Monographs Details:
Authority:
Berg, Cornelius C. 2001. Moreae, Artocarpeae, and (Moraceae): With introductions to the family and and with additions and corrections to Flora Neotropica Monograph 7. Fl. Neotrop. Monogr. 83: 1-346. (Published by NYBG Press)
Berg, Cornelius C. 2001. Moreae, Artocarpeae, and
Family:
Moraceae
Moraceae
Synonyms:
Balanostreblus, Pseudosorocea, Trophisomia, Paraclarisia, Sorocea bonplandii (Baill.) W.C.Burger, Lanj. & Wess.Boer, Balanostreblus ilicifolius Kurz, Sorocea guilleminiana Gaudich., Pseudosorocea bonplandii Baill., Sorocea bonplandii (Baill.) W.C.Burger, Lanj. & Wess.Boer, Trophisomia edulis Rojas Acosta, Sorocea sprucei subsp. saxicola (Hassl.) C.C.Berg, Paraclarisia amazonica Ducke, Sorocea duckei W.C.Burger
Balanostreblus, Pseudosorocea, Trophisomia, Paraclarisia, Sorocea bonplandii (Baill.) W.C.Burger, Lanj. & Wess.Boer, Balanostreblus ilicifolius Kurz, Sorocea guilleminiana Gaudich., Pseudosorocea bonplandii Baill., Sorocea bonplandii (Baill.) W.C.Burger, Lanj. & Wess.Boer, Trophisomia edulis Rojas Acosta, Sorocea sprucei subsp. saxicola (Hassl.) C.C.Berg, Paraclarisia amazonica Ducke, Sorocea duckei W.C.Burger
Description:
Genus Description - Trees or shrubs, dioecious; twigs usually with conspicuous lenticels; uncinate hairs lacking. Leaves alternate and distichous; lamina pinnately veined; margin (spinulose-)dentate to entire; stipules free, lateral. Inflorescences in pairs or solitary (or several together on short-shoots) in the axils of the leaves or below the leaves, racemose to spicate to subcapitate (or uniflorous), bracteate; bracts small, basally attached to peltate, cordiform, ovate, or suborbiculate; staminate flowers with 4 tepals; tepals decussate-imbricate, mostly basally connate; stamens (3-)4, free, straight in the bud, at anthesis often bent inwards or outwards; anthers basi- to dorsifixed, extrorse (to latrorse), the connective mostly broad, often (gland-like) apiculate; pistillode normally absent; pistillate flowers with a tubular, 4-lobed to subentire perianth, its upper and lower part usually ± different (in diameter, surface, indumentum); lower part of the ovary adnate to the lower part of the perianth, the ovary gradually narrowed into the style; stigmas 2, equal, short, mostly tongue-shaped; the rachis, the pedicel, or sometimes also the peduncle more or less carnose and red (or orange) in fruit; fruiting perianth enlarged, fleshy, red, pink, or orange, (always?) turning black(ish) at full maturity, mostly on a(n elongated) pedicel. Fruit large, mostly globose to ellipsoid, adnate to the perianth; endocarp thin, subcoriaceous to subcrustaceous; seed large, without endosperm; testa thin, with a thickened, (sub)orbicular part below the hilum; embryo (always?) green, longitudinally aligned; cotyledons very unequal, the smaller one minute and plane, enclosed by the larger, thick, curved, conduplicate, and lobed cotyledon; radicle short, apical.
Genus Description - Trees or shrubs, dioecious; twigs usually with conspicuous lenticels; uncinate hairs lacking. Leaves alternate and distichous; lamina pinnately veined; margin (spinulose-)dentate to entire; stipules free, lateral. Inflorescences in pairs or solitary (or several together on short-shoots) in the axils of the leaves or below the leaves, racemose to spicate to subcapitate (or uniflorous), bracteate; bracts small, basally attached to peltate, cordiform, ovate, or suborbiculate; staminate flowers with 4 tepals; tepals decussate-imbricate, mostly basally connate; stamens (3-)4, free, straight in the bud, at anthesis often bent inwards or outwards; anthers basi- to dorsifixed, extrorse (to latrorse), the connective mostly broad, often (gland-like) apiculate; pistillode normally absent; pistillate flowers with a tubular, 4-lobed to subentire perianth, its upper and lower part usually ± different (in diameter, surface, indumentum); lower part of the ovary adnate to the lower part of the perianth, the ovary gradually narrowed into the style; stigmas 2, equal, short, mostly tongue-shaped; the rachis, the pedicel, or sometimes also the peduncle more or less carnose and red (or orange) in fruit; fruiting perianth enlarged, fleshy, red, pink, or orange, (always?) turning black(ish) at full maturity, mostly on a(n elongated) pedicel. Fruit large, mostly globose to ellipsoid, adnate to the perianth; endocarp thin, subcoriaceous to subcrustaceous; seed large, without endosperm; testa thin, with a thickened, (sub)orbicular part below the hilum; embryo (always?) green, longitudinally aligned; cotyledons very unequal, the smaller one minute and plane, enclosed by the larger, thick, curved, conduplicate, and lobed cotyledon; radicle short, apical.
Discussion:
HistoryThe genus Sorocea was established by Saint-Hilaire (1821) in a study of the very unequal cotyledons in the genus. In 1844, Gaudichaud published plates of some species, without descriptions. The first treatment with descriptions of species, seven in total, was by Miquel (1853) in Martius Flora Brasiliensis. Some additional species were published by various authors up to 1962, when a revision of the genus was published by Burger, Lanjouw, and Wessels Boer. This revision comprised 22 species, of which seven were new. In 1966, Cuatrecasas added three species. A precursor to the present revision was published by Berg and Akkermans (1985): 16 species were recognized, one of them new; and in four species, two subspecies were recognized. Conclusions based on more recent collections have reduced the number of species to 14, including two species and one new subspecies, which have been more recently described (Berg in Berg & Franco, 1996).Some genera recognized as distinct from Sorocea, Balanostreblus Kurz (1873), Pseudosorocea Baillon (1875b), and Paraclarisia Ducke (1939), were included in the genus by Burger et al. (1962), Paraclarisia being reduced to a subgenus. Trophisomia Rojas Acosta (1914) proved to be a synonym of Sorocea as well.MorphologyHabit: Most Sorocea species are shrubs or small trees, but several species are trees, which can reach a height of 20 or even 25 m. The majority of the Sorocea species are evergreen; only S. duckei and S. sprucei are deciduous. The lenticels are mostly conspicuous and prominent. The latex is white and usually copious.Leaves: The lamina is slightly to pronouncedly inequilateral, mostly coriaceous, sometimes subcoriaceus to chartaceous. The leaf margin is mostly dentate or denticulate. In Sorocea guilleminiana and S. bonplandii, the teeth, as well as the leaf acumen, are spinulose. In S. hilarii and S. sprucei, the teeth and leaf acumen are sometimes very shortly spinulose. In most species, the tertiary venation is scalariform, but in some, including S. affinis and S. muriculata, it is reticulate.Indumentum: Two types of trichomes are found:1. Unicellular hairs on all parts of the plant, but often sparse. 2. Globose-capitate, pluricellular (glandular) hairs on young parts of the plant, often frequent on the lower leaf surface. In some species (e.g., Sorocea briquetii, S. guilleminiana, and S. ruminata), they are abundantly present on the lower part of the perianth of the pistillate flower. There they form or exude a substrate for a fungus with a dense white mycelium, which can be found as white blotches on the enlarged fruiting perianth. The fungus sometimes produces "fruit-bodies."Inflorescences: The inflorescences are borne solitary or in pairs in the leaf axils, or several (to 4 or 6) together on axillary short-shoots. In most species (such as in Sorocea hilarii), the inflorescences also appear on the older wood, occasionally on the trunk. The inflorescences are basically racemose with the flowers ± less distinctly adaxial on the rachis. The staminate flowers are sessile in most species, but are pedicellate in affinis, S. bonplandii, S. guilleminiana, and S. hilarii. In S. affinis, the flower can be either pedicellate or sessile. Pistillate flowers are often sessile at anthesis, but afterwards a pedicel is developed. Pedicels present at anthesis usually elongate afterwards. The bracts are small and basally attached to peltate. They cover the very young flowers, especially in staminate inflorescences. The bracts are subcoriaceous to coriaceous, but submembranaceous in S. duckei and S. sprucei.Staminate Flower: The perianth of the staminate flower consists of four decussate-imbricate tepals, mostly basally or up to halfway connate. Sessile flowers often have a very broad base. The flowers normally have four isomorphic stamens, but the number of stamens is often reduced to three. In species where this reduction occurs, one can also find anisomorphy in the androecium. The pair of stamens opposite the outer tepals can have distinctly shorter filaments and sometimes also smaller anthers. Moreover, aberrations in the length of the filaments and the dimensions of the anthers are not uncommon. In Sorocea duckei and S. sprucei, the filaments are very short and the anthers subsessile. Although in most species, the filaments are rather thick and shorter than the perianth, they are slender and longer than the perianth in S. trophoides. At anthesis, the stamens are straight, bent inwards or outwards. Incurved stamens are, although not very clearly, associated with a broadly sessile flower. When the stamens are straight, the anther is often perpendicular, pointing outwards. The anthers are extrorse (to slightly latrorse). The connective is mostly broad and is often extended into a gland-like apiculum. The pollen grains of several Sorocea species have been described by Niezgoda and Nowaczyk (1976).Fruit: White blotches on the fruiting perianth are caused by fungi. The embryo is (always?) green and has very unequal cotyledons, a phenomenon described by Saint-Hilaire, who established in the same paper the genus Sorocea (1821).POLLINATION AND DISPERSALNothing is known about pollination in Sorocea. None of the features of flowers and inflorescences suggest the occurrence of wind pollination. Moreover, most species are components of the understory of evergreen forest. On the other hand, there are not clear adaptations to insect pollination, certainly not to insect pollination based on breeding. The common occurrence of pluricellular trichomes on the perianth of pistillate flowers suggests that these trichomes might play a role in pollination, as substitutes for pollen grains. These trichomes are apparently nutritious as they provide a substrate for fungi.According to some label data, the fruits (fruiting perianths) are eaten by birds. According to Kubitzki and Ziburski (1994), the fruits (fruiting perianths) of Sorocea duckei float due to buoyancy provided by air enclosed by the folded (large) cotyledon.Systematic Relationships and Subdivision of the GenusThe genus Sorocea is in various respects quite distinct. It is the only genus among the non-anemophilous ones having predominantly racemose inflorescences, which may have implications for pollination (see above). It has clearly defined and ± separate staminate flowers, in contrast to the genera that are probably its closest relatives, such as Clarisia and Batocarpus. Furthermore, the perianth at least in the fruiting state is clearly differentiated into a lower part adnate to the ovary and an upper part free from the ovary.The genus has been divided into two subgenera by Burger et al. (1962: 468): Sorocea and Paraclarisia, the latter comprising only S. duckei and S. sprucei. In subg. Sorocea, one can recognize a group with a spinulose-dentate leaf margin and/or a spinulose acumen of the lamina, comprising S. bonplandii, S. guilleminiana, and S. hilarii, as somewhat distinct from the rest of the subgenus. These three species are morphologically very close and often not easy to distinguish. In the larger group of species, the margin of the lamina is entire or varies from entire to dentate, but neither the teeth nor the acumen of the lamina are spinulose. The species of this group are closely related and it is often difficult to distinguish them. The exception is the clear-cut S. jaramilloi.
HistoryThe genus Sorocea was established by Saint-Hilaire (1821) in a study of the very unequal cotyledons in the genus. In 1844, Gaudichaud published plates of some species, without descriptions. The first treatment with descriptions of species, seven in total, was by Miquel (1853) in Martius Flora Brasiliensis. Some additional species were published by various authors up to 1962, when a revision of the genus was published by Burger, Lanjouw, and Wessels Boer. This revision comprised 22 species, of which seven were new. In 1966, Cuatrecasas added three species. A precursor to the present revision was published by Berg and Akkermans (1985): 16 species were recognized, one of them new; and in four species, two subspecies were recognized. Conclusions based on more recent collections have reduced the number of species to 14, including two species and one new subspecies, which have been more recently described (Berg in Berg & Franco, 1996).Some genera recognized as distinct from Sorocea, Balanostreblus Kurz (1873), Pseudosorocea Baillon (1875b), and Paraclarisia Ducke (1939), were included in the genus by Burger et al. (1962), Paraclarisia being reduced to a subgenus. Trophisomia Rojas Acosta (1914) proved to be a synonym of Sorocea as well.MorphologyHabit: Most Sorocea species are shrubs or small trees, but several species are trees, which can reach a height of 20 or even 25 m. The majority of the Sorocea species are evergreen; only S. duckei and S. sprucei are deciduous. The lenticels are mostly conspicuous and prominent. The latex is white and usually copious.Leaves: The lamina is slightly to pronouncedly inequilateral, mostly coriaceous, sometimes subcoriaceus to chartaceous. The leaf margin is mostly dentate or denticulate. In Sorocea guilleminiana and S. bonplandii, the teeth, as well as the leaf acumen, are spinulose. In S. hilarii and S. sprucei, the teeth and leaf acumen are sometimes very shortly spinulose. In most species, the tertiary venation is scalariform, but in some, including S. affinis and S. muriculata, it is reticulate.Indumentum: Two types of trichomes are found:1. Unicellular hairs on all parts of the plant, but often sparse. 2. Globose-capitate, pluricellular (glandular) hairs on young parts of the plant, often frequent on the lower leaf surface. In some species (e.g., Sorocea briquetii, S. guilleminiana, and S. ruminata), they are abundantly present on the lower part of the perianth of the pistillate flower. There they form or exude a substrate for a fungus with a dense white mycelium, which can be found as white blotches on the enlarged fruiting perianth. The fungus sometimes produces "fruit-bodies."Inflorescences: The inflorescences are borne solitary or in pairs in the leaf axils, or several (to 4 or 6) together on axillary short-shoots. In most species (such as in Sorocea hilarii), the inflorescences also appear on the older wood, occasionally on the trunk. The inflorescences are basically racemose with the flowers ± less distinctly adaxial on the rachis. The staminate flowers are sessile in most species, but are pedicellate in affinis, S. bonplandii, S. guilleminiana, and S. hilarii. In S. affinis, the flower can be either pedicellate or sessile. Pistillate flowers are often sessile at anthesis, but afterwards a pedicel is developed. Pedicels present at anthesis usually elongate afterwards. The bracts are small and basally attached to peltate. They cover the very young flowers, especially in staminate inflorescences. The bracts are subcoriaceous to coriaceous, but submembranaceous in S. duckei and S. sprucei.Staminate Flower: The perianth of the staminate flower consists of four decussate-imbricate tepals, mostly basally or up to halfway connate. Sessile flowers often have a very broad base. The flowers normally have four isomorphic stamens, but the number of stamens is often reduced to three. In species where this reduction occurs, one can also find anisomorphy in the androecium. The pair of stamens opposite the outer tepals can have distinctly shorter filaments and sometimes also smaller anthers. Moreover, aberrations in the length of the filaments and the dimensions of the anthers are not uncommon. In Sorocea duckei and S. sprucei, the filaments are very short and the anthers subsessile. Although in most species, the filaments are rather thick and shorter than the perianth, they are slender and longer than the perianth in S. trophoides. At anthesis, the stamens are straight, bent inwards or outwards. Incurved stamens are, although not very clearly, associated with a broadly sessile flower. When the stamens are straight, the anther is often perpendicular, pointing outwards. The anthers are extrorse (to slightly latrorse). The connective is mostly broad and is often extended into a gland-like apiculum. The pollen grains of several Sorocea species have been described by Niezgoda and Nowaczyk (1976).Fruit: White blotches on the fruiting perianth are caused by fungi. The embryo is (always?) green and has very unequal cotyledons, a phenomenon described by Saint-Hilaire, who established in the same paper the genus Sorocea (1821).POLLINATION AND DISPERSALNothing is known about pollination in Sorocea. None of the features of flowers and inflorescences suggest the occurrence of wind pollination. Moreover, most species are components of the understory of evergreen forest. On the other hand, there are not clear adaptations to insect pollination, certainly not to insect pollination based on breeding. The common occurrence of pluricellular trichomes on the perianth of pistillate flowers suggests that these trichomes might play a role in pollination, as substitutes for pollen grains. These trichomes are apparently nutritious as they provide a substrate for fungi.According to some label data, the fruits (fruiting perianths) are eaten by birds. According to Kubitzki and Ziburski (1994), the fruits (fruiting perianths) of Sorocea duckei float due to buoyancy provided by air enclosed by the folded (large) cotyledon.Systematic Relationships and Subdivision of the GenusThe genus Sorocea is in various respects quite distinct. It is the only genus among the non-anemophilous ones having predominantly racemose inflorescences, which may have implications for pollination (see above). It has clearly defined and ± separate staminate flowers, in contrast to the genera that are probably its closest relatives, such as Clarisia and Batocarpus. Furthermore, the perianth at least in the fruiting state is clearly differentiated into a lower part adnate to the ovary and an upper part free from the ovary.The genus has been divided into two subgenera by Burger et al. (1962: 468): Sorocea and Paraclarisia, the latter comprising only S. duckei and S. sprucei. In subg. Sorocea, one can recognize a group with a spinulose-dentate leaf margin and/or a spinulose acumen of the lamina, comprising S. bonplandii, S. guilleminiana, and S. hilarii, as somewhat distinct from the rest of the subgenus. These three species are morphologically very close and often not easy to distinguish. In the larger group of species, the margin of the lamina is entire or varies from entire to dentate, but neither the teeth nor the acumen of the lamina are spinulose. The species of this group are closely related and it is often difficult to distinguish them. The exception is the clear-cut S. jaramilloi.
Distribution and Ecology: The genus ranges from southern Mexico to Argentina. It is absent from the West Indies (except for Curaçao) and poorly represented in the Guianas. Most of the species are elements of lowland rain forest, being understory treelets or shrubs or canopy trees. Only Sorocea trophoides is a (sub)montane species, although it is also found at low elevations. Sorocea bonplandii, in the southern part of the genus range, and S. sprucei, with a disjunct distribution in South America, are species of (rather) dry forest or shrub vegetation. Sorocea duckei is an element of varzea vegetation in the Amazon Basin. Three elements can be distinguished: a group of species in Central America and the Pacific Coastal regions of Colombia and Ecuador: a group of species in the Amazon Basin, in particular the upper Amazon Basin: and a small group of species south of the Amazon Basin. Sorocea pubivena and S. guilleminiana are the most widespread species, occurring in more than one phytogeographical region. The other species have smaller ranges of distribution, and some, such as S. jaramilloi, S. ruminata, and S. sarcocarpa, have very small ranges, occupying only a part of the phytogeographic regions they are associated with.