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Acta Botanica Brasilica 28(3): 422-433

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Acta Botanica Brasilica 28(3): 422-433 Acta Botanica Brasilica 28(3): 422-433. 2014. doi: 10.1590/0102-33062014abb3341 Morphology of fruits, seeds, seedlings and saplings of three species of Macrolobium Schreb. (Leguminosae, Caesalpinioideae) in the Brazilian Amazon floodplain1 Gildo Vieira Feitoza2,5, João Ubiratan Moreira dos Santos3, Ely Simone Cajueiro Gurgel2 and Denise Maria Trombert Oliveira4 Received: 29 August, 2013. Accepted: 26 March, 2014 ABSTRACT The evaluation of the post-germination phases of plants allows the recognition of transitional structures that support the relationships among taxa and their establishment. Together with fruit and seeds, seedlings and saplings provide useful characters for identifying many species. This study aimed to describe and characterize the morphology of the fruits, seeds, seedlings and saplings of Macrolobium acaciifolium, M. bifolium and M. pendulum, as well as to evaluate the ecological, taxonomic and phylogenetic value of the structures identified. In this work, the seeds were sown in plastic trays with sterilized sand and sawdust, without pre-germination treatment. The fruits of the three species are woody legumes, dehiscent fruit or indehiscent fruit. The seeds have seed coats with or without venation and variable embryos, with plumules differing among the three species; this trait can be helpful in distinguishing among the species studied. The seedlings are of the cryptocotylar-hypogeal-reserve or phanerocotylar-epigeal-reserve type. Macrolobium bifolium and M. pendulum are similar species, differing in only a few characters. These characters have taxonomic value and aid species identification. The characteristics of M. pendulum are described here for the first time. Key words: Detarieae, embryo, Fabaceae, plumule, post-germination development Introduction mosoideae, Calliandra and Mimosa have been evaluated (Parra 1984). In Papilionoideae, there have been studies The morphological aspects of the initial stages of plant of Phaseoleae (Baudet 1974); Dalbergieae (Lima 1989- development have been poorly investigated in various 90); Genisteae (López et al. 1998); Phaseoleae, Sophoreae, families, principally when compared with the structure of Swartzieae and Tephrosieae (Oliveira 2001); Vatairea (Rod- mature vegetative organs. The study of the initial morphol- rigues & Tozzi 2007a); the genistoid clade (Rodrigues & ogy provides knowledge about transitional or ephemeral Tozzi 2007b); Acosmium, Guianodendron and Leptolobium structures that can supply relevant insights into phyloge- (Rodrigues & Tozzi 2008); and Rhynchosia (Rodrigues et netic relationships (Moreira-Coneglian & Oliveira 2006). al. 2012). The studies of Duke & Polhill (1981), Oliveira Likewise, the morphological characters of fruits and seeds (1999a) and Gurgel et al. (2012) evaluated various genera constitute highly reliable criteria for botanical identifica- within the three subfamilies. tion, because they normally have little phenotypic plastic- Although Leguminosae is a well-known family, there ity and are therefore good taxonomic characters (Oliveira have been few studies of its representatives in the Amazon, & Pereira 1984; Oliveira 1999a). In addition, data about notable exceptions including studies conducted in the seedlings, fruits and seeds can contribute information that Venezuelan Amazon (Ricardi 1996; 1999a; 1999b) and in is valuable for analyzing the systematics and ecology of the Brazilian Amazon (Silva et al. 1988; Vieira et al. 1996; various groups of tropical plants (Garwood 1995). Aguiar & Mendonça 2000; Gurgel et al. 2002; Ramos & Fer- Morphological studies of fruits, seeds, seedlings and raz 2008; and Rodrigues et al. 2012). There is also a lack of saplings have helped the characterization of several taxo- data on seedlings that occur in specific environments of the nomic groups in Leguminosae. In Caesalpinioideae, details Amazon, such as the floodplain forests, where species are have been presented for Copaifera (Gurgel 2009). In Mi- subjected to periodic flooding (Silva et al. 1988; Moreira & 1 Based on the Master’s dissertation of the first author 2 Museu Paraense Emílio Goeldi, Av. Magalhães Barata 376, São Braz, CP 399, 66040-170, Belém, PA, Brazil 3 Universidade Federal Rural da Amazônia, Instituto de Ciências Agrárias, Belém, PA, Brazil 4 Universidade Federal de Minas Gerais, Instituto de Ciências Biológicas, Departamento de Botânica, Belo Horizonte, MG, Brazil 5 Author for correspondence: [email protected] Morphology of fruits, seeds, seedlings and saplings of three species of Macrolobium Schreb. (Leguminosae, Caesalpinioideae) in the Brazilian Amazon fl oodplain Moreira 1996; Gama et al. 2003; Maia et al. 2005; Schöngart The present study describes the morphology of fruits, et al. 2005). Such knowledge is important to protecting and seeds and the post-germination phases of Macrolobium understanding the regeneration of these ecosystems. acaciifolium, M. bifolium and M. pendulum, aiming to aid Macrolobium Schreb. (Leguminosae, Caesalpinioideae) the identification of native species of Amazon floodplain is currently included in Detarieae s.l. (Bruneau et al. 2000; forests and to provide a foundation for further phylogenetic 2001; Herendeen et al. 2003). The tribe Detarieae s.l., and ecological studies involving this complex group. comprises all of the genera included in Detarieae, Cyno- metreae and Amherstieae (Léonard 1957); in Detarieae and Amherstieae (Cowan & Polhill 1981); in Detarieae (Polhill Material and methods 1994); and in Detarieae and Macrolobieae (Breteler 1995), Fruits and seeds of Macrolobium acaciifolium, M. bifo- excluding the monotypic genus Umtiza Sim, which occurs lium and M. pendulum were collected from natural popula- in Southern Africa (Herendeen et al. 2003). tions in the Combu Island EPA (48°25’W; 1°25’S). The area Macrolobium comprises approximately 80 species and is is entirely constituted of flooded forest (alluvial rain forest) strictly distributed in the tropics (De la Estrella et al. 2012). and presents Humic Gleysols, with high percentages of silts Approximately 40 species occur in Brazil, 15 of which are and clay and a low percentage of sand (Silva & Sampaio endemic (Martins-da-Silva & Lima 2013). Macrolobium is 1998). According to the Köppen classification system, the the second most representative genus of Detarieae s.l (Lewis climate of the region is type Am, with a mean annual rain- et al., 2005), being used as an ornamental, for timber, to fall of 2500 mm and a mean annual temperature of 27°C produce fish poison and for lumber in construction (Gama (Jardim & Vieira 2001). et al. 2003; Mackinder 2005; Felix-da-Silva et al. 2013). The Two matrix trees were selected for each Macrolobium genus is common in the Brazilian Amazon, mostly in the species studied. The fruits were collected between January Negro River Basin, where it constitutes one of the dominant and April 2012, from directly under the trees or with a floristic elements of floodplain forests (Ducke 1949). trimmer. The fruits were packed into polyethylene bags and The phylogenetic relationships of Detarieae s.l. were transported to the Laboratory of Fruit and Seed Morphology analyzed by Bruneau et al. (2000; 2001), on the basis of trnL of the Museu Paraense Emílio Goeldi for detailed analysis nucleotide sequence data, and by Herendeen et al. (2003), and measurement. Voucher specimens were deposited in the who used morphological data merged with the aligned Herbarium of the Museu Paraense Emílio Goeldi, in Belém, trnL data. Detarieae s.l. has a pantropical distribution, ap- Brazil (MG), under the accession numbers MG-204242 proximately 58% of the species being restricted to Africa, (M. acaciifolium), MG-204243 (M. bifolium) and MG- including Madagascar, 20% being endemic in the Neotrop- 204244 (M. pendulum). ics; and 12% being endemic in tropical Asia (Mackinder Samples (30 fruits and 50 seeds of each species) were 2005). The genus Macrolobium is at the base of the “higher” randomly collected and analyzed. The dimensions (length, Detarieae s.l., constituting a group together with species of width and thickness) were measured with an UltraTech™ the genera Brownea Jacq., Browneopsis Huber, Elizabetha digital fractional caliper (0.01 mm accuracy; General Tools, Schomb. ex Benth., Paloue Aubl. and Ecuadendron D.A. New York, NY, USA). The length was considered the dis- Neill. (Herendeen et al. 2003). This underscores the state- tance from the apex to the base; the width was measured ment made by Bruneau et al. (2000): that the Macrolobium from the dorsal to ventral regions, and the thickness between group (sensu Cowan & Polhill 1981) or complex (sensu valves was determined. For observation and illustration of Léonard 1957) has a complicated history. Bruneau et al. fruits and seeds, we used a stereomicroscope (Motic, SMZ- (2000) also stated that the relationships of the Macrolobium 168 series, Vancouver, Canada) and a camera lucida. The group requires a critical evaluation of morphological and terminology was based on Gunn (1981; 1991), Barroso et anatomical characters in order to examine the possibility al. (1999) and Oliveira (1999b). of a close affinity between New World Macrolobium and For the description of plumule development, three members of the South American Brownea group. embryos of each species were selected and samples of the The Environmental Protection Area of Combu Island embryo axis were fixed in 2.5% glutaraldehyde in 0.1 M (EPA of Combu Island), in the state of Pará, Brazil, presents sodium phosphate buffer pH 7.2
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