Taxon Details: Lecythis pisonis Cambess.
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Family:

Lecythidaceae (Magnoliophyta)
Scientific Name:

Lecythis pisonis Cambess.
Primary Citation:

Fl. Bras. Merid. 2: 377. 1829
Accepted Name:

This name is currently accepted.
Description:

Author: Scott A. Mori

Type: Brazil. Espírito Santo: Without locality, 1816-1821 (fl), St.-Hilaire 365 (lectotype, P, designated Mori & Prance, 1990)

Description: Large trees, to 50 m tall, with or less frequently without buttresses. Bark grayish to dark brown, with deep reticulately arranged more-or-less vertically oriented fissures, the outer bark laminated, the inner bark whitish, the sapwood white to yellowish-white, the heartwood reddish-brown. Leaves deciduous, the new leaves often reddish or cream-colored, flushed just before flowering; petioles 4-13 mm long; blades narrowly to widely ovate or elliptic to widely elliptic, glabrous, (6) 8-15 x 3-8 cm, the base obtuse to rounded, the margins crenate, the apex acuminate; secondary veins in 10-20 pairs. Inflorescences racemes, arising on stems below leaves; pedicels 5-12 mm long, glabrous or puberulous. Flowers 3-7 cm diam.; calyx with 6, widely ovate purplish lobes, 3.5-8 x 3.5-8 mm; petals 6, subequal, narrowly to widely obovate, 17-36 x 14-27 mm, usually purple or white tinged with purple, fading to white after falling; androecial hood flat, uni-dimensional as seen in longitudinal section, usually purple, sometimes white when old, the proximal staminodes often bearing vestigial anthers with fodder pollen, the distal ones often without anthers, the appendage-free ligule present; ovary 4-locular, with 6-15 ovules in each locule, the style 1-2 mm long, with annular ring toward apex. Fruits globose, oblongoid, or turbinate, (6)10-15 x (8.5)10-20(30), the calycine zone prominent or not. Seeds 10-30 per fruit, fusiform, sulcate, 4-6 x 2.5-3 cm, the basal, cord-like funicle surrounded by fleshy, white aril. X = 17.

Common names: Common names. Brazil: sapucaia (tree), castanha de sapucaia (seeds). English. Monkey pot (tree or fruit), paradise nut (seed).

Distribution: Native to the Brazilian coastal forest from Pernambuco to São Paulo and in Amazonia, especially in eastern Amazonia along the Amazon River. It has been commonly collected in eastern Amazonia where it is often planted near homes for its edible seeds. It is cultivated as an ornamental in São Paulo and Rio de Janeiro and in tropical botanical gardens throughout the world, including gardens in Cuba. Jamaica, Panama, and Trinidad.

Ecology: A canopy to emergent tree found in both periodically flooded and non-flooded forest habitats. This species is considered to be a late successional species by restoration ecologists (Pers. comm. R. A. Sartori to S. A. Mori, Oct 2013.

Phenology: In a paper by Mori et al. (1980), data on the fall and flush of leaves, flowering, and fruiting of four individuals of Lecythis pisonis (sapucaia) in the moist forests of southern, coastal Bahia were collected, and in 1978 and 1979 total flower production of five individuals was recorded. Fall and flushing of leaves occured at the end of winter to the end of spring (September-November) during the course of the study. In this season, the sapucaia loses all of its leaves, remains leafless for 10 to15 days, and then produces leaves and flowers at the same time. The fruits mature seven months after fertilization in the months of March and April. Flowering took place independent of rain fall at the same time each year during the course of the study which suggests that this species flowers in response to lengthening days at the end of the winter into spring. There are 3,699 pollen grains produced for each ovule. The number of fruits produced did not pass 0.02% of the number of flowers for any of the trees.

Pollination: Pollination. The flowers are known to be pollinated by female carpenter bees (Xylocopa fontalis) in Bahia and in Amazonia (Mori & Prance, 1990). The bees land on the androecial hood and take fodder pollen (i.e., non-germinating pollen) from the androecial hood. As they collect fodder pollen, fertile pollen is placed on their heads and backs and this is presumably rubbed off on the stigmas of subsequent flowers visited.

Dispersal: The seeds are dispersed by the bat Phyllostomus hastatus. The bats are attracted to the fruits by the fleshy aril surrounding the funicle (Greenhall, 1965). The bats drop the seeds under their night roosts after eating the aril or accidentally while in flight from the trees to their roosts. Although this observation was made outside of the native range of the species in Trinidad, our conversations with local Amazonian people indicate that dispersal of this species by bats occurs there as well. Other frugivorous bats may also be involved in seed dispersal of L. pisonis but this has not been demonstrated.

Predation: Nelson Wisnick has witnessed what he has identified as a crimson-crested woodpecer (Campephilus melanocucos) pecking a large hole in the fruit of this species (http://www.youtube.com/watch?v=WXYJb5oPd9s). The woodpecker is either seeking larvae that are preying on the fruit or for seeds or other tissue to eat. In either case, the end result is the same as preying upon the seeds. Monkys have been reported to eat the seeds.

Field characters: This species is easily recognized by its tendency to bruise bluish-green (bark, flowers, fruits) when damaged; tall stature; fissured bark; laminated outer bark; flat, uni-dimensial androecial hood, often with fodder pollen in the proximal-most staminodes; petals and androecial hood with at least some tinges of purple (both of which fade to white after anthesis); large fruits reaching the size of a human head; and fusiform, sulcate seeds with a long basal aril.

Taxonomic notes: Lecythis pisonis belongs to a group of species known as the sapucaia group (Mori & Prance, 1981). These species are all 1) large trees with fissured, laminated bark, 2) leaves , fruits, and flowers that bruise bluish-green, 3) flat androecial hoods, 4) with fodder pollen in the staminodial anthers of the proximal staminodes, 5) large fruits, often the size of a human head, and 6) sulcate seeds subtended by a yellowish-white, large aril. In a molecular study by Mori et al. (2007) and Huang et al. (2015), this group, consisting of L. ampla, L. lanceolata, L. pisonis, and L. zabucajo, form a distinct clade. This group has been recognized taxonomically as Lecythis sect. Pisonis (Huang et al., 2015). The taxonomy of this group is currently undergoing revision and all indications are that it will be recognized as a separate genus and that there will be modifications in the current species circumscriptions as well as the description of several new species.. As in all other species of Lecythis sect.pisonis, the fruits of L. pisonis have considerable variation in size, shape, and the degree of conspicuousness of the calycine ring. The largest fruits almost always have the most conspicuous calycine rings (see images below). Moreover, it seems like the largest fruited individuals have a preference for periodically flooded habitats in Amazonia. The earliest, validly published name for the larger fruited individuals with conspicuous calycine rings is Lecythis paraensis Huber ex Ducke (see drawing in the protologue of this species), and the largest known fruits are found in a population described by Ledoux as Pachylecythis egleri. The fruits of L. pisonis from eastern extra-Amazonian Brazil where the type was collected do not have expanded calycine rings. Pubescence of the petiole and hypanthium also shows considerable variability. Some poplulations are essentially glabrous (e.g., in populations from eastern extra-Amazonian Brazil) whereas others are pubescent. Finally flower color is not always purple. For example, I have collected flowers from an individual in central Amazonia with white petals and yellow androecial hoods (see image of flowers from Mori et al. 27266 attached to this number in the specimen catalog). In summary, further collections and study may show that some of these populations merit recognition as subspecies or even species.

Conservation: IUCN Red List: not on list (IUCN, 2009): Plantas Raras do Brasil. not on list (Giulietti et al., 2009).

Uses: The seeds are edible and in the past were imported into the United States under the name of Paradise nuts. They are, however, difficult to harvest because bats carry away the seeds soon after the fruits open. Lecythis pisonis is planted near homesteds in eastern Amazonia for its edible seeds and as an ornamental and botanical curiosity, especially in botanical gardens, throughout the tropics. Lecythis pisonis is raised in nurseries and used for reforestation in the state of Rio de Janeiro, Brazil (Pers. comm. R. A. Sartori, Oct. 2013).

Etymology: Etymology. This species is named after William Piso, a German physician interested in medicinal plants who lived in Pernambuco during the time of the Dutch occupation (1630-1654) of this part of Brazil. He and George Marcgrave, the latter posthumously, published Historia naturalis Brasiliae, studied and illustrated what may be this species but, pending further study, may represent another described species currently placed in synonymy withthis species. The second part of the Historia naturalis Brasiliae entitled Historia rerum naturalium Brasiliae was authored by Marcgrave and has been considered by some as the most outstanding natural history documents published in the 17th century in Brazil (MacBryde, 1970). See bibliography entires for Marcgrave and Piso for further information about these early explorers..

Source: Based on Mori in Mori and Prance (1990).

Acknowledgements: We are grateful to B. Angell, C. Gracie, M. Nee, C. Potascheff, and M. Rothman for allowing us to use their images to illustrate the characters of this species.

Flora and Monograph Treatment(s):

Lecythis pisonis Cambess.: [Article] Prance, Ghillean T. & Mori, S. A. 1979. Lecythidaceae - Part I. The actinomorphic-flowered New World Lecythidaceae (Asteranthos, Gustavia, Grias, Allantoma & Cariniana). Fl. Neotrop. Monogr. 21: 1-270.
Lecythis pisonis Cambess.: [Article] Mori, S. A. & Lepsch da Cunha, Nadia M. 1995. The Lecythidaceae of a central Amazonian moist forest. Mem. New York Bot. Gard. 75: 1-55.
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