of the world. Columbia University Press. New York, USA. BARROW, S. 1998. A revision of Phoenix. Kew bulletin, Vol. 53, part 3. Royal Botanic Gardens, Kew. Inglaterra. COSTA, M., J. GÜEMES, C. BENTO, E. ESTRELLES, J. RIERA, Mª. J. CARRAU. 2001. El Jardín Botánico de la Universidad de Valencia. Ed. Universitat de València. Valencia, España. DEL CAÑIZO, J. A. 2011. Palmeras: Todos los géneros y 565 especies. Ed. MundiPrensa. Madrid, España. DOVE, J. 2010. Australian palms: biogeografy, ecology and systematics. CSIRO Publishing Collinwood, Victoria, Australia. DRANSFIELD, J., N. W. UHL, C. B. ASMUSSEN, W. J. BAKER, M. M. HARLEY y C. E. LEWIS. 2008. Genera palmarum. The evolution and classiﬁcation of palms. Royal Botanic Gardens, Kew. Londres, Inglaterra. FONT QUER, P. 1953. Diccionario de botánica. Ed. Labor S.A. Madrid, España. GOVAERTS, R y J. DRANSFIELD. 2005. World Checklist of Palms. Royal Botanic Gardens, Kew. Londres, Inglaterra. HALLE, F. 2004. Architectures de plantes. Ed. First Palavas. Montpellier, Francia. HENDERSON, A. 2009. Palms of southern Asia. Ed. Princeton University Press & The New York Botanic Garden. New Jersey, UK., New York, USA. HENDERSON, A., G. GALEANO y R. BERNAL. 1995. Field guide to the palms of the Americas. Ed. Princeton University Press. New Jersey, UK. HODEL, D. R. 1992. Chamaedorea palms, the species and their cultivation. Ed. Allen Spress. Lawrence, USA. IUCN. 1988. Rare and threatened palms of the new Word. IUCN Botanic Gardens Conservation Secretariat, Cambridge, Inglaterra. IUCN. Red List of Threatened Species. http://www.iucnredlist.org. IUCN Red List

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Unit. IUCN UK Ofﬁce, 219c Huntingdon Road. Cambridge CB3 0DL. Inglaterra. JACQUEMIN, D. 1999. Les palmiers ornamentaux pour le climats mediterranéens. Ed. Editions Champﬂour. Marly-le-Roi. Francia MOYA, B., C. LITTARDI y J. PLUMED. 2005. La poda de las palmeras ornamentales. Biología, ecología y gestión. Ed. Asociación Española de Arboricultura. Valencia, España. QUERO, H. 2000. El complejo Brahea-Erythea (Palmae: Coryphoideae). Proyecto CONABIO L 216. Universidad Nacional Autónoma de México, Instituto de Biología, Jardín Botánico. México. RIVAS -MARTÍNEZ, S., G. NAVARRO, A. PENAS and M. Costa. Biogeographic Map of South America. A preliminary survey. International Journal of Geobotanical Research, Vol. nº 1, December 2011, pp. 21-40 + Map. ROOD, A. N. 1998. Revision of Livistona in Australia. Revista Telopea, Vol. 8 (1). Ed. National Herbarium of New South Wales, Royal Botanic Gardens, Sydney, Australia. TOMLINSON, P. B. 1990. The extructural biology of palms. Ed. Oxford University Press. New York, USA. ZONA, S. 1990. A monograph of Sabal. Revista Aliso, Vol. 2, Nº 4. Rancho Santa Ana Botanic Gardens, Claremont, California, USA.

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PALM SPECIES GROWING IN THE BOTANIC GARDEN 1. Acoelorrhaphe wrightii (Griseb. & H. Wendl.) H. Wendl. ex Becc. 2. Adonidia merrillii (Becc.) Becc. 3. Aiphanes horrida (Jacq.) Burret. 4. Allagoptera arenaria (Gomes) Kuntze. 5. Archontophoenix alexandrae (F. MĂźell.) H. Wendl. & Drude. 6. Archontophoenix cunninghamiana (H. Wendl.). H. Wendl. & Drude. 7. Areca catechu L. 8. Areca triandra Roxb. ex F. Buch.-Ham. 9. Arenga engleri Becc. 10. Bismarckia nobilis. Hildebr. & H. Wendl. 11. Brahea aculeata (Brandegee) H. E. Moore. 12. Brahea armata S. Watson. 13. Brahea brandegeei (Purpus) H. E. Moore. 14. Brahea edulis H. Wendl. ex S. Watson. 15. Brahea sarukhanii H. J. Quero. 16. Butia capitata (Mart.) Becc. 17. Caryota mitis Lour. 18. Caryota urens L. 19. Chamaedorea cataractarum Mart. 20. Chamaedorea costaricana Oerst. 21. Chamaedorea elegans Mart. 22. Chamaedorea metallica O. F. Cook ex H. E. Moore. 23. Chamaedorea microspadix Burret. 24. Chamaedorea oblongata Mart. 25. Chamaedorea potchutlensis Liebm. ex Mart. 26. Chamaedorea radicalis Mart. 27. Chamaedorea seifrizii. Burret. 28. Chamaerops humilis L. 29. Chambeyronia macrocarpa (Brongn.) Vieill. ex Becc. 30. Coccothrinax crinita (Griseb. & H. Wendl. ex C. H. Wright) Becc. 31. Copernicia alba Morong. 32. Dypsis lutescens (H. Wendl.) Beentje & J. Dransf. 33. Gaussia maya (O. F. Cook) H. J. Quero & Read. 34. Gaussia spirituana Moya & Leiva. 35. Howea belmoreana (C. Moore & F. Muell.) Becc. 36. Howea forsteriana (F. Muell.) Becc. 37. Hyophorbe lagenicaulis (L. H. Bailey) H. E. Moore. 38. Hyophorbe verschaffeltii H. Wendl. 39. Jubaea chilensis (Molina) Baill. 40. Licuala grandis H. Wendl.

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41. Livistona australis (R. Br.) Mart. 42. Livistona chinensis (Jacq.) R. Br. ex Mart. 43. Livistona decora (W. Bull) Dowe. 44. Livistona fulva Rodd. 45. Livistona muellery F.M. Bailey. 46. Livistona nitida Rodd. 47. Nannorrhops ritchieana (Griff.) Aitch. 48. Phoenix canariensis Chabaud. 49. Phoenix dactylifera L. 50. Phoenix reclinata Jacq. 51. Phoenix roebelenii J. O’Brien 52. Phoenix rupicola T. Anders. 53. Phoenix sylvestris (L.) Rosb. 54. Phoenix theophrasti Greuter. 55. Pritchardia hillebrandii Becc. 56. Pritchardia munroi Rock. 57. Ptychosperma macarthurii (H. Wendl. ex H. J. Veitch) H. Wendl. ex Hook. f. 58. Ravenea glauca Jum. & H. Perrier. 59. Ravenea rivularis Jum. & H. Perrier. 60. Rhapidophyllum hystrix (Fraser ex Thouin) H. Wendl. & Drude. 61. Rhapis excelsa (Thunb.) Henry. 62. Rhapis humilis Blume. 63. Rhapis multifida Burret. 64. Sabal bermudana L. H. Bailey. 65. Sabal causiarum (O. F. Cook) Becc. 66. Sabal domingensis Becc. 67. Sabal mauritiiformis (H. Karst.) Griseb. & H. Wendl. 68. Sabal mexicana Mart. 69. Sabal minor (Jacq.) Pers. 70. Sabal palmetto (Walter) Lodd. ex Schult. & Schult. f. 71. Serenoa repens (W. Bartram) Small. 72. Syagrus coronata (Mart.) Becc. 73. Syagrus romanzoffiana (Cham.) Glassman. 74. Syagrus schizophylla (Mart.) Glassman. 75. Thrinax radiata Lodd. ex Schult. & Schult. f. 76. Trachycarpus fortunei (Hook.) H. Wendl. 77. Trithrinax campestris (Burmeist.) Drude & Griseb. 78. Washingtonia filifera (Linden ex André) H. Wendl. 79. Washingtonia robusta H. Wendl. 80. x Butyagrus nabonnandii (Prosch.) Voster.

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