| Podocarpus | |
|---|---|
 | |
| Podocarpus totara | |
Scientific classification | |
| Kingdom: | Plantae |
| Clade: | Tracheophytes |
| Clade: | Gymnospermae |
| Division: | Pinophyta |
| Class: | Pinopsida |
| Order: | Araucariales |
| Family: | Podocarpaceae |
| Genus: | Podocarpus L'HérexPers.[2] |
| Type species | |
| Podocarpus elongatus L'Hérex Pers.[2] | |
| Species[3] | |
116, see text | |
| Synonyms[3] | |
MargbensoniaBobrov & Melikian | |
Podocarpus (/ˌpoʊdəˈkɑːrpəs/[4]) is agenus ofconifers, the most numerous and widely distributed of the podocarp family, thePodocarpaceae.Podocarpus species areevergreenshrubs ortrees, usually from 1 to 25 m (3 to 82 ft) tall, known to reach 40 m (130 ft) at times. Thecones have two to five fused cone scales, which form a fleshy,berry-like, brightly colouredreceptacle at maturity. The fleshy cones attractbirds, which then eat the cones and disperse the seeds in their droppings.Plants of the World Online accepts 116 species.[3] Other authorities place 97 to 107 species in the genus depending on thecircumscription of the species.[2][5][6][7]
Species are cultivated as ornamental plants for parks and large gardens. Thecultivar 'County Park Fire' has won theRoyal Horticultural Society'sAward of Garden Merit.[8]
The name comes from Greekπούςpoús meaning "foot" andκαρπόςkarpós meaning "fruit".[further explanation needed]
Common names for various species include "yellowwood" and "pine",[5] as in the plum pine (Podocarpus elatus)[9] or the Buddhist pine (Podocarpus macrophyllus).[10]
Podocarpus species areevergreen woody plants. They are generally trees, but may also be shrubs.[2] The trees can reach a height of 40 metres (130 ft) at their tallest.[5] Some shrubby species have a decumbent growth habit. The primarybranches form pseudowhorls around the trunk. The bark can be scaly or fibrous and peeling with vertical strips. Terminalbuds are distinctive with bud scales that are often imbricate and can be spreading.[2]
Theleaves are simple and flattened, and may be sessile or short petiolate. Thephyllotaxis or leaf arrangement is spiral, and may be subopposite on some shoots.[2][11] The leaves are usually linear-lanceolate or linear-elliptic in shape, though they can be broader lanceolate, ovate, or nearly elliptic in some species.[2][5][11] Juvenile leaves are often larger than adult leaves, though similar in shape.[11] The leaves are coriaceous and have a distinct midrib. Thestomata are usually restricted to the abaxial or underside of the leaf, forming two stomatal bands around the midrib.[2]
Podocarpus spp. are generallydioecious, with the male pollen cones and female seed cones borne on separate individual plants, but some species may bemonoecious. Thecones develop fromaxillary buds, and may be solitary or form clusters.[2]
The pollen cones are long andcatkin-like in shape. They may be sessile or short pedunculate. A pollen cone consists of a slender rachis with numerous spirally arranged microsporophylls around it. Each triangular microsporophyll has two basalpollen-producing pollen sacs. The pollen is bisaccate.[2]
The seed cones are highly modified with the few cone scales swelling and fusing at maturity. The cones are pedunculate and often solitary. The seed cone consists of two to five cone scales of which only the uppermost one or rarely two nearest the apex of the cone are fertile. Each fertile scale usually has one apical ovule. The infertile basal scales fuse and swell to form a succulent, usually brightly colored receptacle. Each cone generally has only oneseed, but may have two or rarely more. The seed is attached to the apex of the receptacle. The seed is entirely covered by a fleshy modified scale known as an epimatium. The epimatium is usually green, but may be bluish or reddish in some species.[2][11]
The natural distribution of the genus consists of much of Africa, Asia, Australia, Central and South America, and several South Pacific islands. The genus occurs from southern Chile north to Mexico in the Americas and from New Zealand north to Japan in the Asia-Pacific region.[2] As recently as the Late Pliocene, Podocarpus had a range in the Northern United States.[12]
Podocarpus and the Podocarpaceae wereendemic to the ancient supercontinent ofGondwana, which broke up intoAfrica,South America,India,Australia-New Guinea,New Zealand, andNew Caledonia between 105 and 45 million years ago.Podocarpus is a characteristic tree of theAntarctic flora, which originated in the cool, moist climate of southern Gondwana, and elements of the flora survive in the humid temperate regions of the former supercontinent. As the continents drifted north and became drier and hotter, podocarps and other members of the Antarctic flora generally retreated to humid regions, especially in Australia, wheresclerophyll genera such asAcacia andEucalyptus became predominant. The flora ofMalesia, which includes theMalay peninsula, Indonesia, thePhilippines, and New Guinea, is generally derived from Asia, but includes many elements of the old Gondwana flora, including several other genera in the Podocarpaceae (Dacrycarpus,Dacrydium,Falcatifolium,Nageia,Phyllocladus, and the Malesian endemicSundacarpus), and alsoAgathis in theAraucariaceae.
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The two subgenera,Podocarpus andFoliolatus, are distinguished by cone and seed morphology.[13]
InPodocarpus, the cone is not subtended by lanceolate bracts, and the seed usually has an apical ridge. Species are distributed in the temperate forests ofTasmania,New Zealand, and southernChile, with a few occurring in the tropical highlands of Africa and the Americas.
InFoliolatus, the cone is subtended by two lanceolate bracts ("foliola"), and the seed usually lacks an apical ridge. The species are tropical and subtropical, concentrated in eastern and southeastern Asia and Malesia, overlapping with subgenusPodocarpus in northeastern Australia andNew Caledonia.
Species in family Podocarpaceae have been reshuffled a number of times based on genetic and physiological evidence, with many species formerly assigned toPodocarpus now assigned to other genera. A sequence of classification schemes has moved species betweenNageia andPodocarpus, and in 1969, de Laubenfels divided the huge genusPodocarpus intoDacrycarpus, Decussocarpus (an invalid name he later revised to the validNageia),Prumnopitys, andPodocarpus. Some species of genusAfrocarpus were formerly inPodocarpus, such asAfrocarpus gracilior.
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In 1985David J. de Laubenfels divided the genus into two subgenera,Podocarpus andFoliolatus, and further divided each subgenus into nine sections.[16] Robert Reid Mill wrote (2015) that while the two subgenera have been strongly supported by subsequent evidence, the sections are "mostly are poorly supported by molecular evidence".[17] In 2015 de Laubenfels revised the sections of subgenusFoliolatus.[18]
MalePodocarpus spp. are extremely allergenic, and have anOPALS allergy-scale rating of 10 out of 10. Conversely, completely femalePodocarpus plants have an OPALS rating of 1, and are considered "allergy-fighting", as they capture pollen while producing none.[20]
Podocarpus resembleyews, and as with yews, the stems, leaves, flowers, and pollen ofPodocarpus are all poisonous. Additionally, the leaves, stems, bark, and pollen arecytotoxic. The malePodocarpus blooms and releases this cytotoxic pollen in the spring and early summer.
The earliest use ofP. elongatus dates back to the southern AfricanMiddle Stone Age where it was used to produce an adhesive by distillation.[21] Today, several species ofPodocarpus are grown as garden trees, or trained into hedges, espaliers, or screens. In the novelJurassic Park byMichael Crichton,Podocarpus trees (misspelled as "protocarpus") were used on Isla Nublar, Costa Rica, to conceal electric fences from visitors.[22] Common garden species used for their attractive deep-green foliage and neat habits includeP. macrophyllus, known commonly as Buddhist pine, fern pine, orkusamaki,P. salignus from Chile, andP. nivalis, a smaller, red-fleshy-coned shrub. Some members of the generaNageia,Prumnopitys, andAfrocarpus are marketed under the genus namePodocarpus.
The red, purple, or bluish fleshy cone (popularly called a "fruit") of most species ofPodocarpus are edible, raw or cooked into jams or pies. They have a mucilaginous texture with a slightly sweet flavor. They are slightly toxic, so should be eaten only in small amounts, especially when raw.[23]
Some species ofPodocarpus are used in systems oftraditional medicine for conditions such asfevers,coughs,arthritis,sexually transmitted diseases, andcanine distemper.[24]
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