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Adult tree

Not Threatened (NZ TCS)[1]
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Clade: Asterids
Order: Apiales
Family: Araliaceae
Genus: Pseudopanax
Species:
P. crassifolius
Binomial name
Pseudopanax crassifolius
(Sol. ex A.Cunn.) C.Koch

Pseudopanax crassifolius, also known as horoeka or lancewood, is a heteroblastic tree belonging to the family Araliaceae. It is endemic to New Zealand and found throughout the country from sea level up to about 750 m in lowland to montane forests and shrublands.[2][3]

The juvenile form, which lasts between 15 and 20 years, is easily recognized. The leaves are stiff and leathery with a prominent central rib, about 1 cm wide and up to 1 m long with irregular teeth, all growing downwards from a central stem. The young trunk has characteristic vertical swollen ridges.

As the tree gets older the stem begins to branch, producing a bushy top, and the leaves become wider and shorter, losing their teeth. It is only when the tree is mature that it adopts a typical tree shape.

The difference between the juvenile and adult tree's appearance is so drastic that the two forms were initially described as separate species and even genera by botanists,[4][5] including the botanists accompanying Captain James Cook's during his first voyage to New Zealand.[6]

One of the theories about this curious change of appearance is that the young plant had to protect itself against browsing by the moa, the giant flightless bird that once roamed New Zealand's bush. Once above moa height, it was out of danger and turns into a "regular" tree. A study of leaf colour development in P. crassifolius found that leaves of seedlings would blend with leaf litter, while juvenile leaf colouration would draw attention to their spines. A closely related Chatham Island species, which evolved in the absence of moa, did not display these changes.[7]

Closely related is Pseudopanax ferox, the toothed lancewood. It is similar in appearance to P. crassifolius except the leaves are more abundant and severely toothed, resembling remotely a bandsaw blade.

Description

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Pseudopanax crassifolius is a heteroblastic plant, meaning that it has different leaf forms and growth habit depending on its growth stage.[8][9] These growth stages, which are largely related to the plant height, are: seedling, juvenile, transitional, and adult.[9]

A juvenile tree is single stemmed with long and narrow deflexed leaves,[2][10] typically below the height of 3–5 m.[9][7]

As an adult, P. crassifolius can reach a height of 15 m.[2] It appears as a round-headed tree due to being multi-branched,[2] making it look more like a "typical tree."

Leaves

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Seedling leaves are smaller and thinner compared to the following growth stages, but they gradually become longer as the plant grows out from the leaf litter, starting from cotyledon.[9] Leaves are initially green, but become more mottled brown and leathery as the plant grows.[9]

Juvenile leaves, which are produced up to 4 m or before the tree branches,[9] are especially long and narrow (1 m long by 1–1.5 cm wide)[11] – similar in shape to a lance.[5] They are dark green with a sharply toothed margin and prominent yellow or orange midrib[9][12] Leaves are also rigid and straight, but angled downwards from the stem.[2]

Intermediate leaves appear once the tree has branched, typically above the height of 4–5m.[9] They are an intermediate between juvenile and adult leaves, being shorter and wider than juvenile leaves.[9]

Adult leaves are shorter and wider (10–20 cm long by 2–3 cm wide), with a less prominent midrib and toothed or entire margin.[9][11] Additionally, leaves are simple, alternate,[13] and grow outwards or upwards from the branch.[14] They typically appear once the tree is above 5 m and branched.[9] Adult leaves also have a greater frost resistance than juvenile leaves.[15]

Trunk

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The trunk of a juvenile is slender,[12] straight,[5] unbranched,[16] and ridged down its length.

For adult trees, the trunk can be up to 50 cm in diameter.[4] It has horizontal lenticels[13] and smooth, grey[12] or mottled bark.[17] At this life stage, the top of the tree has branched and the juvenile leaves have fallen off, producing a rounded head with a naked stem.[12]

Flowers

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The tiny, 4 mm diameter, pale yellow to green[11] flowers are clustered in large terminal umbels, 30 cm across.[18] The flowers are bisexual, so have five stamens and five styles.[4]

Fruits

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The fleshy fruits, which are usually dark purple to black when ripe, are 3–4 mm in diameter and round. They contain five seeds per berry,[19] with the five locules each containing one seed.[10]

Range

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Natural global range

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Pseudopanax crassifolius is endemic to New Zealand.[3]

New Zealand range

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Within New Zealand, P. crassifolius is abundant and widespread in North Island, South Island, and Stewart Island.[2]

Habitat

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Pseudopanax crassifolius occurs from sea level up to about 750 m in lowland to montane forests and shrublands.[2] It does not have a strong affinity for wet habitats, as it usually occurs in uplands/non wetlands.[20] Instead, this species is often growing along forest margins or gaps, in regenerating forests, and as a subcanopy or canopy tree.[10]

Specifically, P. crassifolius is a common subcanopy or canopy species in a Kāmahi - silver fern forest (Beech-broadleaved-podocarp forest Alliances),[21] Hard beech – kāmahi forest (Beech forest Alliances),[22] Silver beech-red beech-kāmahi forest (Beech-broadleaved forest Alliances),[23] Silver fern - mahoe forest,[24] Towai – tawa forest,[25] and Kāmahi-podocarp forest (Broadleaved-podocarp forest Alliances including kauri).[26]

Ecology

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Phenology and reproduction

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The juvenile form of P. crassifolius lasts between 15 and 20 years before it transitions into its adult form.[8][12]

The tree almost only flowers and fruits when it is an adult.[9] Its bisexual flowers are in full bloom from January to April,[27][28] and are pollinated by insects.[29] Fruits are produced in January to April, ripen through the winter, and persist until August.[10] These fruits are eaten and then dispersed by birds.[10] Passing through a bird's gut also strips the seed of its fleshy mesocarp coat, which helps it germinate[30] Subsequently, these seeds were found to germinate after five weeks[10] but this may be delayed if the fleshy mesocarp is still intact.[30]

Pseudopanax crassifolius is evergreen.[10]

Predators, Parasites, and Diseases

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Kererū feeding on Pseudopanax crassifolius berries
 
Kererū feeding on Pseudopanax crassifolius berries (img placeholder)

Birds

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The berries serve as an important food source for many birds.[8] This includes birds like the hihi,[31] kokako,[32] whitehead, tūī,[8] kererū, silvereye, New Zealand bellbird,[10][19] New Zealand kākā, brown creeper, grey warbler, tomtit, blackbird, and chaffinch.[33]

Its flowers also provide nectar, which honey-eating birds like the hihi,[34] tūī, and New Zealand bellbird use for food.[33]

As well as the fruit, kererū will also eat leaf buds of P. crassifolius.[33]

Birds are also known to forage for invertebrates on P. crassifolius. This includes the kākā, rifleman, brown creeper, grey warbler, tomtit, New Zealand fantail, silvereye, and New Zealand bellbird.[33]

Although extinct, it is also believed that the moa would have once predated this plant, specifically its leaves.[7]

Invertebrates

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Bees introduced and native to New Zealand, like the western honey bee[35] or Lasioglossum sordidum,[36] will use this species for nectar when it is flowering. As a result, beekeepers believe P. crassifolius to be good bee forage, and recommend it on farms.[28]

 
Eumetriochroa panacivagans leaf mine on Pseudopanax crassifolius leaf

Moth species Acrocercops panacivagans and Eumetriochroa panacivagansis are known to use Pseudopanax species like P. crassifolias as a host.[37][38] This is evident through the mines they create in the leaves.[37][38] Another moth species, Izatha blepharidota, will also take this plant as a host, with its larvae successfully reared and feeding on the dead branches of P. crassifolius.[39]

The New Zealand praying mantis will deposit its ootheca on P. crassifolius.[40]

Many beetle species are known to parisitise P. crassifolius, commonly by using it as a host. Weevil species from the family Curculionidae and Belidae have larvae that live in dead, or dying, woody parts of P. crassifolius, including Dendrotrupes vestitus, Ectopsis ferrugalis, Platypus apicalis, Scolopterus penicillatus, Stephanorhynchus curvipes,[41] Psepholax simplex, Psepholax mediocris, Psepholax crassicornis, Mesoreda sulcifrons, Crisius semifuscus, Clypeolus pascoei,[42] Phronira osculans,[43] and Dendrotrupes costiceps[44] and Aralius wollastoni.[45] Two longhorn beetle species Tetrorea cilipes and Hexatricha pulverulenta from the family Cerambycidae have larvae that tunnel in the woody parts of trees, and are believed to use P. crassifolius as a host.[41][46]

Scale insects will use P. crassifolius as a host plant by living on its leaves, petioles, fruit, and stems. Notably, Trioza panacis, also known as the lancewood psyllid, will feed on the leaves of P. crassifolius and a few other Pseudopanax species.[47] Additional scale insect species include those from the family Diaspididae, such as Leucaspis gigas, Leucaspis mixta, Leucaspis morrisi,[48] and Hemiberlesia lataniae.[49] From the family Coccidae, this includes the soft scale insect species Ctenochiton viridis and Coccus hesperidum.[50] Felted scale insects from the family Eriococcidae, like Scutare fimbriata and Eriococcus parabilis.[51] Finally, from the family Rhyparochromidae, Woodwardiana evagorata will feed on the leaves of P. crassifolia.[52]

Mammals

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Pseudopanax crassifolius leaves are palatable and preferred by mammals introduced to New Zealand, such as cattle,[53] deer, brushtailed possums, and feral goats.[32][54][55] The leaves of a mature P. crassifolius tree are known to have a higher nutritive value compared to its juvenile form, which may make them more appealing to these mammals.[56] Alongside leaves, the berries of P. crassifolius are eaten by brushtail possums.[57] Rats will also predate P. crassifolius, as they are known to eat its seeds.[58]

Plants

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The parasitic plant Dactylanthus taylorii has been reported to grow on the roots of P. crassifolius, where it steals nutrients by causing the roots to enlarge.[59]

Fungi

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Chaenothecopsis schefflerae will live on P. crassifolius.[60]

 
Pseudopanax crassifolius × lessonii hybrid leaf morphology variation (placeholder img)

Hybrids

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Pseudopanax crassifolius commonly hybridises with Pseudopanax lessonii.[61] Given the two species' dissimilar appearances, they produce a range of morphologically diverse offspring with leaves often appearing different to both parent species.[61] Despite this, Pseudopanax crassifolius hybridises with P. lessonnii significantly more frequently than with morphologically similar Pseudopanax species, such as Pseudopanax ferox or Pseudopanax linearis.[61] One theory relating to this process is that P. crassifolius is more closely related to P. lessonnii and its allies than other Pseudopanax species.[61]

Evolution

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The heteroblastic nature and unique appearance of this species has lead many to question why it has evolved this way. A prominent theory is that P. crassifolius evolved its unique traits as to deter or prevent moa browsing. A P. crassifolius seedling emerging from the leaf litter is typically brown, eventually producing green leaves as it grows. small seedlings' brown colour is thought to have helped the seedling camouflage in the leaf litter it emerges from, potentially helping it avoid being seen by moa. Additionally, the Chatham Island lancewood, which is a very close relative to P. crassifolius endemic to the Chatham Islands, does not share

Uses

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Urban design

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Pseudopanax crassifolius is a popular plant in urban design not only for its iconic and charismatic appearance, but also for its ecological benefits. Its unique look can be used to create textural contrast amongst other plants.[62] Because of its thin and tall juvenile form, this species is also great in small spaces.[63] For these reasons, this species is recommended along streets, entrance ways, traffic islands, or in the central island of roundabouts.[64]

Early European settler use

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The midribs of juvenile leaves used to be used as bootlaces and in fixing horse bridles and harnesses.[8] Its dense, light brown wood can be used for fence posts or piles,[12] but is not known to be very durable.[8] In fact, its wood was once used for timber in Otago.[8] Additionally, the flexible trunk would sometimes be used as a whip.[8]

Traditional Māori use

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Māori from the South Island would use P. crassifolius leaves to make paint brushes for rock paintings.[8] Due to its flexibility, the trunk of a young P. crassifolius could be twisted and used as a tokotoko (ceremonial walking stick).[65]

In Māori culture

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All three lancewood species (P. crassifolius, P. ferox, and P. linearis) are known as horoeka by Māori.[8] Māori thought that a flowering P. crassifolius indicated that birds would be abundant the following year because they would be attracted to the ripened fruit.[8]

Conservation

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Pseudopanax crassifolius is a non-threatened species, meaning it has large, stable populations.[1]

Mammalian browsing is a current threat to P. crassifolius. For example, brushtailed possum browsing is known to cause defoliation and death to mature trees.[8] Cattle browsing also impacts the abundance of P. crassifolius, with one study finding the plant only in areas devoid of cattle.[53]

References

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  65. ^ "Native trees and their connections with people" (PDF). Department of Conservation. Department of Conservation. Retrieved 21 September 2024.
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