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''Solanum lycocarpum'', or wolf apple, is common in the Brazilian savanna, the Cerrado ecoregion. The plant is called ''lobeira'' ("wolf's plant") or ''fruta-do-lobo'' ("wolf's fruit") in Portuguese. The name "wolf apple" comes from the fact that they account for more than 50% of the maned wolf's diet. Likewise, the scientific name "''lycocarpum''" is formed from Latinized Greek elements "''lyco-''", meaning "wolf", and "''carpum''" meaning "fruit".


Range

The native range of the wolf apple tree is the Brazilian savannah, but it grows also on pastures and disturbed land, such as highway margins, in various parts of Brazil. It prefers moist, clay soil, full sun, and mild temperatures.


Description

The wolf apple plant is a flowering
shrub A shrub (often also called a bush) is a small-to-medium-sized perennial woody plant. Unlike herbaceous plants, shrubs have persistent woody stems above the ground. Shrubs can be either deciduous or evergreen. They are distinguished from trees ...
or small tree with round open crown, ranging in height from . The large leaves are long, simple but deeply lobed, tough, and covered in a soft grey-white fuzz, in alternate disposition. Plants flower through the year, but more intensely during the dry season, from late fall to early spring. The flowers are similar to those of the
bitter nightshade ''Solanum dulcamara'' is a species of vine in the genus '' Solanum'' (which also includes the potato and the tomato) of the family Solanaceae. Common names include bittersweet, bittersweet nightshade, bitter nightshade, blue bindweed, Amara ...
: star-shaped, with five sepals, five blue or purple
petal Petals are modified Leaf, leaves that surround the reproductive parts of flowers. They are often advertising coloration, brightly colored or unusually shaped to attract pollinators. All of the petals of a flower are collectively known as the ''c ...
s fused at the base, and five large yellow
anther The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filam ...
s that release
pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametophyt ...
through an opening at the tip. The flowers are arranged in helical monochasal cyme, opening from the base up. Flowers lack
nectar Nectar is a sugar-rich liquid produced by plants in glands called nectaries or nectarines, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualists ...
, and are either male or hermaphroditic. The latter produce fruit after cross-pollination by several species of pollen-collecting
bee Bees are winged insects closely related to wasps and ants, known for their roles in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey. Bees are a monophyly, monophyletic lineage within the ...
s, including ''
Xylocopa virescens Carpenter bees are species in the genus ''Xylocopa'' of the subfamily Xylocopinae. The genus includes some 500 bees in 31 subgenera. The common name "carpenter bee" derives from their nesting behavior; nearly all species burrow into hard plant m ...
'', '' X. frontalis'', '' Oxaea flavescens'', '' Centris scopipes'', '' C. fuscata'', '' Bombus morio'', '' Exomalopsis'' sp., '' Pseudaugochloropsis graminea'', ''
Apis mellifera The western honey bee or European honey bee (''Apis mellifera'') is the most common of the 7–12 species of honey bees worldwide. The genus name ''Apis'' is Latin for "bee", and ''mellifera'' is the Latin for "honey-bearing" or "honey carrying", ...
'', and '' Megachile benigna''. Fruit production occurs mostly between December and January. The fruits are large, up to in diameter and weighing or more, and contain up to 500 dark brown comma-shaped seeds, about . The rind is thin and lustrous, and remains green even after ripening. The pulp is yellowish, soft, sweetish and extremely aromatic. They resemble the tomato in external shape and internal morphology, but also the
eggplant Eggplant ( US, Canada), aubergine ( UK, Ireland) or brinjal (Indian subcontinent, Singapore, Malaysia, South Africa) is a plant species in the nightshade family Solanaceae. ''Solanum melongena'' is grown worldwide for its edible fruit. Mos ...
for the texture and color of the flesh. The maned wolf is an important seed-spreading agent.


Uses


Food

The ripe fruits are edible to humans, and are consumed by local populations in jams and preserves, but the unripe fruits are quite rich in tannin. The fruits are used as a fallback source of food for cattle in the dry season. At the onset of maturation, the whole fruit contains 85% moisture (by weight), 5% starch, and 10% other residue. The dry pulp, without peel and seeds, yields over 50% starch, more than cassava. The starch is locally separated by blending the fruit with water and straining. The pulp of the ripe fruit contains about 75% moisture, 11% carbohydrate, 1% fat, 1% protein, 1% ash, 2.5% soluble fiber, and 2% insoluble fiber. The main volatiles in the ripe fruit are hexanal (which has the scent of grass) and
ethyl butanoate Ethyl butyrate, also known as ethyl butanoate, or butyric ether, is an ester with the chemical formula CH3CH2CH2COOCH2CH3. It is soluble in propylene glycol, paraffin oil, and kerosene. It has a fruity odor, similar to pineapple, and is a key in ...
(scent of apple), although dozens of other compounds contribute to the fruit's aroma. All other parts of the plant are
poison Poison is a chemical substance that has a detrimental effect to life. The term is used in a wide range of scientific fields and industries, where it is often specifically defined. It may also be applied colloquially or figuratively, with a broa ...
ous to humans. However, the leaves are eaten by cattle, and are also an important food source for the bat ''
Platyrrhinus lineatus The white-lined broad-nosed bat (''Platyrrhinus lineatus'') is a bat species found in southern and eastern Brazil, Paraguay, Uruguay, northern Argentina, Bolivia, Peru, Ecuador, Colombia, French Guiana and Suriname. References White ...
''.


Medicine

Currently, there are no proven medicinal uses for the wolf apple, but the plant is used in the local folk medicine for obesity, cholesterol reduction, and as a treatment for diabetes. About 30% of the starch extracted from almost-ripe fruits, which commercialized as capsules, is digestion-resistant, which seems to be the rationale for these popular uses. High-fiber ''Solanum lycocarpum'' flour was found to reduce diabetes symptoms in diabetic rats; however, the starch did not show such effect. The fruit contains about 1.5% of glycoalkaloids in its dried fruits, chiefly solamargine and solasonine, mostly in the peel and seeds. Extracts have been shown to kill in vitro the human parasites ''
Schistosoma mansoni A paired couple of ''Schistosoma mansoni''. ''Schistosoma mansoni'' is a water-borne parasite of humans, and belongs to the group of blood flukes (''Schistosoma''). The adult lives in the blood vessels ( mesenteric veins) near the human inte ...
'' and '' Giardia lamblia'', '' Leishmania amazonensis'', and '' Leishmania infantum''. The main
phenolic compounds In organic chemistry, phenols, sometimes called phenolics, are a class of chemical compounds consisting of one or more hydroxyl groups (— O H) bonded directly to an aromatic hydrocarbon group. The simplest is phenol, . Phenolic compounds are c ...
in ethanol extracts of the ripe fruits are caffeic acid and chlorogenic acid. It is believed that the fruits protect the maned wolf from infection of the giant kidney worm, which is usually fatal for the animal. ''Solanum lycocarpum'' fruit extracts have also been shown to kill larvae of the ''Culex'' mosquito.


Pests

The wolf apple tree is the natural host of the larva of the butterfly '' Leucanella memusae''. Contact with the urticating bristles of this caterpillar may result in severe dermatitis and hemorrhage. The weevil '' Collabismus clitellae'' lays eggs into stems of the plant, causing the formation of
gall Galls (from the Latin , 'oak-apple') or ''cecidia'' (from the Greek , anything gushing out) are a kind of swelling growth on the external tissues of plants, fungi, or animals. Plant galls are abnormal outgrowths of plant tissues, similar to be ...
s, with measurable harmful effect on the plant's growth and fruit production.


References

Fernando Tatagiba (2006):
Lobeira, fruta-de-lobo
. Online article on the ''Biólogo'' website, published 2006-10-06. Accessed on 2020-06-30.
J. C. Motta-Junior, S. A.Talamoni, J. A. Lombardi, and K. Simokomaki (1996): "Diet of the maned wolf, ''Chrysocyon brachyurus'', in central Brazil". ''Journal of Zoology'', volume 240, issue 2, pp. 277–84. M. Motidome, M. E. Leekning, and O. R. Gottlieb (1970): "A química de Solanáceas brasileiras. 1 - A presenca de solamargina e de solasonina no juá e na lobeira." ''Anais da Academia Brasileira de Ciências'', volume 42, pp. 375–76 J. A. Lombardi and J. C. Motta Jr. (1993): "Seed dispersal of ''Solanum lycocarpum'' St. Hil. (Solanaceae) by the maned wolf, ''Chrysocyon brachyurus'' Illiger (Mammalia, Canidae)." ''Ciência e Cultura'', volume 45, pp. 126–27. A.T. Oliveira-Filho and L. C. A. Oliveira (1988):
Biologia floral de uma população de ''Solanum lycocarpum'' St. Hil. (Solanaceae) em Lavras
. ''Revista Brasileira de Botânica, volume 11, pp. 23–32. Abstract accessed on 2020-06-30
Fabiane Farina (2010): "Glycemic and urinary volume responses in diabetic mellitus rats treated with ''Solanum Lycocarpum''." ''Applied Physiology, Nutrition, and Metabolism'', volume 35, issue 1, pp. 40–44. Bicalho Silva and Viviane de Cassia (2015): "Larvicidal activity of oils, fatty acids, and methyl esters from ripe and unripe fruit of ''Solanum lycocarpum'' (Solanaceae) against the vector ''Culex quinquefasciatus'' (Diptera: Culicidae)". Mariza A. Miranda, Lizandra G. Magalhaes, Renata F. J. Tiossi, Christian C. Kuehn, Luiz G. R. Oliveira, Vanderlei Rodrigues, James D. McChesney and Jairo K. Bastos (2012): "Evaluation of the schistosomicidal activity of the steroidal alkaloids from ''Solanum Lycocarpum'' fruits". ''Parasitology Research'', volume 111, pp. 257–62. M. T. P. S. Clerici, C. Kallmann, F. O. G. Gaspi, M. A. Morgano, F. Martinez-Bustos, and Y. K. Chang (2011): "Physical, chemical and technological characteristics of ''Solanum lycocarpum'' A. St. HILL (Solanaceae) fruit flour and starch". ''Food Research International'', volume 44, issue 7, pp. 2143–50. Rodrigo Dall’Agnol and Gilsane L. von Poser (2000): "The use of complex polysaccharides in the management of metabolic diseases: the case of ''Solanum lycocarpum'' fruits". ''Journal of Ethnopharmacology'', volume 71, issues 1–2, pp. 337–41. Aline M. Pascoal, Maria C. B. Di-Medeiros, Karla A. Batista, Maria I. G. Leles, Luciano Moraes Lião, and Kátia F. Fernandes (2013): "Extraction and chemical characterization of starch from ''S. lycocarpum'' fruits". ''Carbohydrate Polymers'', volume 98, issue 2, pp. 1304–10. Melissa G. Morais, Guilherme A. F. da Costa, Álan A. Aleixo, Graziela T. de Oliveira, Lucas F. Alves, and Joaquim M. Duarte-Almeida (2015): "Antioxidant, antibacterial and cytotoxic potential of the ripe fruits of ''Solanum lycocarpum'' A. St. Hil. (Solanaceae)". ''Journal of Natural Product Research'', volume 29, issue 5, pp. 480–83 A. C. P. Oliveira, D. C. Endringer, R. J. P. Araújo, M. G. L. Brandão, and M. M. Coelho (2003): "The starch from ''Solanum lycocarpum'' St. Hill. fruit is not a hypoglycemic agent". ''Brazilian Journal of Medical and Biological Research'', volume 36, issue 4, pp. 525–30. Ludmilla M. de S. Aguiar (2005): "First record on the use of leaves of ''Solanum lycocarpum'' (Solanaceae) and fruits of ''Emmotum nitens'' (Icacinacea) by ''Platyrrhinus lineatus'' (E. Geoffroy) (Chiroptera, Phyllostomidae) in the Brazilian Cerrado". ''Revista Brasileira de Zoologia'', volume 22, issue 2, pp. 509–10.
Edmund C. Jaeger Edmund Carroll Jaeger, D.Sc.,University of California, Riverside, Science Library (January 28, 1887 – August 2, 1983) was an American biologist known for his works on desert ecology. He was born in Loup City, Nebraska to Katherine (née Gunther ...
(1944):
A Source-Book Of Biological Names And Terms
'. pp. 41, 153. Published by Charles C. Thomas, Illinois.
Gilmarcio Z. Martins, Raquel R. D. Moreira, Cleopatra S. Planeta, Adélia E. Almeida, Jairo K. Bastos, Lígia Salgueiro, Carlos Cavaleiro, and Maria do Céu Sousa (2015): "Effects of the extract and glycoalkaloids of ''Solanum lycocarpum'' St. Hill on ''Giardia lamblia'' trophozoites" ''Pharmacognosy Magazine'', volume 11, suplement 1, pp. S161–65. Ana P. A. Pereira, Célio F. F. Angolini, Bruno N. Paulino, Leonardo B. C. Lauretti, Eduardo A. Orlando, Joyce G. S. Silva, Iramaia A. Neri-Numa, Jane D. R. P. Souza, Juliana A. L. Pallone, Marcos N. Eberlin, and Glaucia M. Pastore (2019): "A comprehensive characterization of Solanum lycocarpum St. Hill and Solanum oocarpum Sendtn: Chemical composition and antioxidant properties". ''Food Research International'', volume 124, pp. 61–69. Mariza A. Miranda, Renata F. J. Tiossi, Mariana R. da Silva, Kelly C. Rodrigues, Christian C. Kuehn, Luiz G. R. Oliveira, Sérgio Albuquerque, James D. McChesney, Claudio M. Lezama‐Davila, Angelica P. Isaac‐Marquez, and Jairo K. Bastos (2013): "In vitro Leishmanicidal and cytotoxic activities of the glycoalkaloids from ''Solanum lycocarpum'' (Solanaceae) fruits". ''Chemistry & Biodiversity'', volume 10, issue 4, pp. 642–48. Lilian V. A. Pinto, Edvaldo A. A. Da Silva, Antonio C. Davide, Valquíria A. Mendes De Jesus, Peter E. Toorop, Henk W. M. Hilhorst (2007): "Mechanism and control of ''Solanum lycocarpum'' seed germination". ''Annals of Botany'', volume 100, issue 6, pp. 1175–1187. L. C. Clementino, A. M. A. Velásquez, T. G. Passalacqua, L. Almeida, M. A. S. Graminha, G. Z. Martins, L. Salgueiro, C. Cavaleiro, M. C. Sousa, R. R. D. Moreira (2018): "In vitro activities of glycoalkaloids from the ''Solanum lycocarpum'' against ''Leishmania infantum''". ''Revista Brasileira de Farmacognosia'', volume 28, issue 6, pp. 673–77. M. A. Soares, L. P. Sánchez, J. B. Isaac Júnior, E. S. Pereira, E. M.s Pires, G. L. D. Leite, and J. C. Zanuncio (2019): "''Solanum lycocarpum'' Saint Hilaire (Solanaceae) is host plant of ''Leucanella memusae'' (Walker) (Lepidoptera: Saturniidae: Hemileucinae) in Brazilian mountain grasslands". ''Florida Entomologist'', volume 102, issue 3, pp. 624–26. A. L. T. De Souza, G. W. Fernandes, J. E. C. Figueira, and M. O. Tanaka (1998): "Natural history of a gall-inducing weevil ''Collabismus clitellae'' (Coleoptera: Curculionidae) and some effects on its host plant ''Solanum lycocarpum'' (Solanaceae) in Southeastern Brazil". ''Annals of the Entomological Society of America'', volume 91, issue 4, pp. 404–09.
{{Taxonbar, from=Q3280462 lycocarpum Flora of Brazil Flora of the Cerrado Tropical fruit Edible Solanaceae