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Neotropical Moraceae

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Jos Eduardo Lahoz da Silva Ribeiro


Universidade Estadual de Londrina, Paran, Brazil.

Description
Trees, shrubs, hemi-epiphytes or herbs and very rarely lianas; plants monoecious, dioecious or androdioecious, unarmed (except Maclura Nutt.), latex present in all plant parts, white, brown (coffee with milk), yellow or orange. Leaves simple, alternate or rarely opposite (Bagassa Aubl.) arranged in spiral in Ficus L. and Dorstenia L. (rosulate) or distichous, at least in the reproductive branches in other genera, venation pinnate, rarely trinerved (Bagassa), brochidodromous; stipules usually conspicuous, terminal, fully amplexicaul, partially amplexicaul to lateral, their scars forming a complete ring or, if not amplexicaul, with a scar opposite the insertion of the leaves. Inflorescences unisexual or bisexual, usually in pairs in the leaf axil or more on short shoots, forming unisexual or bisexual synflorescences (Tribe Ficeae and Castilleae), structurally complex, racemose, spicate, globose, capitate or urceolate receptacles (= syconium, fig).Flowers unisexual, free or immersed in the receptacle, monochlamydeous or achlamydeous, staminate flowers with 2-4 (-7) free or connate tepals or tepals absent, stamens (3-) 4 (-5), or reduced to 1, filaments straight or inflexed before anthesis; pistillode generally absent; pistillate flowers with (3-) 4 (-7) free or connate tepals; ovary free or adnate to tepals, stigmas 2 or 1, ovule 1 with subapical placentation. Fruits achenes, usually associated with afleshy perianth or other structures that become fleshy, like bracts and/or receptacle.

Notes on delimitation

The family Moraceae is currently placed in the order Rosales (APG III) as sister to the familyUrticaceae (Sytsma et al., 2002), here including the genera Cecropia Loefl., Pourouma Aubl. and Coussapoa Aubl., previously placed in the family Cecropiaceae.

The family Moraceae is currently divided into six tribes: Artocarpeae, Morea, Maclureae, Dorstenieae, Castilleae and Ficeae (Ribeiro, 2007; Clement & Weiblen, 2009).

Distribution in the Neotropics

The family is distributed throughout the Neotropics, with a major center of diversity in northern South America, specifically the Amazon rainforests, where all Neotropical genera are represented. The genus Dorstenia has a concentration of species mainly in northern Central America and the Greater Antilles and in the Atlantic forests of southeastern Brazil. In the genus Morus L., M. celtidifolia Kunth is distributed primarily from Mexico to Honduras and M. insignis Bureau in Central America and the Andean region of South America.

Distinguishing characters (always present)



The most obvious features of the family are the presence of latex in all parts of the plant, andterminal stipules. In times of high water stress, some species may not exude evident latex. The stipules are small and often caducous in some species of Clarisia Ruiz & Pav. andSorocea A.St-Hil. Even in the absence of stipules, it is possible to observe the scars left by these structures. The flowers are small, tapaloid or naked; unisexual; gynoecium with one or two stigmas; ovaryuniovulate and placentation subapical.

Other important characters



Generally woody plants, from trees to hemiepiphytes and lianas. Leaves alternate. Inflorescences structurally complex, racemose, spicate, globose, capitate or an urceolatereceptacle (= syconium, fig).

Key differences from similar families

The presence of terminal stipule or stipule scar, associated with the presence of latex in all parts of the plant, makes Moraceae difficult to confuse with other families. Some species ofCoussapoa (Urticaceae) have aqueous whitish latex in the branches and stipules terminals, besides the usual hemi-epiphytic habit, which can be confused with species of Ficus, although waxy glandular spots in the base of blade underside do not occur in Coussapoa spp.

Number of genera
19 genera: Tribe Artocarpeae Batocarpus H.Karst. (3 spp.) Clarisia Ruiz & Pav. (3 spp.) Tribe Moreae Bagassa Aubl. (1 spp.) Morus L. (2 spp.) Sorocea A. St-Hil. (14 spp.) Trophis P.Browne (5 spp.) Tribe Maclureae Maclura Nutt. (2 spp.)

Tribe Dorstenieae Brosimum Sw. (15 spp.) Dorstenia L. (46 spp.) Helianthostylis Baill. (2 spp.) Trymatococcus Poepp. & Endl. (2 spp.) Tribe Castilleae Castilla Cerv. (3 spp.) Helicostylis Trcul (7 spp.) Maquira Aubl. (4 spp.) Naucleopsis Miq. (22 spp.) Perebea Aubl. (9 spp.) Poulsenia Eggers (2 spp.) Pseudolmedia Trcul (15 spp.) Tribe Ficeae Ficus subg. Urostigma (Gasparrini) Miq. sect. Americana Miq.(ca. 100 spp.) Ficus subg. Pharmacosycea (Miq.) Miq. sect. Pharmacosycea (ca. 20 spp.)

Useful tips for generic identification

Images of some genera and species of the Central Brazilian Amazon can be accessed at:http://brahms.inpa.gov.br/bol/content/Groups/PFRD/resources/PFRD_1999_Moraceae.pdf

Notable genera and distinguishing features


Bagassa: leaves opposite, venation trinerved. Batocarpus: canopy tree, stems and the shallow roots covered with elevated red lenticels. Brosimum: inflorescences usually bisexual and globose, covered with peltate bracts. Clarisa racemosa Ruiz & Pav.: canopy tree, stems and the shallow roots covered with elevated red lenticels. Clarisa ilicifolia (Spreng.) Lanji. & Rossberg: understory treelets; leaf margin and apex spinulose. This species is often confused with Sorocea guilleminiana Gaudich., differentiated by theprominent midrib on the upper surface of the blade and by the presence of uncinate trichomes. Dorstenia: herbaceous habit; inflorescence a receptacle with variable shape - orbicular, elliptic,obovate, angulate, stellate or furcate. Ficus subg. Urostigma sect. Americana: habit hemi-epiphyte; inflorescence a syconium; with a waxy glandular spot at the base of the midrib of the lower surface of the leaf. Ficus subg. Pharmacosycea sect. Pharmacosycea: trees; inflorescence a syconium; with 2 waxyglandular spots in the axils of the basal lateral veins of the lower surface of the leaf. Maclura: tree or liana; often armed with axillary thorns. Morus: leaves alternate distichous, the margin crenate to serrate. Sorocea: lamina is slightly to pronouncedly inequilateral; leaf margin mostly dentate todenticulate, sometimes the teeth and acumen spinulose. Trymatococcus: leaves with globose-capitate pluricellular hairs; inflorescences bisexual.

Genera of tribe Castilleae (Castilla, Helicostylis, Maquira, Naucleopsis, Perebea, Poulsenia,Pseudolmedia): trees with the architectural model of Cook and the inflorescences usually with involucres of bracts.

Status

All genera are endemic and restricted to the neotropics, with the exception of Morus, Trophis,Dorstenia and Ficus, which have a pantropical distribution pattern. The genus Artocarpus J.R. & G. Foster and Morus alba L. are naturalized in the Neotropics, grown for their edible fruits.

General notes

The inflorescences of the family are among the most complex in Angiosperms. These are often associated with pollination systems in which plants provide breeding sites as a reward for the pollinator. For the genus Ficus entomophily is well documented where the pollination is carried out by wasps of the family Agaonidae (Chalcidoidae, Hymenoptera). The two neotropical sections Americana and Pharmacosycea are pollinated by the wasp generaPegoscapus and Tetrapus respectively.

The Neotropical Castilla elastica Sess (Sakai, 2001), the New Guinea Antiaropsis decipiensK. Shum. (Zerega et al., 2004) and probably all species of Tribe Castilleae are pollinated by thrips (Thysanoptera). Although the family always have been cited, even in the recent scientific literature, as an example of a wind-pollinated group, probably only 6% of all species of the family are really anemophilous (Ribeiro, 2007).

Important literature
Berg, C.C. 1972. Olmedieae and Brosimeae. Flora Neotropica 7. Hafner Publishing Company, New York.

Berg, C.C. Moraceae. 1992. In: A.R.A. Grts-van Rijn (ed.). Flora of the Guianas. vol. 21. Koenigstein, Germany. 82p.

Berg, C.C. 1998b. Moraceae (excl. Ficus). In: G. Harling & L. Andersson (eds.). Flora of Ecuador. Vol. 27B. Arlv, Sweden. 128p.

Berg, C.C. 2001. Moreae, Artocarpeae, and Dorstenia (Moraceae). With introductions to the family and Ficus and with additions and corrections to Flora Neotropica Monograph 7. Flora Neotropica Monograph 83. The New York Botanical Garden, New York. p. 347.

Berg, C.C. & M.E.E. Hijman. 1999. The genus Dorstenia (Moraceae). Ilicifolia 2:1-211.

Berg, C.C. & X. Villavicencio. 2004. Taxonomic studies on Ficus (Moraceae) in the West Indies, extra-Amazonian Brazil, and Bolivia. Ilicifolia 4:1-132.

Carauta, J.P.P. 1978. Dorstenia L. (Moraceae) do Brasil e Pases Limtrofes. Rodriguesia 29(44): 52-223.

Clement, W.L. & Weiblen, G.D. 2009. Morphological Evolution in the Mulberry Family (Moraceae). Systematic Botany 34(3): 530-552.

Miquel, F.A.W. 1853 (1965). Urticineae. In: C.F.P. von Martius (ed.). Flora Brasiliensis vol. 4 part. 1, pp. 79-170. J. Cramer, Weinheim.

Ribeiro, J.E.L.S. 2007. Studies on the phylogeny, taxonomy and evolution of reproductive characters in Moraceae Gaudich. Universidade Estadual de Campinas (Ph.D. thesis). 124p. (in Portuguese).

Rohwer, J.G. 1993. Moraceae. In: K. Kubitzki, J.G. Rohwer & V. Bittrich (eds.). The Families and Genera of Vascular Plants. Vol. II. Spring-Verlag, Berlin. 438-453.

Sakai, S. 2001. Thrips pollination of androdioecious Castilla elastica (Moraceae) in a seasonal tropical forest. American Journal of Botany 88(9): 1527-1534. Sytsma, K.J.; Morawetz, J.; Pires, J.C.; Nepokroeff, M.; Conti, E.; Zjhra, M.; Hall, J.C. & Chase, M.W. 2002. Urticalean Rosids: circumscription, Rosid ancestry, and phylogenetics based on rbcL, trnL-F, and ndhF sequences. American Journal of Botany 89(9): 1531-1546. Zerega, N.J.C., L.A. Moundy & G.D. Weiblen. 2004. Pollination in the New Guinea endemicAntiaropsis decipiens (Moraceae) is mediated by a new species of thrips, Thrips antiaropsidis sp. nov. (Thysanoptera: Thripidae). International Journal of Plant Sciences 165(6):1017-1026.

How to cite
Ribeiro, J.E.L.S. (2011). Neotropical Moraceae. In: Milliken, W., Klitgrd, B. & Baracat, A. (2009 onwards), Neotropikey - Interactive key and information resources for flowering plants of the Neotropics. http://www.kew.org/science/tropamerica/neotropikey/families/Moraceae.htm.