Brexia

Brexia 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 leaf blades. The pentamerous flowers occur in cymes. The petals are greenish white, the samens are alternating with wide, incised staminodes. The superior ovary develops in a long ribbed fruit. Brexia naturally grows on the coast of East Africa, on Madagascar, the Comoros and Seychelles. Opinions differ about the number of species in Brexia. Sometimes the genus is regarded monotypic, B. madagascariensis being a species with a large variability, but other authors distinguish as many as twelve species.[3] Common names for B. madagascariensis include jobiapototra, tsimiranjana, tsivavena, vahilava, voalava, voankatanana, voantalanina, voatalanina and votalanina (all Malagasy), and mfukufuku (Swahili), mfurugudu (Shambala, Tanzania) and bwa kato (Seychelles).[2][4][5][6]

Brexia
Brexia madagascariensis, ↑flower, ↓leaves of a young plant
Scientific classification
Kingdom:	Plantae
Clade:	Tracheophytes
Clade:	Angiosperms
Clade:	Eudicots
Clade:	Rosids
Order:	Celastrales
Family:	Celastraceae
Genus:	Brexia Noronha ex Thouars
Species
Brexia madagascariensis (Lam.) Ker Gawl. type species Brexia alaticarpa G.E.Schatz & Lowry Brexia australis G.E.Schatz & Lowry Brexia marioniae G.E.Schatz & Lowry[1]
Synonyms
Thomassettia, Venana[2]

Description

Stems and leaves

Brexia is a shrub or small tree, usually 3–7, but occasionally up to 10 m high with many branches, that are smooth with ridges early on but later becoming cylindrical. The leaves are evergreen, with a leaf stem mostly 1–2 cm long, but sometimes very short. The leathery leaf blades are between 3½ and 50 cm long and 2 and 11 cm wide, with large differences in shape, narrowly inverted egg-shaped to linear with teeth or even spines along the edges in young growth, while on mature shoots they may be narrowly to broadly inverted egg-shaped and the edge toothy to entire, with a rounded to indented tip, and wedged along the leaf stem or rounded at the base. The stipules are very narrow.

Flowers

Each inflorescence may consist of usually between three and twelve flowers, but may occasionally contain as few as one and as many as seventeen flowers, usually on a flattened 1½−5 mm wide common stem of up to 9 cm long, and growing from a leaf axil, but in some forms on thick branches or the trunk (so-called cauliflory). Sometimes near the tip are one or two persistent leaf-like bracteoles of about 1 cm, but mostly the small scale-like bracteoles fall off quickly. Individual flowers are on stems of up to 2 mm long. Each flower has five sepals of about 2½ × 3½–4 mm, the lower half of which are fused, the free tips short triangular with a rounded tip and an entire edge. The five fleshy petals are greenish or creamy white, 1¼–1¾ × ⅞–1¼ cm, broadly inverted egg-shaped, with a blunt tip. The five stamens have filaments of about 1¼ mm thick, and are topped by anthers of 5 × 2½ mm. Alternating with the stamens are mostly unequally incised staminodes with three to five incisions. The superior ovary has five sides and is 8–10 mm high including the style.

Fruit and seed

The fully grown fruit is a drupe of 4–10 × 2–3 cm, ovoid to cylindric, with five prominent ribs, with the mesocarp initially woody but supposedly pulpy and edible when fully ripe. It contains brown or almost black, finely wrinkled, irregularly compressed, oval, keeled seeds of 4½–7½ × 3–3½ mm.[2][7]

Differences with other genera

Brexia has been assumed to be closely related to Escallonia. These two genera however differ in many details of the sporogenesis, gametogenesis and fertilisation, such as the ripe pollen which is two-celled in Escallonia and three-celled in Brexia.[8]

Taxonomy

Brexia is a deviant genus, that was assigned early on to the Brexiaceae or Brexioideae, together with two other enigmatic, monotypic genera, Ixerba and Roussea. The common characters between these taxa are few and are shared with many other Pentapetalae. These common characters include that they are all small trees or shrubs with simple leathery evergreen leaves, with an entire or serrated margin, and pentamerous flowers set individually or in cymes in the axil of the leaves. Botanist have differed on the rank and placement of these three genera. They have by some authors also been assigned to the Escalloniaceae, Saxifragaceae, Grossulariaceae, and the Hydrangeaceae. Genetic data indicate that the three genera are not each other's closest relatives, in fact are not closely related at all. Ixerba has now been included in the Strasburgeriaceae. Roussea is most related to Carpodetus, Cuttsia and Abrophyllum, that are nowadays combined in the family Rousseaceae, the sister group of the Campanulaceae, and together representing the basic branche of the Asterales. Brexia is closely related to the large celastracean genera Pleurostylia and Elaeodendron.[9]

Taxonomic history

Francisco Noronha described the genus Brexia before his death in 1788, although this name was only properly published by Thouars in 1806. Jean-Baptiste Lamarck described the same species as Venana madagascariensis in 1797. In 1823, John Bellenden Ker Gawler assigned this taxon to Brexia, creating the combination Brexia madagascariensis.[10] Later, Brexia (the nomen conservandum) was given priority over Venana (nomen rejectum). In 1902 William Hemsley described Thomassetia seychellarum based on a specimen from the Seychelles,[7] but this name was later synonymised with Brexia madagascariensis.

Modern classification

Brexia is together with Polycardia sister to a clade containing of Elaeodendron and Pleurostylia, and less related to the other Celastraceae. Opinions differ about the number of species in Brexia. Sometimes B. madagascariensis is regarded as a species with a large variability, but other authors distinguish as many as twelve species.[3] Populations outside Madagascar are sometimes regarded as separate taxa. Those on the coast of continental Africa with inflorescences that have few flowers and large fruits are referred to as var. mossambicensis, while those growing at higher elevations on the Seychelles characterised by small pinkish petals and small fruits are separated as B. madagascariensis ssp. microcarpa or alternatively B. microcarpa. B. madagascariensis is also spread along the East coast of Madagascar. All other putative taxa occur on Madagascar only, mostly with very limited distributions. These include:

• B. alaticarpa, with three to five flowers per cyme, with petiolate, narrow to lanceolate leaves with entire, rolled-down margins and a rounded tip, and conspicuously winged fruits (former Toamasina Province).
• B. australis, with one or two flowers per cyme, leaves mostly sessile and only up to 12 cm (coastal Anosy Region).
• B. marioniae, with inflorescences on the branches, and staminodes with entire margins or ending in two or three shallow teeth (former Antsiranana and Toamasina Provinces).[2]

Phylogeny

B. madagascariensis showing entire leaf margins of adult branches, distinctly flattened petioles and minute brachteoles

Modern genetic analysis results in the following relationship tree.[11]

	Polycardia         Brexia montana     Brexia australis         Brexia humbertii     Brexia madagascariensis               Brexiella/Evonymopsis     Elaeodendron/Pseudocatha         Halleriopsis       Erythrostegia       Astrocassine       Salvadoropsis       Pleurostylia       Hartogiopsis     Macrynodrys                       Lophopetalum     Kokoona

Etymology

Brexia is derived from the Greek word βρέχω (brecho) "to rain", and refers to the large leathery leaves that are impervious to rain.[12]

Distribution

B. madagascariensis is a widespread species found in coastal areas in Madagascar (Sava, Atsinanana, Analanjirofo, Vatovavy-Fitovinany, Atsimo-Atsinanana, and Anosy regions), Mozambique, Tanzania (including Zanzibar), and Comoro Islands.[13][14] The populations of Brexia that occur at higher elevations in the Seychelles are sometimes regarded as a distinct subspecies B. madagascariensis ssp. microcarpa, but others view it as a separate species B. microcarpa. All other putative species are endemics of Madagascar.[2]

Ecology

During the night, nectar is produced and stored in the cup of the flower. This is commonly associated with flowers specialising in pollination by bats. Common brown lemurs Eulemur fulvus were seen fouraging from Brexia flowers in the early morning. These primates took the inflorescences with their hands and bent these to lick the nectar from the flower cups. This was repeated in other nearby specimens of Brexia immediately after. Flowers were not damaged, and it is likely the lemurs assisted the pollination.[15] B. madagascariensis grows in coastal vegetations on a range of soil types such as coral, sand, loam or coarse rocky ground, in places like the edges of saline water swamp forest, mangrove swamps, evergreen shrubland and other forest types near the sea.[13] In Tanzania it has been found in an almost pure stand of Maesopsis eminii.[16] The fruit can float in sea water for many months and the numerous dark seeds within remain viable.[6]

Use

Fruits can be eaten raw as pulp when fully ripe. The wood can be used for spoons and tool handles, poles and yokes, but also provides fire wood and can be made into charcoal. A potion to treat stomach-ache can be produced by boiling the roots.[6]

References

1. "Brexia". The Plantlist. Retrieved 2016-03-20.
2. Schatz, George E.; Lowry, II, Porter P. (2004). "A synoptic revision of Brexia (Celastraceae) in Madagascar" (PDF). Adansonia. 26 (1): 67–81. Retrieved 2016-03-23.
3. "Brexia". Gateway to African Plants. Retrieved 2016-03-21.
4. "Plant Information for Brexia madagascariensis (Brexiaceae)". Africa tree database. Archived from the original on 2016-04-04. Retrieved 2016-03-21.
5. Government of the Seychelles, Third national report to the Convention on Biological Diversity (PDF), retrieved 2016-03-21
6. Ruffo, C.K.; Birnie, A.; Tengnas, B. (2002). Edible Wild Plants of Tanzania. Nairobi: Regional Land Management Unit. ISBN 978-9966-896-60-5. cited on "Brexia madagascariensis". Useful Tropical Plants. Retrieved 2016-03-21.
7. Robson, N.K.B. (1978). "Brexiaceae". Flora Zambesiaca. 4. cited on "Brexia madagascariensis (Lam.) Ker-Gawl". Kew eFloras. Retrieved 2016-03-25.
8. Mauro Cresti; Paolo Gori; Ettore Pacini, eds. (2012), "Sporo-, gametogenesis and fertilisation of Escallonia and Brexia with comments on their taxonomy", Proceedings of the Tenth International Symposium on the Sexual Reproduction in Higher Plants, 30 May – 4 June 1988, University of Siena, Siena, Italy, Springer Science & Business Media, ISBN 978-3642732713
9. Koontz, Jason A.; Soltis, Douglas E. (1999). "DNA sequence data reveal polyphyly of Brexioideae (Brexiaceae; Saxifragaceae sensu lato)". Plant Systematics and Evolution. 219 (3): 199–208. doi:10.1007/bf00985579. S2CID 758907.
10. "Venana madagascariensis". Tropicos. Retrieved 2016-08-08.
11. Bacon, Christine D.; Simmons, Mark P.; Archer, Robert H.; Zhao, Liang-Cheng; Andriantiana, Jacky (2016). "Biogeography of the Malagasy Celastraceae: Multiple independent origins followed by widespread dispersal of genera from Madagascar". Molecular Phylogenetics and Evolution. 94 (Pt A): 365–382. doi:10.1016/j.ympev.2015.09.013. PMID 26432393.
12. Quattrocchi, Umberto (2012). CRC World Dictionary of Medicinal and Poisonous Plants: Common Names, Scientific Names, Eponyms, Synonyms, and Etymology (5 Volume Set). CRC Press. ISBN 978-1420080445. Retrieved 2016-03-20.
13. "Brexia madagascariensis (Lam.) Ker Gawl". African Plant Database. Retrieved 2016-03-21.
14. "Brexia madagascariensis". Useful Tropical Plants. Retrieved 2016-03-21.
15. Birkinshaw, Chris (2002). "Probable pollination of Brexia madagascariensis (Lam.) Ker Gaul. by Eulemur fulvus at Ambila-Lemaitso, Madagascar". Lemur News. 7: 11.
16. "Brexia madagascariensis (Tree)". The Ecological Register. Retrieved 2016-03-21.