Prumnopitys taxifolia

Prumnopitys taxifolia is a species of dioecious evergreen conifer in the family Podocarpaceae, reaching heights of tall, with a trunk of up to in diameter. Branches are initially ascending and eventually become spreading, forming a domed or rounded crown. The bark is thin and smooth (particularly on younger trees); it has spots, similar to hammer marks, which are red or purplish-brown in colour when fresh, weathering into a dark brown to black colour, and peeling freely into thick flakes. The outer bark itself can vary in colour from a dark brown to an almost black colour. The wood is very dense, hard, and a dark brown to a yellow-brown colour. P. taxifolia has an average lifespan of 600 years, although a specimen from Tongariro National Park is estimated to have lived for more than 1000 years. Prumnopitys taxifolias young form is distinct and the species profoundly exhibits heteroblasty (a significant change in a plant's form). Its young form, resembles a tangled shrub, is so distinct that the first botanists in New Zealand initially thought they were separate species. Unlike the related miro (Pectinopitys ferruginea), P. taxifolia has a distinctive and long-lasting young stage, which is a tangle of slender, flexible, divaricating branchlets that are interspersed with a scattering of leaves. After several decades, the adult tree begins to grow out of the top of the young shrub and the divaricating branchlets will drop off. It is unknown why this species developed this characteristic; one possibility is that the tangled branches are designed to create a warm temperature inside the shrub, hence the species could survive in colder climates, or it may serve as a defence against browsing animals. The young stage can last for up to 60 years. Young leaves are 5–10 × 1–2 mm long and brown to pale yellow in colour, narrow and tapering, with a pointed tip, and they can have long whip-like shoots. Adult leaves are 10–15 × 1–2 mm long, dark green in colour, glaucous, subdistichous and often apiculate in character. They generally have a broad shape, but can be more pectinate in character, and are (6–)10–20 mm long, (1–)1.5–2 mm wide. Leaves are widest near the middle, with nearly parallel sides. The midrib is raised on a bluish-green surface, where there are waxy, pale bluish-green bands of stomata. The Canadian botanist, James Emory Eckenwalder, hypothesised that the leaves of P. taxifolia potentially contain unidentified physiologically active compounds. Like all conifers, P. taxifolia does not produce flowers or true fruits, but instead produces pollen and seed in cones, or 'pseudo-fruits'. The species is dioecious, with male and female reproductive structures on separate trees. The ovules are found on short axillary branches, with 3–10 per a 40 mm long spike. P. taxifolias male pollen cones are found on nearly leafless lateral and axillary spikes near the ends of foliage branches. Each spike is yellow in colour, and can have up to 10–30 cones per spike, each 5–12 mm long and approximately 2 mm wide. The seed cones take 12–18 months to mature and are found throughout the year. They are round to oval in shape, fleshy, globe-shaped or nearly spherical, purplish-black in colour, and about 8–10 mm wide. P. taxifolia has a diploid chromosome count of 38. Phytochemistry The sap of P. taxifolia contains matairesinol, a compound which has been investigated in cancer research. In addition to matairesinol, the heartwood of the tree contains numerous other compounds, which includes, but is not limited to aromadendrin, kaempferol, quercetin, and genistein. Foliage oils of the tree were investigated in 1987 in the scientific journal Phytochemistry. Researchers found seven sesquiterpene hydrocarbons and two oxygenated sesquiterpenes, ten diterpene hydrocarbons, and two oxygenated diterpenes (including ferruginol) were identified. ==Gallery==

File:Prumnopitys taxifolia - Michael Berardozzi - 417974914.jpeg|alt=P taxifolia's hammer-like scar patterned bark.|P. taxifolia has smooth brown bark, with hammer mark-like patterning throughout it. File:Prumnopitys taxifolia - Colin Meurk - 363506166 (cropped).jpeg|alt=Globe-shaped fruits (both ripened and unripened) and foliage of P. taxifolia.|P. taxifolias purple-black-coloured globe-shaped drupes. File:Prumnopitys taxifolia - Leon Perrie - 242595429.jpeg|alt=Strobili and foliage of P. taxifolia.|The pollen cones are 30–50 mm long, with up to 30 cones per spike. File:Young mataī tree - adjacent to Mawaihakona Stream.jpg|alt=P. taxifolia in its young stage; it is a wiry brown-coloured shrub.|P. taxifolia before exhibiting heteroblasty. It resembles a divarciating-wiry shrub. ==Taxonomy==

Classification }} }} }} }}}} Prumnopitys taxifolia is a member of the genus Prumnopitys, which consists of five species across Australasia and South America. P. taxifolia is categorised in the subgenus Prumnopitys. In 2019 the British botanist Chris Page transferred six species from Prumnopitys into a genus Page newly described as Pectinopitys. Page retained three species within the genus Prumnopitys, including P. taxifolia. He based his move on cladistic, morphological, and molecular analysis, concluding that Pectinopitys was distinct enough for it to be described as a new genus, rather than retaining it in Prumnopitys. Page also mentioned that Pectinopitys has a diploid chromosone count of 2n =  36, while Prumnopitys has a count of 2n = 38. Khan et al. (2022) revealed that Prumnopitys montana of South America, is the closest relative to P. taxifolia. A 2012 study based on rbcL data sequencing revealed that both species are nearly allied to species within the genus Pectinopitys (formerly placed within the same genus). History Prumnopitys taxifolia was first described as Dacrydium taxifolium by the British botanist Aylmer Lambert in his work A Description of the Genus Pinus. It is the basionym (original scientific name) of the species. P. taxifolia was described as Dacrydium mai and Podocarpus spicatus in 1838. The species was attempted to be renamed to Stachycarpus spicatus in 1891 by the French botanist Philippe Édouard Léon Van Tieghem. American botanist David John de Laubenfels described it as Prumnopitys taxifolia in 1978, which is its currently recognised binomial name. Etymology The etymology of P. taxifolias genus name, Prumnopitys, is uncertain. One possibility suggests the name means 'plum-fruited pine', which derives from the Greek , which means 'plum', and which means 'fir' or 'pine'. Gledhill (2008) disputes this, and states the genus name means 'hindmost-pine'. The specific epithet refers to the 'yew-like' leaves of the species, deriving from the Latin , which is in reference to Taxus baccata. The species is commonly known as the mataī and or black pine. The word is likely a combination of the Māori language words , meaning 'unripe', and '' meaning 'fermented' or 'sour'; likely referring to the bitter taste of resin exuded by the trunk. In 1908 the New Zealand ethnographer Elsdon Best recorded that young trees are also known in Māori as , , and . Herbert Williams recorded that the trees are also known as . ==Ecology==

, has been recorded as a disperser of P. taxifolias fruits. Prumnopitys taxifolias fruits are dispersed by fruit-eating animals (frugivores). The fruits of P. taxifolia reflect some ultraviolet radiation, which is visible to some species of birds and vertebrates, although they do not reflect as strongly as Dacrycarpus dacrydioides. A 1989 study, published in the New Zealand Journal of Ecology, recorded the kererū (Hemiphaga novaeseelandiae), kōkako (Callaeas wilsoni), tūī (Prosthemadera novaeseelandiae), and weka (Gallirallus australis) as dispersers of the fruits of P. taxifolia. The fruits are also eaten by introduced blackbirds (Turdus merula) and song thrushes (T. philomelos). The kākā (Nestor meridionalis) have been recorded destroying the fruits of P. taxifolia. The fruits of P. taxifolia are eaten by the endemic moth, Heterocrossa iophaea. P. taxifolia is a host to the native beetles, Ambeodontus tristis and Prionoplus reticularis. A 2012 study found that P. taxifolia fruits can still germinate after being excreted by feral pigs (Sus scrofa), with a success rate of 57 percent. A 2021 study hypothesised P. taxifolias fruits were consumed by the extinct little bush moa (Anomalopteryx didiformis), although in low quantities when compared to other native species at the studied site at Borland Burn in Fiordland National Park. P. taxifolia can play host to epiphytes (plants that grow on other plants) of the genus Astelia. In 2023 a study published in MycoKeys, examined three Chaenothecopsis lichen species that grow primarily on the exudates of P. taxifolia. One of these species, C. matai, was named after the tree. P. taxifolia is sensitive to fire and has a moderate flammability rate. Reproduction Like all conifers, the ovules of P. taxifolia are pollinated by the wind. The cones of P. taxifolia have an elongated central axis with about twelve ovules. Each ovule is positioned beneath arranged bracts (specialised leaves). At the base of each bract, there is a groove that forms a small gutter around the ovule. During pollination, pollen collects into the grooves before and after a drop of secretion appears, helping to capture pollen efficiently. ==Distribution==

Prumnopitys taxifolia is endemic to New Zealand; its range primarily covers the North and South Islands. It also occurs on Stewart Island, but it is extremely uncommon there. In The Forest Flora of New Zealand, published in 1889, Thomas Kirk reported only one specimen growing on the southern side of Halfmoon Bay. P. taxifolia commonly coincides with southern beech (Nothofagus) species in the South Island's West Coast Region. Kirk noted in his 1889 revision of New Zealand plants that the species was very common in the central areas of the North Island and in the western and south-western parts of the South Island. The conservation status of P. taxifolia was assessed by the IUCN Red List in 2013 as "Least Concern", and its population trend was evaluated as "Stable". Its assessment in the New Zealand Threat Classification System was evaluated in 2023 as "Not Threatened". Habitat Prumnopitys taxifolia is typically found in lowland and montane forests. It occurs from above sea level. It is commonly found on mountain slopes and grows in fertile, well-drained soils. In the eastern regions of both islands, where rainfall is relatively low, the tree can still grow well in the drier climates. The largest specimens of P. taxifolia have been found growing in deep pumiceous soils or in lime-rich soils beneath limestone escarpments. Similar to Pectinopitys ferruginea, the tree is commonly found on alluvial soils in the West Coast Region of the South Island. P. taxifolia is a slow-growing, wind-resistant tree that is highly shade-tolerant, which can regenerate fully under closed canopy. ==Uses==

In Māori culture The timber is highly valued by the indigenous Māori people for its durability and hardiness and had a variety of uses, including being a prized timber for building (canoes), carving and building tools, and making traditional instruments, such as a , a trumpet, which signals a war. Other uses of the timber included it being used for bowls, containers, combs, spades, troughs, and weapons. The young divaricating form of the tree has flexible stems which were used by Māori to make (eel pots). P. taxifolias fruits are edible and are described as sweet, fragrant, and rather slimy in taste. Another soruce from 1853 described them as "very glutinous". They were collected by Māori in large baskets, and then eaten. According to Best, the Māori seem recognise two different varieties of the tree. In Māori mythology, the blood-red colour of the inner bark of mature trees attributed to the blood of Tunaroa, a mythological god, who was killed by Māui, the cultural hero. In European culture Although P. taxifolia had some uses as a timber, it was historically considered inferior to the kauri (Agathis australis) and rimu (Dacrydium cupressinum). P. taxifolias timber is noted for its strength and durability, and was historically used for building bridges, house building, especially for flooring and weatherboards, and railway sleepers. It was also used, to a lesser degree, for furniture-making. Although commonly known as black pine to early European settlers, an 1853 source mentions that the tree was also known as maiberry to early settlers. Another important resource from the tree is the sap exuded from the trunk, which is also edible, and is described as sweet and bitter in taste, similar to the fruits. The sap produces a rich brown-coloured beverage, termed mataī beer by Europeans. The sap is even brewed into a modern-day alcoholic beverage. ==References==

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