Pentaclethra macroloba

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Pentaclethra macroloba
Pentaclethra macroloba tree in Costa Rica

Least Concern  (IUCN 3.1)[1]
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Clade: Rosids
Order: Fabales
Family: Fabaceae
Subfamily: Caesalpinioideae
Clade: Mimosoid clade
Genus: Pentaclethra
Species:
P. macroloba
Binomial name
Pentaclethra macroloba
Synonyms[3][4]

Acacia macroloba Willd.
Mimosa macroloba (Willd.) Poir.
Acacia aspidioides G.Mey.
Pentaclethra filamentosa Benth.
Pentaclethra brevipila Benth.
Cailliea macrostachya Steud.
Dichrostachys macrostachya (Steud.)

Walp.

Entada werbaeana J.Presl[2]

Pentaclethra macroloba is a large and common

fish poison. Despite their toxicity, the seeds are eaten by variegated squirrels, parrots and macaws, and serve as the nurseries of the larvae of the moth Carmenta surinamensis
.

Etymology and local names

The name Pentaclethra is derived from

Ancient Greek, penta meaning 'five', and cleithro meaning 'bolt', which alludes to the five imbricate sepals and five petals joined at their bases; this defines the species of the genus.[5]

The tree is

Sranantongo it is called kroebara.[8] In Spanish it is called gavilán or quebracho.[9]

Description

Pentaclethra macroloba is a tree which can vary in height and size depending on distribution. In Costa Rica it grows to the greatest dimensions and is usually a canopy tree which reaches a trunk diameter of 130 centimetres (51 in),[3] and heights of 30–40 metres (98–131 ft).[3][9] In Brazil the average height is said to be approximately 14 metres (46 ft),[6] and in the eastern Guianas it is said to grow up to 25 metres (82 ft).[8] In these areas it is a subcanopy tree.[8][10] This may be due to infraspecific variation.[3][10]

The canopy or crown is dense and broad.[6][8]

The twigs are brownish-red, when they are young they are covered in a

puberulous indumentum.[8] The twigs are without spines.[9][11] The 0.7–0.8 cm stipules are deciduous, linear or hair-like.[9]

Leaves

Detail of a leaf

The large leaves are

nectar glands on the rachis, or anywhere on the leaves.[9][13]

Flowers

Illustration of a flower, note the long staminodes
An inflorescence displaying the staminodes.
The flower spikes are held above the foliage

The inflorescence is a 10 to 25 cm, most often 20 cm long spike,[5][8] with approximately 320 flowers, each one perfect and complete (containing all the functional male and female parts).[5] The rachis is 15–20 cm long and has 1–5 cm peduncle.[9] The spikes are terminal or subterminal on the branches.[8][9] Developing spikes are covered in a reddish-brown pubescence.[8]

The about 2 cm

nectary at the base of the ovary.[5]

The apical gland mentioned above is a rather unique and mysterious anatomical feature found only in the anthers of mimosids and related legumes. Such glands occur in four distinct forms, and the

food body or an osmophore.[14]

Fruit, seeds and seedlings

Despite producing hundreds of flowers per inflorescence, each spike only produces a few fruit, which is a pod (or bean).[3] These are held erect above the foliage and branches.[8][9] The pod is (16.1-) 30 to 38 (-45.2)cm in length, (3.3-) 4.1 to 6 (-7.3)cm in width,[6][8] and 0.8 to 1.1 cm thick.[8] It is coloured greenish when fresh, and dark brownish-grey when dry.[9] The shape is falcate or linear-spatulate and flattened, with the greatest width above the middle, the base being attenuated truncate and the apex being rounded.[8][9] The pod is woody, and it opens by elastic dehiscence from the apex to the base.[9] It has longitudinal veins. The dorsal suture is up to 1.1 cm broad and thickened, the ventral suture is up to 0.8 cm thick. As the seedpod dried tension builds until the two valves twist open suddenly when the suture cracks open.[8] The pod has three to eight seeds,[3] more often four to six.[8][9]

The large and flat seeds[8] measure from 3.8 to 5 cm, exceptionally 6.1 cm, in length, and 2.5 to 3.5 cm, exceptionally 5.7 cm in width. The form is lens-like to spoon-shaped or deltoid, or ovoid to elliptical, and laterally flattened.[6][9] They are coloured dark brown. The seeds do not have arils.[9] The cotyledons are composed anatomically of thin-walled parenchyma cells with various oil cells.[6]

When the seeds

stomata on the bottom surface.[6]

Similar species

The plant can be recognised as belonging to this species by: not having any spines, bipinnate leaves without nectar glands, inflorescences as large spikes covered in conspicuous white stingy staminodes, and fruit held above the branches, in the shape of a machete, which open with an elastically explosive mechanism from the apex to the base. Its large seeds are recognisable by their more or less lens-like to spoon-shaped form.

morphology.[6]

Distribution

According to one 1983 source, it exists in three disjunct populations, one through the lowlands of northeast

Darien Province in eastern Panama. The third is found in western Panama, Costa Rica and Nicaragua.[3][10]

Another source considers it introduced to Trinidad and Tobago and the Windward Islands.[4]

In Costa Rica and Nicaragua it primarily grows along the eastern coasts.[9][15] It does not occur in western Nicaragua.[15] The populations in Costa Rica are not disjunct, as it occurs through the central valleys and in certain patches on the western coast.[9] In eastern Panama it is the dominant tree in certain moist, seasonal, swamp forests in coastal areas of the Atlantic.[16] It has been collected in Bocas del Toro and Darién Province.[11]

In Colombia it is only found along the Pacific coast.[17] It is a common tree in northern parts of Brazil. It is found in the states of Acre, Amazonas, Amapá, Pará and Roraima, and perhaps in the state of Mato Grosso.[12]

In the 1993 article Catalogue of the Flowering Plants and Gymnosperms of Peru it was said to occur in the

determined to be P. macroloba in 1993, was re-identified as the species Dimorphandra coccinea in 1997.[20]

Ecology

It grows in lowland forest from sea level to 600 metres above sea level in Costa Rica, and is especially abundant in humid, temporarily inundated rainforests.[3][9] It is restricted to the lowlands in Panama.[11] It occurs at an altitude of 0-290m in Colombia.[17] In Brazil it is known from the seasonally flooded, closed tropical rainforests known as várzea, in both igapó (inundated forest) habitat and forest on 'terra firme' land.[12][21] In the Brazilian Amazon River estuary it is the 'hyperdominant' lifeform, forming vast monospecific stands.[21][22] In the Amazon River estuary the plants are subjected to daily tides. In the wooded floodplains of Mazagão, Brazil, the population density was 18 trees per hectare with a basal area of 49 m2 per hectare.[23] In La Selva Biological Station Pentaclethra macroloba constitutes 40% of the basal area and up to 18% of the stem density in parts of the forest, where it forms monocultural stands.[13]

Seed pods and seeds

Different trees synchronize their flowering in the dry season.[21] In Costa Rica flowering starts in March.[9] It is the conspicuous white staminodes which attract pollinators, not the minute petals.[5] It is cross-pollinated.[3] The main visitors to the flowers are wasps, bees and ants. Immature fruits appear during the dry season and ripe fruits in the rainy season. The species is evergreen, showing no specific changes in the leaves throughout the year, despite flooding.[21]

The tree uses at least three methods of seed dispersal. Seeds are mechanically dispersed when mature pods split, popping open during the rainfall.[21][24][25] The mechanism works by elastic dehiscence. The seeds are expelled up to ten meters from the parent tree.[26] This is the season when rivers overflow their banks, and the lowland floods.[21] In 2001 the discovery was published that submerged seeds develop air pockets that allow them to float; this was then interpreted as evidence that the species has adaptions for dispersal by water (hydrochory).[24][25] Dispersal of seeds during daily river flooding and at the peak of the river flood allows the plant to use the receding waters of the tides to transport them long distances.[21] The population density increases with the distance from the normal edge of the inundation. This is believed to be due to the action of the floodwaters during high tide, the floating seeds end up stuck behind fallen branches, buttress roots or other objects, often in great numbers.[22][23]

Like many leguminous plants, P. macroloba is a nitrogen fixer which forms a symbiotic relationship with Rhizobium, which grows in specialised root nodules. While root nodules are typically found on buried roots, P. macroloba individuals growing in swampy areas produce nodules on aerial roots.[27]

Seed predation

Gary Hartshorn, a

Heteromys desmarestianus. An experiment where the mice were exclusively fed the seeds killed them all. Hartshorn noted the presence of the larvae of a wasp-like, clear-winged moth feeding in the seeds, but noted that their presence did not impede germination, and speculated that the opening of the cotyledons at the surface of the soil was perhaps a defensive tactic of the plant, as it exposes the caterpillars to foraging ground insectivores.[10] The seeds are actually heavily predated upon, frequently ten or eleven insect larvae emerge through small holes on the exposed surface of the opened cotyledons.[3] This appears to be primarily caused by the moth Carmenta surinamensis, which was found in 43.6% of the seeds in one study in Costa Rica. This reduces their viability, but does not always kill them.[13]

Hartshorn himself had noted in his earlier thesis paper that grey squirrels (

Pionus senilis) feed upon the seeds up in the trees while still in the unopened pods. The animals access the seeds by making a hole in one valve of these pods. A small study which looked at 100 pods found that 22% of the fruit had such holes in them. The animals rarely get all the seeds, but often just take one or few from a pod. Only 5% of the total seed production is consumed before the pods open. These are taken in a rather random manner, with no particular selection apparent. An explanation for this may be that the preferences of the squirrels and parrots are precisely diametrically opposed and they might cancel each other out, but it is also possible that the optimal foraging theory, which would have it that predators select the biggest and easiest to forage seeds, may not apply here, as both animals may be able to tolerate some toxins, they are still susceptible to them, and must limit their intake. The pod damage may hinder the elastic mechanism that flings the seeds away from the tree, thus 22% the pod predation may still be quite significant for the population dynamics.[26] Great green macaws have also been recorded to feed on the seeds.[28]

Capuchin monkeys have a strange and mysterious behaviour, known as 'anointing', in which they periodically rub their bodies with specific objects. On Trinidad the small population of white-fronted capuchins at Bush Bush Wildlife Sanctuary use the pods of P. macroloba to anoint themselves. They pluck a pod from the tree, break it open, and rub themselves with the inner part. These Trinidadian monkeys do this often, and always do it in sight of each other. Afterwards they often eat one or more of the seeds found inside the pod, whether these are ripe or not. Other monkeys avoid the bark and seeds of P. macroloba. Perhaps the monkeys use the seedpods as a curative, or some kind of medicine.[29]

Phytosociology

In Mazagão, Brazil, it is found growing together with a high frequency of palms and the trees

Pterocarpus amazonicus, Symphonia globulifera, Licaria mahuba, Hernandia guianensis and others.[23]

It is a common tree in the

Mora excelsa forests of the Guianas and the Orinoco Delta of Venezuela, in this habitat the Mora is the dominant species, and Pentaclethra is a subcanopy tree.[8]

In Central America the species is the dominant canopy tree, growing with

Iriartea gigantea, and the trees Pterocarpus officinalis and Stryphnodendron microstachyum.[3]

Uses

Timber

The

specific gravity of 0.51–0.61.[3] The timber has commercial value.[30] It is used to make furniture, cabinets and for general construction.[3] The timber is traded as 'gavilán'.[31]

Other

Pracaxi oil

Pracaxi oil, which is extracted from the seeds of P. macroloba, is rich in

fish poison.[8]

Traditional medicinal properties ascribed to this plant species by caboclos (Brazilian Portuguese for 'locals') and various native groups are as an antivenom against snakebites,

triterpenoid saponins which they named macrolobin-A and B. Although these compounds do not fully counteract all the venomous effects, they could find use as complements to standard treatment against snakebites, or as molecular models for the development of novel future medicinal compounds.[35]

Conservation

It is a common species where it occurs in the Guianas[8] and Central America.[9][15]

References

  1. doi:10.2305/IUCN.UK.2019-2.RLTS.T62027193A149061882.en
    . Retrieved 25 December 2022.
  2. doi:10.5962/bhl.title.61845
    .
  3. ^ a b c d e f g h i j k l m n o p q Flores, E. M. (2002). "Pentaclethra macroloba" (PDF). In J. A. Vozzo (ed.). Tropical Tree Seed Manual. Agricultural Handbook. Vol. 721. Washington, DC: USDA Forest Service. pp. 601–604.
  4. ^ a b "Pentaclethra macroloba (Willd.) Kuntze". Plants of the World Online. Royal Botanic Gardens, Kew. 2017. Retrieved 23 May 2021.
  5. ^
    doi:10.3732/ajb.1600308
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  6. ^
    doi:10.1590/2317-1545v41n4222721
    . Retrieved 23 May 2021.
  7. ^
    doi:10.1016/j.jep.2005.01.063
    . Retrieved 29 May 2021.
  8. ^
    ISBN 90-9000987-6
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  9. ^ a b c d e f g h i j k l m n o p q r s t u v w Zamora, N. (1 April 2016). "Pentaclethra macroloba (Willd.) Kuntze". Manual de Plantas de Costa Rica - Draft Treatments (in Spanish). Missouri Botanical Garden. Retrieved 17 May 2021.
  10. ^ a b c d Hartshorn, Gary (January 1983). "Species Accounts: Pentaclethra macroloba (Gavilán)". In Janzen, D. H. (ed.). Costa Rican Natural History. University of Chicago Press. pp. 301–303.
  11. ^
    doi:10.2307/2394414
    . Retrieved 25 May 2021.
  12. ^ a b c Morim, M.P. (2020). "Pentaclethra macroloba". Flora do Brasil 2020. Jardim Botânico do Rio de Janeiro. Retrieved 21 May 2021.
  13. ^ a b c McKenna, Duane D.; McKenna, Katherine M. (July 2006). "Sesiid Moths Reduce Germination, Seedling Growth, and Survivorship in Pentaclethra macroloba (Mimosoideae), a Locally Dominant Lowland Neotropical Tree". Biotropica. 38 (4): 508–513. Retrieved 21 May 2021.
  14. doi:10.2307/2446002
    .
  15. ^ a b c Zarucchi, James L. (3 June 2009). "Pentaclethra macroloba (Willd.) Kuntze". Flora de Nicaragua (in Spanish). Missouri Botanical Garden. Retrieved 17 May 2021.
  16. ^ C. Michael Hogan. 2008. Isthmian–Atlantic moist forests. Encyclopedia of Earth and World Wildlife Fund National Council of Science and the Environment, eds. Mark Mcginley and Cutler Cleveland
  17. ^ a b Bernal, R.; Gradstein, S. R.; Celis, M., eds. (2015). Catálogo de plantas y líquenes de Colombia (in Spanish). Bogotá: Instituto de Ciencias Naturales, Universidad Nacional de Colombia.
  18. ^ "Pentaclethra macroloba (Willd.) Kuntze". Peru Checklist. Missouri Botanical Garden. 8 August 2009. Retrieved 23 May 2021.
  19. ^ Brako, L.; Zarucchi, J. L. (1993). "Catalogue of the Flowering Plants and Gymnosperms of Peru". Monographs in Systematic Botany from the Missouri Botanical Garden. 45 (i–xl): 1–1286.
  20. ^ "Alwyn H. Gentry - 56239". Tropicos.org. Missouri Botanical Garden. 2021. Retrieved 25 May 2021.
  21. ^
    doi:10.1007/s00468-021-02095-x
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  22. ^
    doi:10.15517/rbt.v69i1.43446
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  23. ^
    doi:10.1590/1806-90882017000400006
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  24. ^
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    doi:10.1046/j.0269-8463.2001.00586.x
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  26. ^ a b Anhalzer, Gabriela; Fournier, Michelle; O'Connor, Tim; Stevenson, Louise; Yglesias, Mariel (2010). "Traits for predator selection on Pentaclethra macroloba seeds" (PDF). American Journal of Undergraduate Research. 8 (4): 1–8. Retrieved 26 May 2021.
  27. S2CID 10885930
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  28. ^ Humedales de Ramsar (FIR) – Anexo #2 Biodiversidad 2009 (PDF) (Report) (in Spanish). Centro Científico Tropical. 2009. p. 12. Retrieved 31 August 2019.
  29. ^ a b c Lynch Alfaro, Jessica W.; Matthews, Luke; Boyette, Adam H.; MacFarlan, Shane J.; Phillips, Kimberley A.; Falótico, Tiago; Ottoni, Eduardo; Verderane, Michele; Izar, Patrícia; Schulte, Meredith; Melin, Amanda; Fedigan, Linda; Janson, Charles; Alfaro, Michael E. (2012). "Anointing Variation Across Wild Capuchin Populations: A Review of Material Preferences, Bout Frequency and Anointing Sociality in Cebus and Sapajus". American Journal of Primatology. 74: 299–314. Retrieved 26 May 2021.
  30. ^ "ontrato de Realización de los Diseños del Proyecto Específico de Senderización y Aprovechamiento Turístico Sostenible con Participación Comunitaria en el Refugio Nacional de Vida Silvestre Gandoca – Manzanillo" (PDF) (in Spanish). INBio. May 2012.
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  35. ^
    doi:10.1016/j.toxicon.2007.03.024
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