Ixerbaceae
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''Ixerba brexioides'', the sole
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
in the
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
''Ixerba'', is a bushy tree with thick, narrow, serrated, dark green leaves and panicles of white flowers with a green heart. The fruit is a green capsule that splits open to reveal the black seeds partly covered with a fleshy scarlet aril against the white inside of the fruit. ''Ixerba'' is an
endemic Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found else ...
of the northern half of the North Island of New Zealand. Common names used in New Zealand are tawari ( mi, tāwari) for the tree and whakou when in flower. It is assigned to the family
Strasburgeriaceae Strasburgeriaceae is a small family of flowering plants in the order Crossosomatales, only found in New Zealand and New Caledonia. It contains two genera, '' Strasburgeria'' and ''Ixerba''. Both genera have simple, evergreen, alternated leaves, o ...
.


Description


Stem and leaves

Tawari is a small tree of up to 10 m high with a spreading crown. The trunk is usually between 2–4 dm in diameter, and covered by a dark to grayish brown bark. Young branches have few flat-lying pale unicellular T-hairs, while the peduncles,
pedicels In botany, a pedicel is a stem that attaches a single flower to the inflorescence. Such inflorescences are described as ''pedicellate''. Description Pedicel refers to a structure connecting a single flower to its inflorescence. In the absenc ...
,
sepal A sepal () is a part of the flower of angiosperms (flowering plants). Usually green, sepals typically function as protection for the flower in bud, and often as support for the petals when in bloom., p. 106 The term ''sepalum'' was coine ...
s and petals are thickly covered in such hairs, giving them a felty look. The leaves are alternately positioned along the stem and often almost create a whorl at the end of a growth period, with the bud at the end covered in stout triangular scales. These
scales Scale or scales may refer to: Mathematics * Scale (descriptive set theory), an object defined on a set of points * Scale (ratio), the ratio of a linear dimension of a model to the corresponding dimension of the original * Scale factor, a number w ...
have entire margins fringed with simple single celled hairs. The first leaves to appear when growth recommences are an intermediate series between schales and full leaves.
Stipule In botany, a stipule is an outgrowth typically borne on both sides (sometimes on just one side) of the base of a leafstalk (the petiole). Stipules are considered part of the anatomy of the leaf of a typical flowering plant, although in many speci ...
s at the base of the leaf stem are absent. The leaf stalk is stout, fleshy and about 2 cm long. The somewhat fleshy and distinctly leathery simple leaf blades are yellowish to dark green on top and pale green beneath and measure 6-16 × 1–4 cm, are lanceolate to elliptic in shape, while the widest point may be at or beyond midlength, with a pointy tip, and the edges are coarsely serrated. A small gland is present at the tip of each tooth. Young leaves are often reddish and have rolled-in margins. Leaves on young shoots may be relatively narrower. Old leaves discolor to orange or red.


Flowers

Flower buds are formed in March and April and these open from October to the end of December varying according to location and altitude. The
inflorescence An inflorescence is a group or cluster of flowers arranged on a Plant stem, stem that is composed of a main branch or a complicated arrangement of branches. Morphology (biology), Morphologically, it is the modified part of the shoot of sperma ...
sits at the end of the branches in an umbel-like panicle that consists of five to fifteen flowers. Each flower is hermaphrodite, starsymmetric, 2—3 cm in diameter and produces copious amounts of nectar, but apparently does not emit a scent. The five
sepal A sepal () is a part of the flower of angiosperms (flowering plants). Usually green, sepals typically function as protection for the flower in bud, and often as support for the petals when in bloom., p. 106 The term ''sepalum'' was coine ...
s are broadly oval, covered in downy hairs and 5–6 mm long. The five petals are overlapping in the bud, white, felty, spoon-shaped with a narrow base (or claw). The
claw A claw is a curved, pointed appendage found at the end of a toe or finger in most amniotes (mammals, reptiles, birds). Some invertebrates such as beetles and spiders have somewhat similar fine, hooked structures at the end of the leg or tarsus ...
is 1-2 cm long and is inserted on a 5-lobed disc. The five stamens extend well beyond the corolla and alternate with the five lobes of the disc at the centre of the flower. The filaments are very long and white. The anthers are connected to the filaments midlength and open in lengthwise slits towards the middle of the flower. Pollen grains have five slits. The nectar is produced by the ledges of the disc, just above the petal implant. The five styles that rise from the disc are sometimes free from each other at their based and are twisting and fused higher up.


Fruit and seed

The fruit is a leathery capsule, broadly oval in shape. These contain five spaces that split open outwards from the base of the style when the fruit is ripe by April. Each space contains two shiny black seeds that are shaped like a slightly curved tear drop of 6×3 mm, and are partly covered by a conspicuous scarlet aril, both of which starkly contrast with the horny white inside of the fruit (or
mericarp A schizocarp is a dry fruit that, when mature, splits up into mericarps. There are different definitions: * Any dry fruit composed of multiple carpels that separate. : Under this definition the mericarps can contain one or more seeds (the m ...
). Aborted seeds are greyish. The embryo is green. ''Ixerba'' has fifty
chromosome A chromosome is a long DNA molecule with part or all of the genetic material of an organism. In most chromosomes the very long thin DNA fibers are coated with packaging proteins; in eukaryotic cells the most important of these proteins are ...
s (2n = 50).


Taxonomy

''Ixerba'' has been assigned to different families over time. Traditionally it was regarded related to ''
Roussea ''Roussea simplex'' is a woody climber of 4–6 m high, that is endemic to the mountain forest of Mauritius. It is the only species of the genus ''Roussea'', which is assigned to the family Rousseaceae. It has opposing, entire, obovate, gree ...
'' and ''
Brexia ''Brexia'' is a plant genus assigned to the Celastraceae. It is a dense evergreen shrub or small tree of usually around 5  m high, with alternately set, simple, leathery leaves with a short leaf stem and lanceolate to inverted egg-shaped ...
'' and has been combined with them in the
Brexiaceae ''Brexia'' is a plant genus assigned to the Celastraceae. It is a dense evergreen shrub or small tree of usually around 5  m high, with alternately set, simple, leathery leaves with a short leaf stem and lanceolate to inverted egg-shaped l ...
. These three have been placed in different other famililies, such as the
Escalloniaceae Escalloniaceae is a family of flowering plants consisting of about 130 species in seven genera. In the APG II system it is one of eight families in the euasterids II clade (campanulids) that are unplaced as to order. More recent research has prov ...
, but doubt remained about their proper placement. After genetic analysis showed that ''Ixerba'' was not very much related to either ''Roussea'' or ''Brexia'', it was placed in the monogeneric family Ixerbaceae. However, further testing revealed that ''Ixerba'' was closely related to the New Caledonian genus ''
Strasburgeria ''Strasburgeria robusta'' is an evergreen tree with large toothed leaves and large but rather inconspicuous, single, pendulant flowers in a gloomy colorscheme of yellowish with brown markings, with about ten sepals, five petals, ten stamens, a ve ...
'' and it was assigned to the
Strasburgeriaceae Strasburgeriaceae is a small family of flowering plants in the order Crossosomatales, only found in New Zealand and New Caledonia. It contains two genera, '' Strasburgeria'' and ''Ixerba''. Both genera have simple, evergreen, alternated leaves, o ...
by the
APG III APG is an abbreviation with several different meanings: * Aberdeen Proving Ground, a United States Army installation in Aberdeen, Maryland, also ** Phillips Army Airfield, the airfield of the above, from its IATA airport code * Aboriginal Provisiona ...
in 2009.


Phylogeny

Fossil remains of ''Ixerba'' have been found from the Middle
Miocene The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...
. Recent phylogenetic analysis resulted in the following tree.


Etymology

The genus name ''Ixerba'' is an anagram of ''Brexia'', a treelet from Madagascar and the coast of East Africa, to which it bears a superficial likeness. The species epithet ''brexioides'' means "like ''Brexia''".


Distribution

Tawari is a species
endemic Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found else ...
to the North Island of New Zealand, roughly north of a line crossing Taupo. It is said to be widespread from about
Kaitaia Kaitaia ( mi, Kaitāia) is a town in the Far North District of New Zealand, at the base of the Aupouri Peninsula, about 160 km northwest of Whangārei. It is the last major settlement on State Highway 1. Ahipara Bay, the southern end of ...
south to
Waitomo Waitomo is a rural community in the King Country region of New Zealand's North Island. There are several solutional cave systems in the area around the village, which are popular tourist attractions. Restaurants and accommodation are centred i ...
, and
Te Urewera Te Urewera is an area of mostly forested, sparsely populated rugged hill country in the North Island of New Zealand, a large part of which is within a protected area designated in 2014, that was formerly Te Urewera National Park. Te Urewera is ...
, including the Tutamoe Ranges, the
Waitākere Ranges The Waitākere Ranges is a mountain range in New Zealand. Located in West Auckland between metropolitan Auckland and the Tasman Sea, the ranges and its foothills and coasts comprise some of public and private land. The area, traditionally kno ...
,
Waipoua Forest Waipoua Forest is a forest, on the west coast of the Northland Region of New Zealand's North Island. It preserves some of the best examples of kauri forest remaining in New Zealand. It is notable for having two of the largest living kauri t ...
, Te Moehau on the
Coromandel Peninsula The Coromandel Peninsula ( mi, Te Tara-O-Te-Ika-A-Māui) on the North Island of New Zealand extends north from the western end of the Bay of Plenty, forming a natural barrier protecting the Hauraki Gulf and the Firth of Thames in the ...
, higher elevations on the
Barrier Islands Barrier islands are coastal landforms and a type of dune system that are exceptionally flat or lumpy areas of sand that form by wave and tidal action parallel to the mainland coast. They usually occur in chains, consisting of anything from a f ...
and elsewhere around the
Bay of Plenty The Bay of Plenty ( mi, Te Moana-a-Toi) is a region of New Zealand, situated around a bight of the same name in the northern coast of the North Island. The bight stretches 260 km from the Coromandel Peninsula in the west to Cape Runaw ...
.


Ecology

Tawari can often be found together with the New Zealand kauri ''Agathis australis'' in the
understory In forestry and ecology, understory (American English), or understorey (Commonwealth English), also known as underbrush or undergrowth, includes plant life growing beneath the forest canopy without penetrating it to any great extent, but abo ...
of the lowland rain forest. It is common in cloud forest up to 700 m too. The species has a preference for shaded or sheltered locations, often in permanently damp soil and near streams. It is locally abundant but mostly individuals are far between. It is common in regenerating forests along with tawheowheo ''Quintinia serrata''. Tawari has a mass flowering strategy, attracting a range of visitors with the abundant nectar it produces. Each flower produces on average 18 μl of nectar with an average concentration of 20% brix. Pollination is mainly by birds, however in modified forest where birds a few in number, flower visitation is most frequently by large flies and moths, followed by honey bees, bumblebees, native bees, wasps and beetles. Floral morphology, such as its wide spreading anthers, suggest adaptation to pollination by birds. Seeds capsules develop between January and April each year, and dehisce to reveal up to ten glossy, purple-black seeds that are partly covered by an orange fleshy aril. Seeds are primarily dispersed by birds, with reports including
kererū The kererū (''Hemiphaga novaeseelandiae'') or New Zealand pigeon is a species of pigeon native to New Zealand. Johann Friedrich Gmelin described the bird in 1789 as a large, conspicuous pigeon up to in length and in weight, with a white br ...
, pōpokotea, hihi, and
kākā The New Zealand kākā (''Nestor meridionalis'') is a large species of parrot of the family Nestoridae found in New Zealand's native forests. The species is often known by the abbreviated name kākā, although it shares this name with the rece ...
.


Use

Tawari is a rich source of nectar, which is often so abundant that bees produce
monofloral honey Monofloral honey is a type of honey which has a distinctive flavor or other attribute due to its being predominantly from the nectar of a single plant species. It is stored and labeled separately so as to command a premium price. While there may ...
, that is the honey is chiefly made of nectar from tawari flowers. Tawari honey is regularly for sale from food shops in New Zealand. The nectar contains a lot of fructose and much water. The honey may be thin as a consequence, but nonetheless crystallizes quickly. The honey has a light colour and tastes a bit like butterscotch. It contains relatively few pollen, and if at least 20% of the pollen is tawari, it can be marketed as tawari honey. Bark can produce a black dye that is known to be used on flax. Māori people traditionally used the flowers to make necklaces and other adornments to be worn at festivities.


Cultivation

Tawari is known to be very difficult to grow in gardens. It needs a sheltered location, humus-rich soil and good drainage, but the soil should never dry out. The species probably depends on a mycorrhiza, and if planted next to Kapuka ''
Griselinia littoralis ''Griselinia littoralis'', commonly known as kapuka, New Zealand broadleaf or pāpāuma, is a fast-growing small to medium-sized evergreen tree that is native to New Zealand. Description ''Griselinia littoralis'' is a hardy evergreen shrub th ...
'', tawari does much better.


External sources

Photo series illustrating many details of the anatomy


References

{{Taxonbar, from=Q147889 Flora of the North Island Trees of New Zealand Monotypic rosid genera Crossosomatales