Arecaceae

Morphology

Whether as shrubs, trees, or vines, palms have two methods of growth: solitary or clustered. The common representation is that of a solitary shoot ending in a crown of leaves. This monopodial character may be exhibited by prostrate, trunkless, and trunk-forming members. Some common palms restricted to solitary growth include Washingtonia and Roystonea. Palms may instead grow in sparse though dense clusters. The trunk develops an axillary bud at a leaf node, usually near the base, from which a new shoot emerges. The new shoot, in turn, produces an axillary bud and a clustering habit results. Exclusively sympodial genera include many of the rattans, Guihaia, and Rhapis. Several palm genera have both solitary and clustering members. Palms which are usually solitary may grow in clusters, and vice versa. These aberrations suggest the habit operates on a single gene.

Palms have large, evergreen leaves that are either palmately ('fan-leaved') or pinnately ('feather-leaved') compound and spirally arranged at the top of the stem. The leaves have a tubular sheath at the base that usually splits open on one side at maturity. The inflorescence is a spadix or spike surrounded by one or more bracts or spathes that become woody at maturity. The flowers are generally small and white, radially symmetric, and can be either uni- or bisexual. The sepals and petals usually number three each, and may be distinct or joined at the base. The stamens generally number six, with filaments that may be separate, attached to each other, or attached to the pistil at the base. The fruit is usually a single-seeded drupe (sometimes berry-like) but some genera (e.g. Salacca) may contain two or more seeds in each fruit.

Like all monocots, palms do not have the ability to increase the width of a stem (secondary growth) via the same kind of vascular cambium found in non-monocot woody plants. This explains the cylindrical shape of the trunk (almost constant diameter) that is often seen in palms, unlike in true trees. However, many palms, like some other monocots, do have secondary growth, although because it does not arise from a single vascular cambium producing xylem inwards and phloem outwards, it is often called "anomalous secondary growth".

The Arecaceae are notable among monocots for their height and for the size of their seeds, leaves, and inflorescences. Ceroxylon quindiuense, Colombia's national tree, is the tallest monocot in the world, reaching up to 60 m tall. The coco de mer (Lodoicea maldivica) has the largest seeds of any plant, 40–50 cm in diameter and weighing 15–30 kg each. Raffia palms (Raphia spp.) have the largest leaves of any plant, up to 25 m long and 3 m wide. The Corypha species have the largest inflorescence of any plant, up to 7.5 m tall and containing millions of small flowers. Calamus stems can reach 200 m in length.

Range and habitat

Most palms grow in the tropics and subtropics. They are abundant throughout the tropics and subtropics, and thrive in almost every habitat they are in. Their diversity is highest in wet, lowland forests, especially in ecological "hotspots" such as Madagascar, which has more endemic palms than all of Africa. Colombia may have the highest number of palm species in one country. Palms are most commonly seen throughout Africa, South America, the Arabian peninsula, South and Southeast Asia, northern Australia, the islands of tropical and subtropical parts of the Pacific Ocean, Mexico, Puerto Rico, and the U.S sunbelt including California, Florida, and Hawaii.

Only about 130 palm species naturally grow entirely beyond the tropics, mostly in the subtropical highlands and warm temperate lowlands. The northernmost native palm is Chamaerops humilis, which reaches 44°N latitude in southern France. The southernmost palm is the Rhopalostylis sapida, which reaches 44°S on the Chatham Islands where an oceanic climate prevails. Some palms, such as the Trachycarpus fortunei, grow well under cultivation in temperate climates, some at 50°N or even farther north in oceanic climates (Ireland, Scotland, England, Long Island and the Pacific Northwest, from Oregon to Vancouver).

Palms inhabit a variety of ecosystems. More than two-thirds of palm species live in humid moist forests, where some species grow tall enough to form part of the canopy and shorter ones form part of the understory. Some species form pure stands in areas with poor drainage or regular flooding, including Raphia hookeri which is common in coastal freshwater swamps in West Africa. Other palms live in tropical mountain habitats above 1000 m, such as those in the genus Ceroxylon native to the Andes. Palms may also live in grasslands and scrublands, usually associated with a water source, and in desert oases such as the date palm. A few palms are adapted to extremely basic lime soils, while others are similarly adapted to extreme potassium deficiency and toxicity of heavy metals in serpentine soils.

Palms are a monophyletic group of plants, meaning the group consists of a common ancestor and all its descendants. Extensive taxonomic research on palms began with botanist H.E. Moore, who organized palms into 15 major groups based mostly on general morphological characteristics. The following classification, proposed by N.W. Uhl and J. Dransfield in 1987, is a revision of Moore's classification that organizes palms into six subfamilies.

A few general traits of each subfamily are listed below.

The Coryphoideae are the most diverse subfamily, and are a paraphyletic group, meaning all members of the group share a common ancestor, but the group does not include all the ancestor's descendants. Most palms in this subfamily have palmately lobed leaves and solitary flowers with three, or sometimes four carpels. The fruit normally develops from only one carpel.

Subfamily Calamoideae includes the climbing palms, such as rattans. The leaves are usually pinnate; derived characters (synapomorphies) include spines on various organs, organs specialized for climbing, an extension of the main stem of the leaf-bearing reflexed spines, and overlapping scales covering the fruit and ovary.

Subfamily Nypoideae contains only one species, Nypa fruticans, which has large, pinnate leaves. The fruit is unusual in that it floats, and the stem is dichotomously branched, also unusual in palms.

Subfamily Ceroxyloideae has small to medium-sized flowers, spirally arranged, with a gynoecium of three joined carpels.

The Arecoideae are the largest subfamily, with six diverse tribes (Areceae, Caryoteae, Cocoeae, Geonomeae, Iriarteeae, and Podococceae) containing over 100 genera. All tribes have pinnate or bipinnate leaves and flowers arranged in groups of three, with a central pistillate and two staminate flowers.

The Phytelephantoideae are a monoecious subfamily. Members of this group have distinct monopodial flower clusters. Other distinct features include a gynoecium with five to 10 joined carpels, and flowers with more than three parts per whorl. Fruits are multiple-seeded and have multiple parts.

Currently, few extensive phylogenetic studies of the Arecaceae exist. In 1997, Baker et al. explored subfamily and tribe relationships using chloroplast DNA from 60 genera from all subfamilies and tribes. The results strongly showed the Calamoideae are monophyletic, and Ceroxyloideae and Coryphoideae are paraphyletic. The relationships of Arecoideae are uncertain, but they are possibly related to the Ceroxyloideae and Phytelephantoideae. Studies have suggested the lack of a fully resolved hypothesis for the relationships within the family is due to a variety of factors, including difficulties in selecting appropriate outgroups, homoplasy in morphological character states, slow rates of molecular evolution important for the use of standard DNA markers, and character polarization. However, hybridization has been observed among Orbignya and Phoenix species, and using chloroplast DNA in cladistic studies may produce inaccurate results due to maternal inheritance of the chloroplast DNA. Chemical and molecular data from non-organelle DNA, for example, could be more effective for studying palm phylogeny.

Selected genera

See list of Arecaceae genera arranged by taxonomic groups or by alphabetical order for a complete listing of genera.

Evolution

The Arecaceae are the first modern family of monocots appearing in the fossil record around 80 million years ago (Mya), during the late Cretaceous period. The first modern species, such as Nypa fruticans and Acrocomia aculeata, appeared 94 Mya, confirmed by fossil Nypa pollen dated to 94 Mya. Palms appear to have undergone an early period of adaptive radiation. By 60 Mya, many of the modern, specialized genera of palms appeared and became widespread and common, much more widespread than their range today. Because palms separated from the monocots earlier than other families, they developed more intrafamilial specialization and diversity. By tracing back these diverse characteristics of palms to the basic structures of monocots, palms may be valuable in studying monocot evolution. Several species of palms have been identified from flowers preserved in amber, including Palaeoraphe dominicana and Roystonea palaea. Evidence can also be found in samples of petrified palmwood.

Uses

Human use of palms is as old or older than human civilization itself, starting with the cultivation of the date palm by Mesopotamians and other Middle Eastern peoples 5000 years or more ago. Date wood, pits for storing dates, and other remains of the date palm have been found in Mesopotamian sites. The date palm had a tremendous effect on the history of the Middle East. W.H. Barreveld wrote:

One could go as far as to say that, had the date palm not existed, the expansion of the human race into the hot and barren parts of the "old" world would have been much more restricted. The date palm not only provided a concentrated energy food, which could be easily stored and carried along on long journeys across the deserts, it also created a more amenable habitat for the people to live in by providing shade and protection from the desert winds (Fig. 1). In addition, the date palm also yielded a variety of products for use in agricultural production and for domestic utensils, and practically all parts of the palm had a useful purpose.

An indication of the importance of palms in ancient times is that they are mentioned more than 30 times in the Bible, and at least 22 times in the Quran.

Arecaceae have great economic importance, including coconut products, oils, dates, palm syrup, ivory nuts, carnauba wax, rattan cane, raffia, and palm wood.

Along with dates mentioned above, members of the palm family with human uses are numerous.

The southeastern U.S. state of South Carolina is nicknamed the Palmetto State after the sabal palmetto (cabbage palmetto), logs from which were used to build the fort at Fort Moultrie. During the American Revolutionary War, they were invaluable to those defending the fort, because their spongy wood absorbed or deflected the British cannonballs. The sabal palmetto is also the state tree of Florida. Some palms can be grown as far north as the United States' Mid-Atlantic, such as the National Arboretum in Washington, D.C., southern Midwest, and even north along the Pacific coast to Oregon, Washington, and British Columbia, where ocean winds have a warming effect. Species of transplanted palms have even been known to have survived as far north as Devon. The Chinese Trachycarpus fortunei is being grown experimentally on the Faroe Islands at 62°N, with young plants doing well so far.

Endangered species

Like many other plants, palms have been threatened by human intervention and exploitation. The greatest risk to palms is destruction of habitat, especially in the tropical forests, due to urbanization, wood-chipping, mining, and conversion to farmland. Palms rarely reproduce after such great changes in the habitat, and those with small habitat ranges are most vulnerable to them. The harvesting of heart of palm, a delicacy in salads, also poses a threat because it is derived from the palm's apical meristem, a vital part of the palm that cannot be regrown (except in domesticated varieties, e.g. of peach palm). The use of rattan palms in furniture has caused a major population decrease in these species that has negatively affected local and international markets, as well as biodiversity in the area. The sale of seeds to nurseries and collectors is another threat, as the seeds of popular palms are sometimes harvested directly from the wild. At least 100 palm species are currently endangered, and nine species have reportedly recently become extinct.

However, several factors make palm conservation more difficult. Palms live in almost every type of warm habitat and have tremendous morphological diversity. Most palm seeds lose viability quickly, and they cannot be preserved in low temperatures because the cold kills the embryo. Using botanical gardens for conservation also presents problems, since they can only house a few plants of any species or truly imitate the natural setting. Also, the risk of cross-pollination can lead to hybrid species.

The Palm Specialist Group of the World Conservation Union (IUCN) began in 1984, and has performed a series of three studies to find basic information on the status of palms in the wild, use of wild palms, and palms under cultivation. Two projects on palm conservation and use supported by the World Wildlife Fund took place from 1985 to 1990 and 1986–1991, in the American tropics and southeast Asia, respectively. Both studies produced copious new data and publications on palms. Preparation of a global action plan for palm conservation began in 1991, supported by the IUCN, and was published in 1996.

The rarest palm known is Hyophorbe amaricaulis. The only living individual remains at the Botanic Gardens of Curepipe in Mauritius.

Pest species

Pests that attack a variety of species of palm trees include:

Symbolism

The palm branch was a symbol of triumph and victory in pre-Christian times. The Romans rewarded champions of the games and celebrated military successes with palm branches. Early Christians used the palm branch to symbolize the victory of the faithful over enemies of the soul, as in the Palm Sunday festival celebrating the triumphal entry of Jesus into Jerusalem. In Judaism, the palm represents peace and plenty, and is one of the Four Species of Sukkot; the palm may also symbolize the Tree of Life in Kabbalah.

Panaiveriyamman was an ancient Tamil tree deity related to fertility. Named after panai, the Tamil name for the Palmyra palm, she was also known as Taalavaasini, a name that further related her to all types of palms.

Today, the palm, especially the coconut palm, remains a symbol of the tropical island paradise. Palms appear on the flags and seals of several places where they are native, including those of Haiti, Guam, Saudi Arabia, Florida, and South Carolina.

Other plants

Some species commonly called palms, though they are not true palms, include: