Amborella

Amborella is a monotypic genus of understory shrubs or small trees endemic to the main

island, Grande Terre, of New Caledonia.[4] The genus is the only member of the
family Amborellaceae and the order Amborellales and contains a single species, Amborella
trichopoda.[5] Amborella is of great interest to plant systematists because molecular
phylogenetic analyses consistently place it as the sister group to all other flowering plants.

Description[edit]
Amborella is a sprawling shrub or small tree up to 8 m high. It
bears alternate, simple evergreen leaves without stipules.[5][6] The leaves are two-ranked, with
distinctly serrated or rippled margins, and about 8 to 10 cm long.[6]
Amborella has xylem tissue that differs from that of most other flowering plants. The xylem
of Amborella contains only tracheids; vessel elements are absent.[7] Xylem of this form has long been
regarded as a primitive feature of flowering plants.[8]

Buds and staminate ("male") flowers of Amborella

The species is dioecious. This means that each plant produces either "male flowers" (meaning that
they have functional stamens) or "female flowers" (flowers with functional carpels), but not both.[9] At
any one time, a dioecious plant produces only functionally staminate or
functionally carpellate flowers. Staminate ("male") Amborella flowers do not have carpels, whereas
the carpellate ("female") flowers have non-functional "staminodes", structures resembling stamens in
which no pollen develops. Plants may change from one reproductive morphology to the other. In one
study, seven cuttings from a staminate plant produced, as expected, staminate flowers at their first
flowering, but three of the seven produced carpellate flowers at their second flowering.[10]
The small, creamy white flowers are arranged in inflorescences borne in the axils of foliage leaves.
[11]
 The inflorescences have been described as cymes, with up to three orders of branching, each
branch being terminated by a flower.[11] Each flower is subtended by bracts.[11] The bracts transition
into a perianth of undifferentiated tepals.[11] The tepals typically are arranged in a spiral, but
sometimes are whorled at the periphery.
Carpellate flowers are roughly 3 to 4 mm in diameter, with 7 or 8 tepals. There are 1 to 3 (or rarely 0)
well-differentiated staminodes and a spiral of 4 to 8 free (apocarpous) carpels. Carpels bear green
ovaries; they lack a style. They contain a single ovule with the micropyle directed downwards.
Staminate flowers are approximately 4 to 5 mm in diameter, with 6 to 15 tepals. These flowers bear
10 to 21 spirally arranged stamens, which become progressively smaller toward the center. The
innermost may be sterile, amounting to staminodes. Stamens bear triangular anthers on short
broad filaments. An anther consists of four pollen sacs, two on each side, with a small sterile central
connective. The anthers have connective tips with small bumps and may be covered with secretions.
  These features suggest that, as with other basal angiosperms, there is a high degree of
[12]

developmental plasticity.[10]
Typically, 1 to 3 carpels per flower develop into fruit. The fruit is an ovoid red drupe (approximately 5
to 7 mm long and 5 mm wide) borne on a short (1 to 2 mm) stalk. The remains of the stigma can be
seen at the tip of the fruit. The skin is papery, surrounding a thin fleshy layer containing a red juice.
The inner pericarp is lignified and surrounds the single seed. The embryo is small and surrounded
by copious endosperm.[13]

Phylogeny[edit]
Currently plant systematists accept Amborella trichopoda as the most basal lineage in
the clade of angiosperms.[14] In systematics the term "basal" describes a lineage that diverges near
the base of a phylogeny, and thus earlier than other lineages. Since Amborella is apparently basal
among the flowering plants, the features of early flowering plants can be inferred by comparing
derived traits shared by the main angiosperm lineage but not present in Amborella. These traits are
presumed to have evolved after the divergence of the Amborella lineage.
One early twentieth century idea of "primitive" (i.e. ancestral) floral traits in angiosperms, accepted
until relatively recently, is the Magnolia blossom model. This envisions flowers with
numerous parts arranged in spirals on an elongated, cone-like receptacle rather than the small
numbers of parts in distinct whorls of more derived flowers.
In a study designed to clarify relationships between well-studied model plants such as Arabidopsis
thaliana, and the basal angiosperms Amborella, Nuphar (Nymphaeaceae), Illicium, the monocots,
and more derived angiosperms (eudicots), chloroplast genomes using cDNA and expressed
sequence tags for floral genes, the cladogram shown below was generated.[15]
extant seed plants  
gymnosperms
   

angiosperms  
Amborella
   

   
Nuphar
   

   
Illicium
   

   
monocots
   

 
magnoliids
 

 
eudicots
 
This hypothesized relationship of the extant seed plants places Amborella as the sister taxon to all
other angiosperms, and shows the gymnosperms as a monophyletic group sister to the
angiosperms. It supports the theory that Amborella branched off from the main lineage of
angiosperms before the ancestors of any other living angiosperms. There is however some
uncertainty about the relationship between the Amborellaceae and the Nymphaeales: one theory is
that the Amborellaceae alone are the monophyletic sister to the extant angiosperms; another
proposes that the Amborellaceae and Nymphaeales form a clade that is the sister group to all other
extant angiosperms.[15]
Because of its evolutionary position at the base of the flowering plant clade, there was support for
sequencing the complete genome of Amborella trichopoda to serve as a reference for evolutionary
studies. In 2010, the US National Science Foundation began a genome sequencing effort
in Amborella, and the draft genome sequence was posted on the project website in December 2013.
[16]

Classification[edit]
Amborella is the only genus in the family Amborellaceae. The APG II system recognized this family,
but left it unplaced at order rank due to uncertainty about its relationship to the
family Nymphaeaceae. In the more recent APG systems, APG III and APG IV, the Amborellaceae
comprise the monotypic order Amborellales at the base of the angiosperm phylogeny.[3][14]

Older systems[edit]
The Cronquist system, of 1981, classified the family:[17][18]
Order Laurales
Subclass Magnoliidae
Class Magnoliopsida [=dicotyledons]
Division Magnoliophyta [=angiosperms]
The Thorne system (1992) classified it:[19][20]
Order Magnoliales
Superorder Magnolianae
Subclass Magnoliideae [=dicotyledons]
Class Magnoliopsida [=angiosperms]
The Dahlgren system classified it:[21]
Order Laurales
Superorder Magnolianae
Subclass Magnoliideae [=dicotyledons],
Class Magnoliopsida [=angiosperms].

Genomic and evolutionary considerations [edit]

Amborella is of great interest to plant systematists because molecular
phylogenetic analyses consistently place it at or near the base of the flowering
plant lineage.[22][23][24] That is, the Amborellaceae represent a line of flowering plants that
diverged very early on (about 130 million years ago) from all the other extant species of
flowering plants, and, among extant flowering plants, is the sister group to the other
flowering plants.[22] Comparing characteristics of this basal angiosperm, other flowering
plants and fossils may provide clues about how flowers first appeared—what Darwin
called the "abominable mystery".[25] This position is consistent with a number of
conservative characteristics of its physiology and morphology; for example, the wood
of Amborella lacks the vessels characteristic of most flowering plants.[5] Further, the
female gametophyte of Amborella is even more reduced than normal
female angiosperm gametophyte.[26]
Amborella, being an understory plant in the wild, is commonly in intimate contact with
shade- and moisture-dependent organisms such as algae, lichens and mosses. In those
circumstances, some horizontal gene transfer between Amborella and such associated
species is not surprising in principle, but the scale of such transfer has caused
considerable surprise. Sequencing the Amborella mitochondrial genome revealed that
for every gene of its own origin, it contains about six versions from the genomes of an
assortment of the plants and algae growing with or upon it. The evolutionary and
physiological significance of this is not as yet clear, nor in particular is it clear whether
the horizontal gene transfer has anything to do with the apparent stability and
conservatism of the species.[27][28]

Ecology[edit]
Amborella is typically dioecious, but has been known to change sex in cultivation.
[5]
 Amborella has a mixed pollination system, relying on both insect pollinators and wind.
[9]

Conservation[edit]
The islands of New Caledonia are a biodiversity hot-spot, preserving many early
diverging lineages of plants, of which Amborella is but one. This preservation has been
ascribed to climate stability during and since the Tertiary (66 to 3 million years ago),
stability that has permitted the continued survival of tropical forests on New Caledonia.
In contrast, drought conditions dominated the Australian climate towards the end of the
Tertiary. Current threats to biodiversity in New Caledonia include fires, mining,
agriculture, invasion by introduced species, urbanization and global warming.[23] The
importance of conserving Amborella has been dramatically stated by Pillon: "The
disappearance of Amborella trichopoda would imply the disappearance of a genus, a
family and an entire order, as well as the only witness to at least 140 million years of
evolutionary history."[29] Conservation strategies targeted on relic species are
recommended, both preserving a diversity of habitats in New Caledonia and ex-situ
conservation in cultivation.[23]