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Dacrydium cupressinum

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Dacrydium cupressinum
A mature Dacrydium cupressinum specimen is at centre of the image, surrounded by several apparent Dicksonia fibrosa specimens and areas of tussock grassland in the foreground.
A mature D. cupressinum specimen in the South Island's West Coast Region
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Gymnospermae
Division: Pinophyta
Class: Pinopsida
Order: Araucariales
Family: Podocarpaceae
Genus: Dacrydium
Species:
D. cupressinum
Binomial name
Dacrydium cupressinum
Synonyms
Thalamia cupressina Spreng

Dacrydium cupressinum, commonly known as rimu, is a species of tree in the family Podocarpaceae. It is a dioecious evergreen conifer, reaching heights of up to 60 metres (200 feet) with a stout trunk up to 2 m (6 ft 7 in) in diameter. It is endemic to New Zealand; its range covers the North, South, and Stewart Islands, and it typically inhabits lowland to montane forests. D. cupressinum has an estimated lifespan of 600–800 years, although it may live as long as 1,200 years.

D. cupressinum has a distinctive flavonoid glycoside profile, and has twelve known sesquiterpenes, which include high-levels of the unique sesquiterpene terpene compound of 9-epi-β-caryophyllene. D. cupressinum was first described in 1786 by Daniel Solander and was later given a full description in 1803 by Aylmer Lambert. D. cupressinum's fruits are consumed by various birds such as: bellbirds, kererū, and the tūī. Its fruits also provide an important source of food and vitamins for the endemic flightless parrot, the kākāpō, which will only mate during years of heavy fruiting.

In Māori culture, D. cupressinum is of great importance. D. cupressinum had several medicinal purposes for Māori, and the British missionary, Richard Taylor, described the fruit as "much prized" by the natives. In 1773, the British explorer, James Cook, brewed the first native beer at Tamatea / Dusky Sound, extracting the tips of D. cupressinum's branches, to create what was known as "spruce beer". D. cupressinum's conservation status was assessed by the IUCN Red List in 2013 as "Least Concern", and its population trend was assessed as "Stable".

Description

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Dacrydium cupressinum (rimu) is a species of dioecious evergreen conifer in the family Podocarpaceae, reaching heights of up to 60 metres (200 feet) tall with a stout trunk up to 2 m (6 ft 7 in) in diameter, its trunk is usually smooth and cylindrical.[2][3] Its bark is dark-brown in colour, shedding in large, thick flakes, while the wood is typically a dark-red colour.[2][4] D. cupressinum has an estimated lifespan of 600–800 years, although it may live as long as 1,200 years.[5][6]

Juvenile branches are numerous, slender, with pendulous (hanging) branchlets. In adult specimens, there are fewer branches, spreading, with slender, pendulous branchlets. D. cupressinum's leaves differ in colour, from dark-green to bronze-green, red-green, or in some cases orange in colour, they are typically a yellowish-green colour and are keeled and imbricated (or arranged in an overlapping pattern). Juvenile leaves measure 4–7–(10) mm in length and 0.5–1 mm in width, they are acute, linear (long and narrow) to subulate (slender), slightly curved, and subfalcate (vaguely sickle-shaped) in character.[2][7] Subadult leaves are ascending, incurved, 4–6 mm long, and is rhomboid (diamond-shaped) in character. Adult leaves are similar, but they are appressed (flattened down), measuring 2–3 mm, rigid, and triangular in a cross-section.[2][4]

As a conifer, D. cupressinum does not have flowers, but instead has cones (strobili); male and female cones are first seen on subadult specimens. In male specimens, cones are solitary or paired, terminal, measuring 5–10 mm, and are oblong (tectangular) in character, producing yellow pollen. Its ovules are solitary and are positioned terminally on up-curved branchlets. Its receptacles form a fleshy cup that is red or deep-orange in colour and measures about 1–2 mm in length. Its seeds are dark brown to black in colour, are tiny, and measure about 3–3.8–(4) mm in length.[2][7]

Fruiting takes over a year to fully mature and often coincides with the presence of young female cones. Its fruits are most commonly seen between February and May.[2] D. cupressinum is morphologically a distinctive species, although it is sometimes misidentified with the seedlings of Manoao colensoi, which instead have glossier, less fine leaves.[2] D. cupressinum has a diploid chromosome count of 20.[4]

Phytochemistry

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The foliage of D. cupressinum contains various sesquiterpenes, with their levels showing significant variation in specimens, which is similar to the differences observed in diterpene levels. There are twelve known sesquiterpenes in D. cupressinum that have been identified by a 1985 study, but none of which show any structural similarity to lauren-1-ene.[8] The distinctive diterpene lauren-1-ene has been identified only in the foliage of D. cupressinum, however, the concentration of this compound, along with other diterpenes, which also shows significant variation between specimens.[9]

D. cupressinum contains high-levels of the unique sesquiterpene terpene compounds of 9-epi-β-caryophyllenes.[10] D. cupressinum also has a distinctive flavonoid glycoside profile; it possesses flavonol-3-O-rhamnoglucosides and also, to a lesser extent, 3-0-methyl-myricetin glycosides.[11]

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Taxonomy

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Phylogeny of Dacrydium

D. cupressinum Solander ex Forster

D. balansae Brongniart & Gris

D. nausoriense de Laubenfels

D. araucarioides Brongniart & Gris

D. nidulum de Laubenfels

D. guillauminii Buchholz

D. lycopodioides Brongniart & Gris

D. elatum (Roxburgh) Wallich ex Hooker

D. pectinatum de Laubenfels

D. magnum de Laubenfels

D. xanthandrum Pilger

D. gibbsiae Stapf

D. beccarii Parlatore

D. comosum Corner

D. gracile de Laubenfels

D. cupressinum was first described in 1786 by European naturalists Daniel Solander and Georg Forster, in their publication titled, De Plantis Esculentis Insularum Oceani Australis Commentatio Botanica.[12][13] The British botanist Aylmer Lambert provided a more comprehensive description in his 1803 work, A Description of the Genus Pinus.[14] It was given the name Thalamia cupressina by Kurt Sprengel in an 1826 volume of Systema Vegetabilium.[15][13] D. cupressinum is the type species of the genus Dacrydium.[2] D. cupressinum's cladistic placement can be summarised in the phylogenetic tree at the right.[16]

Evolution

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A 1998 study on the phylogeny of the Podocarpaceae family examined the evolution of D. cupressinum and various other species. The research identified it as sister to a clade consisting of various other Podocarpaceae species, including members of the genus Dacrycarpus. This discovery revealed that D. cupressinum shares a common biological ancestor with these species, while maintaining its evolutionary uniqueness.[17] A 2020 study argued that the Dacrydium genus originated in New Caledonia based on a historical biogeographic reconstruction. Despite this, macrofossil evidence "strongly supports" an Australasian origin and another study argues that Dacrydium originated in Australasia during the Late Cretaceous epoch and dispersed into Southeast Asia in the Early Oligocene.[18]

Etymology

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The etymology of D. cupressinum's genus, Dacrydium, means 'tear drop' and the specific epithet, cupressinum, translates simply to 'cypress'. D. cupressinum is commonly known as rimu and red pine.[1][2] There are a few more names that Māori had for specific parts of the tree; for example, the heartwood was known as: kāpara, māpara, or ngāpara. Its fruit was known as "huarangi", and the seed inside the fruit was known as "matawhanaunga".[19]

Ecology

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A large, greenish to yellowish coloured bird, called the kākāpō, can be seen standing on soil, and looking directly towards the camera.
D. cupressinum's fruits are an important food resource for the endemic flightless parrot, the kākāpō.

D. cupressinum's fruits are an important food and vitamin resource for the endemic flightless parrot, the kākāpō (Strigops habroptilus), which feeds on them.[20] The fruits are particularly high in calcium and vitamins, kākāpō may have adapted to survive despite prolonged periods of limited access to calcium and vitamins. It is likely kākāpō seek out sources rich in these nutrients when their physiological needs are heightened, such as during reproduction in females or growth in chicks.[21] A 2006 study recorded that female individuals would forage on D. cupressinum specimens, and would even climb trunks to collect up to 500–600 g (18–21 oz) of ripened fruit for a single chick every day.[22] The kākāpō will only mate during years of heavy fruiting.[23][24]

Its seeds are dispersed by gravity and by frugivory.[25] Its seeds are well-adapted to be dispersed by birds; the primary birds which disperse the seeds include: bellbirds (Anthornis melanura), kererū (Hemiphaga novaeseelandiae), and the tūī (Prosthemadera novaeseelandiae).[26][27] Other birds, such as white-eyes (Zosterops lateralis) and house sparrows (Passer domesticus), also feed on the fruits fleshy receptacles. Excrements of the wētā (Hemedeina thoracica) are frequently found in seed traps, which could indicate that wētā consume the seeds of D. cupressinum.[28] The most common fungi associated with D. cupressinum are Armillaria novae-zelandiae and Ganoderma applanatum, and to a lesser extant, Rigidoporus concrescens.[29]

Phenology

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D. cupressinum's ovules are pollinated by the wind.[26] D. cupressinum's seeds typically remain dormant for at least 30 days and sometimes much longer prior to germination.[30] A 1964 study estimated that a mature "vigorous" specimen would possess about 18 kg (40 lb) of ripe seed sand receptacles or 1.8 kg (4.0 lb) of clean sound seed (or 200,000 seeds).[5] The seed ripens between March and May, about 15–18 months following pollination and are later dispersed for over 1–2 months, although infertile seeds may continue to be released for a significantly longer duration.[26] The seedlings will not establish themselves where there is moderate to severe root competition and a moderately dense canopy because of under low light intensities.[26]

The growth rates of D. cupressinum vary significantly, especially in its early stages. Height growth often occurs at the expense of diameter growth, with saplings under 10 cm (3.9 in) in diameter reaching up to 12 m (39 ft) tall. In open conditions, annual increments can be 15–30 cm (5.9–11.8 in) in height and 1–3 cm (0.39–1.18 in) in diameter, but growth slows in shaded forest environments, with seedlings growing 1–2 cm (0.39–0.79 in) per year.[5]

Distribution

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A dense, native New Zealand forest, with a river in the foreground of the image and a mountain range in the background. D. cupressinum is the predominant species in the forest, with lesser components of mataī (Prumnopitys taxifolia) and halls tōtara (Podocarpus laetus).
D. cupressinum, along with other predominantly conifer species, growing along the banks of the Haupiri River in the West Coast Region.

D. cupressinum is endemic to New Zealand. Its range covers the North Island, South Island, and Stewart Island. In the South Island, it is uncommon, or not present, in parts of the eastern side of the island, it is almost entirely extinct on the Banks Peninsula, where only a single natural specimen remains.[2] D. cupressinum was previously common around the town of Little River.[31] In the South Island, D. cupressinum's range predominantly covers the western side of the island, and it is largely not present in: Canterbury, Marlborough, Otago, and more generally, the inland South Island.[31][32]

D. cupressinum was likely naturally not present in only a few specific areas of the North Island prior to European colonisation, which includes: Tongariro National Park, Kaingaroa Forest, and the southern parts of Hawke's Bay.[31] On the eastern side of the South Island, its presence is scattered, particularly in the areas in between Blenheim and Waikouaiti. D. cupressinum does occur, however, in smaller groups, such as in Ashley Forest, the Hunter Hills, the Kaikōura Ranges, and only a few other locales.[31]

Habitat

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D. cupressinum typically inhabits lowland to montane forests, it sometimes inhabits near subalpine scrub.[2] D. cupressinum is a dominant or codominant emergent tree in mixed conifer or conifer-angiosperm forests reaching elevations of up to 700 m (2,300 ft) above sea level. D. cupressinum is commonly associated with other conifers in its natural habitat, such as: kauri (Agathis australis), and tōtara (Podocarpus totara), as well as angiosperms, which could include: taraire (Beilschmiedia tarairi) and various Metrosideros species.[1] D. cupressinum has a very wide climatic range and can grow in most soil contents.[31]

Uses

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... this fruit is much prized by the natives, and the smallness of the size is made up by its abundance; this tree produces a resin very bitter, but eatable. The wood also possesses the same qualities, an infusion might be used for beer.

D. cupressinum has played a significant role in both the Māori and European cultures of New Zealand. In 1773, the British explorer, James Cook, brewed the first native beer at Tamatea / Dusky Sound, extracting the tips of the branches of D. cupressinum to create what was known as "spruce beer".[2][33]

The British missionary, Richard Taylor, published a book on the cultural and natural history of New Zealand plants in 1848. Taylor mentioned that D. cupressinum's fruit was "much prized" by the natives and an infusion of the wood was used for crafting beer.[34] D. cupressinum was also a significant source of timber in New Zealand as it was present throughout the lowland areas of the country.[1][35]

In Māori culture

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D. cupressinum is of great importance to Māori.[34] Its fruit, is was abundant during each fruiting season, served as a significant food source to Māori.[19][33] The timber of D. cupressinum was crafted into both war spears and hunting spears. D. cupressinum also possessed some medicinal purposes to Māori, such as, its gum (sap) is very "astringent" and was used as a styptic to stop bleeding from wounds. Its bark was utilised for decoction to treat wounds, and the inner parts of the bark were crushed into a pulp to treat burns. Its leaves was used to treat sores.[19] Another name Māori used for the tree was puaka, which the ethnographer Elsdon Best documented was used by the Te Arawa tribe (or known as iwi in Māori).[36]

Conservation

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D. cupressinum's conservation status was assessed by the IUCN Red List in 2013 as "Least Concern", and its population trend was assessed as "Stable".[1] Its assessment in the New Zealand Threat Classification System was evaluated in 2023 as "Not Threatened".[2] As a native tree, D. cupressinum is protected from logging under law and its natural regeneration is vigorous in locations with mature trees and is further supported by revegetation efforts.[1] The current threats that D. cupressinum possesses are introduced species, such as deer and possums.[1]

See also

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