Flora - Morphology, Distribution, Functional Ecology of Plants
Volume 206, Issue 6, June 2011, Pages 575-584
Attalea phalerata and Bactris glaucescens (Arecaceae, Arecoideae): Phenology and pollination ecology in the Pantanal, Brazil
Abstract
We compared the reproductive and vegetative phenology and pollination ecology of sympatric Attalea phalerata and Bactris glaucescens palms (Arecaceae) in the Pantanal, Brazil, in a riparian forest area subject to periodic flooding. Attalea phalerata has a solitary stem and produces staminate, pistillate and rarely bisexuals inflorescences that open during the day. Bactris glaucescens has multiple stems and has bisexual flowers with crepuscular/nocturnal anthesis. Both species present bud break and leaf-budding during the year. Attalea phalerata shows continual annual flowering with ripening of fruits during the dry season. For B. glaucescens flowering occurs simultaneously with fruiting for about seven months, and fruit production can be influenced by temperature and level of flooding. The difference in the timing of anthesis of the flowers ensures production of floral resources during both day and night when both species of palms are flowering. The floral structures of both species have morphological characteristics associated with pollination by insects (mainly beetles), such as the presence of odor, stigmatic secretion, heat production, and adhesive substances ("pollenkitt") in the pollen. In A. phalerata, the main pollinators were Mystrops sp. (Nitidulidae) and Madarini (Curculionidae). Derelomus sp. (Curculionidae) and Paratenetus sp. (Tenebrionidae) beetles visited B. glaucescens during the day and could have pollinated the flowers at these occasions.
Introduction
The family Arecaceae comprises 440 genera and 3000 species with pantropical distribution and is represented in Brazil by 36 genera and 195 species (Giulietti et al., 2005). Information on phenology, pollination and/or reproductive system of Arecaceae are published in studies with individual species (e.g. Aguirre and Dirzo, 2008, Borchsenius, 2002, Inkrot et al., 2007, Listabarth, 2001, Martén and Quesada, 2001, Meekijjaroenroj and Anstett, 2003, Meléndez-Ramírez et al., 2004, Miller, 2002, Oliveira et al., 2003, Voeks, 2002), with two or three congeneric species or not (Adler and Lambert, 2008, Borchsenius, 1997, Ervik et al., 1999, Listabarth, 1996, Núñez et al., 2005) or in palm communities formed by seven to 27 species of the same (Listabarth, 1992) or different genera (De Steven et al., 1987, Genini et al., 2009, Henderson et al., 2000a).
Most studies have been done in an environment of humid tropical forest and/or plantations mainly in the Americas (Bolivia, Brazil, Colombia, Costa Rica, Ecuador, Mexico, Panama, Peru) and more rarely in countries on other continents (e.g. Australia – Inkrot et al., 2007, France – Meekijjaroenroj and Anstett, 2003). In Brazil, the studies are concentrated in the northern region, in the Amazon (Henderson et al., 2000a, Henderson et al., 2000b) and Pará (Oliveira et al., 2003), with few records from other regions or states (e.g. Bahia – Voeks, 2002).
Many Arecaceae species exhibit several characteristics associated with wind pollination syndrome (Faegri and Pijl, 1979) such as small and numerous flowers, light-colored, and abundant, smooth monosulcate pollen (Anderson et al., 1988). However, Henderson (1986) in a review on pollination biology in palm trees reported the occurrence of various types of pollination, but cantharophily, mellitophily, and myophily are the most common syndromes in the family. The more specialized relationship between palms and pollinators seem to be those involving beetles, particularly those belonging to the families Nitidulidae and Curculionidae (Henderson, 1986, Howard et al., 2001, Núñez et al., 2005, Voeks, 2002).
Attalea Kunth is one of the most important Neotropical genera of Arecaceae, occurring from Mexico to Bolivia, Paraguay, southern Brazil and the Caribbean. The number of species in the genus ranges from 27 to 67, according to the author considered, but there is consensus for validity of at least 20 species (Moraes, 2004, Pintaud et al., 2008). For Bactris Jacq. ex Scop there are at least 200 recognized species that are distributed from Mexico to Paraguay, with greater diversity in Brazil (Granville, 1992). Here, Attalea and Bactris are represented, respectively, by 13 and 23 species distributed throughout the national territory (Lorenzi et al., 1996). In the Brazilian Pantanal there is record (Pott and Pott, 1994, Pott and Pott, 1999) of at least two species of Attalea (A. phalerata Mart. ex Spreng. and A. speciosa Mart. ex Spreng.) and three species of Bactris (B. cf. cuyabensis Barb. Rodr., B. glaucescens Drude and B. aff. setosa Mart.).
In general, species of Attalea bloom and bear fruit throughout the year (Moraes et al., 1996, Reys et al., 2005, Voeks, 1988, Voeks, 2002), although these phenophases may vary depending on the season (Voeks, 2002) or according to the level of disturbance to the environment (Salm, 2005, Voeks, 1988). In this genus, monoecy predominates, and there is only occasional occurrence of androgyny (Anderson et al., 1988, Núñez et al., 2005). However, in monoecious species the male and female flowers are rarely produced simultaneously in the same individual, therefore being functionally dioecious (Núñez et al., 2005, Voeks, 1988, Voeks, 2002).
Monoecy also occurs in species of Bactris (Moraes, 2004), in which the flowering and fruiting period usually lasts 1–5 months (De Steven et al., 1987, Henderson et al., 2000a, Henderson et al., 2000b, Listabarth, 1996), with flowering and/or fruiting throughout the year being the rare case (e.g. B. major Jacq.: De Steven et al., 1987). In the species of Bactris the reproductive cycle usually occurs in the rainy season (De Steven et al., 1987, Henderson et al., 2000a, Henderson et al., 2000b, Listabarth, 1996), since prolonged periods of drought seem to negatively affect the development of some species (Chaimsohn et al., 2002, Moraes and Sarmiento, 1992). In regions with higher rainfall and without pronounced dry periods, flowering in this genus can occur up to twice a year (Ferreira, 2005).
Consistent data on pollination ecology of species of Attalea are available for A. allenii H. E. Moore (Núñez et al., 2005) and A. amygdalina Kunth (Blair et al., 2007) in Colombia and A. funifera Mart. in Brazil (Voeks, 2002). The main pollinators of these species are small Nitidulidae beetles (e.g. Mystrops spp.) and Curculionidae (Celetes bipunctata, Phyllotrox tatianae), although the inflorescences are visited by a wide variety of insects such as bees, wasps, ants, flies, bedbugs and cockroaches (Núñez et al., 2005, Voeks, 2002). Núñez et al. (2005) indicate pollination by nitidulid beetles of the genus Mystrops for A. attaleoides (Barb. Rodr.) Wess. Boer, A. microcarpa Mart. and A. speciosa. For Bactris the number of studies on pollination is higher, even compared to other genera of palms (Listabarth, 1996). Among them we can cite the studies of Listabarth, 1992, Listabarth, 1996 with three species of Bactris in the Peruvian Amazon rainforest and the community study of Henderson et al. (2000b) developed in the Brazilian Amazon forest with 10 sympatric species. In both studies, beetles of the genus Phyllotrox/Curculionidae were important pollinators followed by species of Epurea/Nitidulidae (Listabarth, 1992, Listabarth, 1996) or Colopterus (Henderson et al., 2000b). Pollination of Bactris by scarab beetles (Cyclocephala amazona, Scarabaeidae) in Costa Rica has also been recorded (Beach, 1984).
The palm trees are highly productive and in wetland forests are dominant in some environments, playing an important role as a source of organic matter due to their persistent and fibrous sheaths and the presence of dry leaves on their stems. Additionally, they provide food and shelter for local wildlife, such as arthropods, birds and rodents (Genini et al., 2009, Kahn and de Granville, 1995). In the Pantanal, the fruits of Attalea phalerata and Bactris glaucescens are important food resources for fish, birds, wild and domesticated mammals, as well as the local human populations (Pott and Pott, 1994). Since an understanding of phenology and pollination ecology is essential to furnish data for breeding programs, conservation and species management (Oliveira et al., 2003), we compared the phenology and pollination ecology of the palms Attalea phalerata and Bactris glaucescens occurring sympatrically in area of riparian vegetation in the Pantanal, Brazil.
Section snippets
Study area and studied species
The study lasted from August/2008 to October/2009 (except August/2009) in a strip of riparian forest adjacent to the Miranda River, near the Base for Studies of the Pantanal (BEP), of the Universidade Federal de Mato Grosso do Sul (19°34′ 36.4′′S and 57°01′ 07.8′′W), located in an area called "Passo do Lontra" sub-region of Miranda, in the Pantanal, Corumbá, Mato Grosso do Sul. The climate is Aw, hot and humid (Köppen, 1948), with two well-defined seasons: a rainy season from November to March
Morphology
Attalea phalerata has erect, staminate (Fig. 2A), pistillate (Fig. 2B) and bisexual inflorescences. Bisexual inflorescences can be of two types: (i) predominantly staminate with 3–5 fertile female flowers present at the base of some male rachillae (Fig. 2 C) or (ii) functionally pistillate, with 3–4 sterile male flowers that occur at the basis of some female flowers (Fig. 2B, detail). The number of staminate inflorescences per plant was similar to that of pistillate inflorescences, but the last
Morphology and biology of the inflorescence and flower
Attalea phalerata, although morphologically monoecious, is functionally dioecious, because the staminate and pistillate inflorescences do not occur simultaneously in the same individual, as also reported by Moraes et al. (1996) for this species in Bolivia. In several species of Attalea sexual alternation is common; sometimes they produce staminate inflorescences, sometimes pistillate and rarely occurs the simultaneous combination of the two sexes (Uhl and Dransfield, 1987). In the studied
Acknowledgments
To CAPES (Coordenação de Aperfeiçoamento de Pessoal de Nível Superior), for the scholarship awarded to the first author; the Graduate Program in Plant Biology, Federal University of Mato Grosso do Sul for the logistic support; Ayr de Moura Bello for the identification of beetles; Berinaldo Bueno, for help in statistical treatments; CEMTEC/AGRAER/INMET (Centro de Monitoramento de Tempo, do Clima e dos Recursos Hídricos de Mato Grosso do Sul/Agência de Desenvolvimento Agrário e Extensão
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