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_AMBORELLA_ is a monotypic genus of understory shrubs or small trees endemic to the main island, Grande Terre , of New Caledonia . The genus is the only member of the family AMBORELLACEAE and the order AMBORELLALES and contains a single species , _AMBORELLA TRICHOPODA_. _Amborella_ is of great interest to plant systematists because molecular phylogenetic analyses consistently place the genus at or near the base of the flowering plant lineage.

CONTENTS

* 1 Description * 2 Phylogeny

* 3 Classification

* 3.1 Older systems

* 4 Genomic and evolutionary considerations * 5 Ecology * 6 Conservation * 7 Gallery * 8 References * 9 Further reading * 10 External links

DESCRIPTION

_Amborella_ is a sprawling shrub or small tree up to 8 m high. It bears alternate or decussate , simple evergreen leaves without stipules . The leaves are two-ranked, with distinctly serrated or rippled margins, and about 8 to 10 cm long.

_Amborella_ has xylem tissue that differs from that of most other flowering plants. The xylem of _Amborella_ contains only tracheids ; vessel elements are absent. Xylem of this form has long been regarded as a "primitive " feature of flowering plants. _ 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. 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.

The small, creamy white, flowers are arranged in inflorescences borne in the axils of foliage leaves. The inflorescences have been described as cymes , with up to three orders of branching, each branch being terminated by a flower. Each flower is subtended by bracts . The bracts transition into a perianth of undifferentiated tepals . 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 developmental plasticity.

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.

PHYLOGENY

Currently plant systematists accept _ Amborella trichopoda_ as the most basal lineage in the clade of angiosperms . 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.

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.

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.

CLASSIFICATION

_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 .

OLDER SYSTEMS

The Cronquist system , of 1981, classified the family: Order Laurales Subclass Magnoliidae Class Magnoliopsida Division Magnoliophyta

The Thorne system (1992) classified it: Order Magnoliales Superorder Magnolianae Subclass Magnoliideae Class Magnoliopsida

The Dahlgren system classified it: Order Laurales Superorder Magnolianae Subclass Magnoliideae , Class Magnoliopsida .

GENOMIC AND EVOLUTIONARY CONSIDERATIONS

_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. 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. 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". 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 .

_Amborella_ in the wild, being an understory plant, commonly is in intimate contact with shade- and moisture-dependent organisms such as algae, lichens and mosses. In the circumstances, it is not surprising in principle to find that there should have been horizontal gene transfer between _Amborella_ and such associated species, but the scale of such transfer has in fact 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.

ECOLOGY

_Amborella_ is typically dioecious , but has been known to change sex in cultivation. _Amborella_ has a mixed pollination system, relying on both insect pollinators and wind.

CONSERVATION

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. 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." Conservation strategies targeted on relic species are recommended, both preserving a diversity of habitats in New Caledonia and ex-situ conservation in cultivation.

GALLERY

*

_ Amborella trichopoda_ plant *

Young leaves with serrated margins

REFERENCES

* ^ "IPNI Plant Name Query Results for Amborella". _The International Plant Names Index _. Retrieved 2013-09-03. * ^ _A_ _B_ _C_ Angiosperm Phylogeny Group III (2009). "An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III". _Botanical Journal of the Linnean Society_. 161 (2): 105–121. doi :10.1111/j.1095-8339.2009.00996.x . * ^ "IPNI Plant Name Query Results for _ Amborella trichopoda_". _The International Plant Names Index _. Retrieved 2013-09-03. * ^ Jérémie, J. (1982). "Amborellacées". In A. Aubréville and J. F. Leroy. _Flore de La Nouvelle-Calédonie et Dépendances_ (in French). 11. Paris: Muséum National d’Histoire Naturelle. pp. 157–160. CS1 maint: Uses editors parameter (link ) * ^ _A_ _B_ _C_ _D_ Große-Veldmann, B.; Korotkova, N.; Reinken, B.; Lobin, W. & Barthlott, W. (2011). "_ Amborella trichopoda_ — Cultivation of the most ancestral angiosperm in botanic gardens". _The Journal of Botanic Garden Horticulture_. 9: 143–155. Retrieved 2016-10-21. * ^ _A_ _B_ Simpson, M.G. (2010). _ Plant Systematics_ (2nd ed.). Elsevier. p. 186 * ^ Carlquist, S. J. ">(PDF). _Pacific Science_. 55 (3): 305–312. doi :10.1353/psc.2001.0020 . Retrieved 2013-06-23. * ^ Sporne, K.R. (1974). _The Morphology of Angiosperms_. London: Hutchinson. ISBN 978-0-09-120611-6 . p. 98. * ^ _A_ _B_ Thien, L. B.; Sage, T. L.; Jaffré, T.; Bernhardt, P.; Pontieri, V.; Weston, P. H.; Malloch, D.; Azuma, H.; Graham, S. W.; McPherson, M. A.; Rai, H. S.; Sage, R. F. & Dupre, J.-L. (2003). "The Population Structure and Floral Biology of _ Amborella trichopoda_ (Amborellaceae)". _Annals of the Missouri Botanical Garden_. Missouri Botanical Garden Press. 90 (3): 466–490. JSTOR 3298537 . doi :10.2307/3298537 . * ^ _A_ _B_ Buzgo, M.; Soltis, P. S. & Soltis, D. E. (2004). "Floral Developmental Morphology of _ Amborella trichopoda_ (Amborellaceae)". _International Journal of Plant Sciences_. 165 (6): 925–947. doi :10.1086/424024 . * ^ _A_ _B_ _C_ _D_ Posluszny, U.; Tomlinson, P.B. (2003), "Aspects of inflorescence and floral development in the putative basal angiosperm _ Amborella trichopoda_ (Amborellaceae)", _Canadian Journal of Botany_, 81 (1): 28–39, doi :10.1139/b03-004 * ^ Endress, P. K. & Igersheim, Anton (2000). "The Reproductive Structures of the Basal Angiosperm _ Amborella trichopoda_ (Amborellaceae)". _International Journal of Plant Sciences_. Current Perspectives on Basal Angiosperms. The University of Chicago Press. 161 (S6): S237–S248. doi :10.1086/317571 . * ^ Floyd, S.K. & Friedman, W.E. (2001). "Developmental evolution of endosperm in basal angiosperms: evidence from _Amborella_ (Amborellaceae), _Nuphar_ (Nymphaceae), and _Illicium_ (Illiciaceae)". _ Plant Systematics and Evolution_. 228: 153–169. doi :10.1007/s006060170026 . * ^ _A_ _B_ The Angiosperm Phylogeny Group (2016), "An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV", _Botanical Journal of the Linnean Society_, 181 (1): 1–20, doi :10.1111/boj.12385 CS1 maint: Uses authors parameter (link ) * ^ _A_ _B_ Albert, V. A.; Soltis, D. E. ; Carlson, J. E.; Farmerie, W. G.; Wall, P. K.; Ilut, D. C.; Solow, T. M.; Mueller, L. A.; Landherr, L. L.; Hu, Y.; Buzgo, M.; Kim, S.; Yoo, M.-J.; Frohlich, M. W.; Perl-Treves, R.; Schlarbaum, S. E.; Zhang, X.; Tanksley, S. D.; Oppenheimer, D. G.; Soltis, P. S. ; Ma, H.; dePamphilis, C. W. & Leebens-Mack, H. (2005). "Floral gene resources from basal angiosperms for comparative genomics research". _BMC Plant Biology_. 5. * ^ Amborella Genome Database * ^ Cronquist, A. (1981). _An integrated system of classification of flowering plants_. New York: Columbia University Press. ISBN 9780231038805 . * ^ Cronquist, A. (1988). _The evolution and classification of flowering plants (2nd ed.)_. Bronx, NY: New York Botanical Garden. * ^ Thorne, Robert F. (1992). "Classification and geography of flowering plants". _Botanical Review_. 58 (3): 225–348. doi :10.1007/BF02858611 . * ^ Thorne, Robert F. (1992). "An updated phylogenetic classification of the flowering plants". _Aliso_. 13: 365–389. * ^ Dahlgren, R.M.T. (1980). "A revised system of classification of the angiosperms". _Botanical Journal of the Linnean Society_. 80 (2): 91–124. doi :10.1111/j.1095-8339.1980.tb01661.x . * ^ _A_ _B_ Soltis, P. S. Greilhuber, Johann; Doležel, Jaroslav & Wendel, Jonathan F. _ Plant Genome Diversity. Volume 2_. Springer. pp. 1–11. ISBN 978-3-7091-1160-4 . * ^ _A_ _B_ _C_ Pillon, Y. (2008). _Biodiversité, origine et évolution des Cunoniaceae : implications pour la conservation de la flore de Nouvelle-Calédonie_ (PDF) (PhD) (in French and English). University of New Caledonia. Retrieved 2013-06-22. * ^ Drew, B. T.; Ruhfel, B. R.; Smith, S. A.; Moore, M. J.; Briggs, B. G.; Gitzenda