Characterization of unisexual flower development in the endangered mahogany tree Swietenia macrophylla King. (Meliaceae)

The selection of candidate plus trees of desirable phenotypes from tropical forest trees and the rapid devastation of the natural environments in which these trees are found have created the need for a more detailed knowledge of the floral and reproductive biology of tropical tree species. In this article, the organogenic processes related to unisexual flower development in tropical mahogany, Swietenia macrophylla , are described. Mahogany inflorescences at different developmental stages were evaluated using scanning electron microscopy or optical microscopy of histological sections. The unisexual flowers of S. macrophylla are usually formed in a thyrse, in which the positions of the female and male flowers are not random. Differences between male and female flowers arise late during development. Both female and male flowers can only be structurally distinguished after stage 9, where ovule primordia development is arrested in male flowers and microspore development is aborted in female flower anthers. After this stage, male and female flowers can be distinguished by the naked eye as a result of differences in the dimensions of the gynoecium. The floral characteristics of S. macrophylla (distribution of male and female flowers within the inflorescence, and the relative number of male to female flowers) have practical implications for conservation strategies of this endangered species.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 156 , 529–535.     

In vitro rhizogenesis: histoanatomy of Cedrela odorata (Meliaceae) microcuttings

Cedrela odorata (Meliaceae) is considered as one of the most valuable forest tree in the tropics. Clonal propagation of this species provide an alternative method to propagate superior genotypes, being the production of good quality adventitious roots one of the most important steps in micropropagation techniques. The sequence of anatomical changes that takes place during the formation of adventitious roots in shoots of Cedrela odorata cultured in vitro is described in this study. Eigth-week-old shoots, from multiplication cultures, were rooted in Murashige and Skoog's medium (1962) with half-strength macronutrients and with 0 or 1 mg/l indole-3-butyric acid (IBA). Between 12 and 24h after the start of rooting, some cambium, phloem and interfascicular parenchyma cells became dense cytoplasm, nuclei with prominent nucleoli and the first cell divisions were observed, especially in shoots treated with auxin (dedifferentiation phase). After 3-4 days, the number of dedifferentiated cells and mitotic divisions increased considerably, and the formation of groups of some 30-40 meristematic cells (meristemoids) was observed (induction phase). The first primordial roots developed from the 4th-5th day. The vascular tissues of these primordia connected to those of the explant, and roots began to emerge from the base by day 6. Development of the primordial roots was similar in the control shoots and shoots treated with 1 mg/l IBA, although there were more roots per explant in the latter.     

Imbibition of Swietenia macrophylla (Meliaceae) seeds: the role of stomata

BACKGROUND AND AIMS: The occurrence of stomata in seed coats is uncommon and there is limited information about their function(s). The aim of this study was to verify the distribution of stomata in seed coats of Swietenia macrophylla and to relate it to the imbibition process and aspects of the structure of the outer integument layers. METHODS: For the structural and ultrastructural studies, the seeds were processed using the usual techniques and studied under light and scanning electron microscopes. Histochemical tests were employed to identify the cell wall composition in the different seed coat portions. To assess the role of the stomata in the imbibition, non-impervious seeds were compared with partially impervious ones, in which only the embryo, median or hilar regions were left free. Further, the apoplastic pathway marker was employed to confirm the role of the stomata as sites of water passage during imbibition. KEY RESULTS: A positive relationship was observed between seed coat thickness and stomata density. The stomata were devoid of movement, with a large pore. They occurred in large numbers in the embryo region and extended with lower frequency towards the wing. Imbibition rates were related to stomata density, suggesting that the stomata act as preferential sites for water entry in the S. macrophylla seeds. CONCLUSIONS: At maturity, the stomata in the seed coat play a significant role in seed imbibition. The data may also infer that these permanently opened stomata have an important role in gas exchange during seed development, aiding embryo respiration.     

Population genetic structure of mahogany (Swietenia macrophylla King, Meliaceae) across the Brazilian Amazon, based on variation at microsatellite loci: implications for conservation

Mahogany (Swietenia macrophylla, Meliaceae) is the most valuable and intensively exploited Neotropical tree. No information is available regarding the genetic structure of mahogany in South America, yet the region harbours most of the unlogged populations of this prized hardwood. Here we report on the genetic diversity within and the differentiation among seven natural populations separated by up to 2100 km along the southern arc of the Brazilian Amazon basin. We analysed the variation at eight microsatellite loci for 194 adult individuals. All loci were highly variable, with the number of alleles per locus ranging from 13 to 27 (mean = 18.4). High levels of genetic diversity were found for all populations at the eight loci (mean HE = 0.781, range 0.754-0.812). We found moderate but statistically significant genetic differentiation among populations considering both estimators of FST and RST, theta = 0.097 and rho = 0.147, respectively. Estimates of theta and rho were significantly greater than zero for all pairwise population comparisons. Pairwise rho-values were positively and significantly correlated with geographical distance under the isolation-by-distance model. Furthermore, four of the populations exhibited a significant inbreeding coefficient. The finding of local differentiation among Amazonian mahogany populations underscores the need for in situ conservation of multiple populations of S. macrophylla across its distribution in the Brazilian Amazon. In addition, the occurrence of microgeographical genetic differentiation at a local scale indicates the importance of maintaining populations in their diverse habitats, especially in areas with mosaics of topography and soil.     

A combination of molecular markers identifies evolutionarily significant units in Cedrela odorata L. (Meliaceae) in Costa Rica

The necessity for conservation of the geneticcomponent of biodiversity is now widelyrecognised. A broad genetic base is required tomaintain evolutionary potential and thepopulation erosion occurring in much of theworld's forests threatens the genetic integrityof many tree species. Spanish Cedar (Cedrela odorata L.) has been under severepressure for generations and is now the focusof a study aimed at assessing the levels anddistribution of genetic diversity in remainingpopulations. Ten Costa Rican populations wereanalysed using chloroplast and AFLP markers.The overall level of diversity was as expectedfor an outcrossing, long-lived, woody species(H _T = 0.27). However, this concealeda deep divergence within the species, forchloroplast and AFLP (_CT = 0.83)markers. Populations were differentiated in twogroups that exhibited contrasting habitatpreferences and two ecotypes, wet and dry, wereidentified. Within the ecotypes, all but onepopulation were fixed for a single chloroplasthaplotype and within populations, total genomicdiversity levels were low (H _S= 0.03–0.13). Populations possessing the dryecotype maintained significantly more diversitythan those from wet regions. Within the wetecotype group, pairwise genetic distancebetween populations fitted an isolation bydistance model. The group was stronglysubdivided and showed isolation by distancearound the southern edge of the centralmountain ranges. The genetic divergence of thetwo ecotypes, observed at both organellar andnuclear loci, identifies evolutionarilysignificant units that, taken together withprevious studies of the species, provide arational basis on which to build a conservationpolicy for the species.     

Chloroplast DNA Microsatellites Reveal Contrasting Phylogeographic Structure in Mahogany (Swietenia macrophylla King, Meliaceae) from Amazonia and Central America

Big-leaf mahogany (Swietenia macrophylla King) is one of the most valuable and overharvested timber trees of tropical America. In order to better characterize geographic patterns of genetic variation, we performed a phylogeographic analysis of S. macrophylla based on six polymorphic chloroplast genome simple sequence repeat loci (cpSSRs) analyzed in 16 populations (N?=?245 individuals) distributed across Central America and the Brazilian Amazon. Of the 31 total cpDNA haplotypes identified, 16 occurred in Central America and 15 in Amazonia with no single haplotype shared between the two regions. Populations from Central America showed moderate differentiation (F _ST ?=?0.36) while within population genetic diversity was generally high (mean Nei’s H _E ?=?0.639). In contrast, the Amazonian populations were strongly differentiated (F _ST ?=?0.91) and contained relatively low genetic diversity (mean H _E ?=?0.176), except for one highly diverse population (H _E ?=?0.925) from eastern Amazonia. Spatial analysis of molecular variance (SAMOVA) identified a single Central American phylogroup and four Amazonian phylogroups, indicating stronger phylogeographic structure within Amazonia. The results demonstrate distinctive regional patterns of S. macrophylla differentiation, and the first evidence of a strong phylogeographic break between Central American and South American mahogany populations. We suggest that the frequent occurrence of hurricanes in Central America, the differences in the glacial histories and in the duration and intensity of anthropogenic disturbance during the late Holocene may have played important roles in the geographic structuring of cpDNA lineages in the two regions. The high private haplotype diversity in Brazilian populations suggests that cpSSRs can be used as DNA barcodes for regional timber certification.     

The tree Cedrela odorata (Meliaceae): a morphologically subdivided species in Costa Rica

In Latin America. Cedrela odorata is a wide ranging species that occurs in several environments, where it shows significant morphological variation. A common garden experiment was established with seedlings from 63 families of ten populations from two habitat types (mesic and dry), distributed throughout Costa Rica, to examine the relationships between quantitative variation and site of population origin. Seedlings from dry areas tended to be distinct from those from mesic areas, with climatic grouping of provenance explaining a mean of 52% of the total variance and 80% of the genetic variance. Cluster analysis for seedling traits showed two natural groupings of families, which corresponded for the most part with the regional population groupings into mesic (Atlantic and South Pacific) and dry (North Pacific) groups. Cluster analysis based on seed weight and size also separated populations into mesic and dry climatic groups. Seeds from populations in dry areas were 43% heavier, and seedlings were 61% taller, 117% greater in diameter, and with leaflets 39% longer and 81% wider. These differences may be related to fast growth in the dry zone for taking advantage of early life cycle moisture availability. These findings may indicate incipient speciation in C. odorata in Costa Rica. Evaluation of reproductive isolating mechanisms between populations from the mesic and dry zones, and of clines at potential zones of hybridization would assist in testing a speciation hypothesis.     

CITES and the sustainable management of Swietenia macrophylla King

The merits and applicability of CITES to the management of mahogany ( Swietenia macrophylla King) are assessed. Emphasis is placed on the necessity for trade regulation in order to encourage the sustainable management of S. macrophylla , including the protection of adequate population and genetic resources throughout its range.     

Anatomy,ultrastructure, and secretory activity of the floral nectaries in Swietenia macrophylla (Meliaceae)

• Premise of the study: While mahogany (Swietenia macrophylla) is one of the most important forest species in the Amazon region, little is known about its reproductive biology. Knowledge about the nectary structure and dynamics of nectar production of this species represent a key step toward understanding its relationship with pollinators. • Methods: Mahogany tree floral buds and flowers in anthesis were collected, fixed, and processed for study by light and transmission and scanning electron microscopy. The chemical composition of nectar and the nectary pigments was also studied. • Key results: Both staminate and pistillate flowers have nectaries, which contain a papillose epidermis and stomata. The nectariferous tissue is parenchymatous, with the cell cytoplasm primarily containing mitochondria and plastids. Secretory activity initiates at the beginning of anthesis, which occurs at nightfall. Flowers undergoing anthesis become structurally modified, with starch grains in the plastids disappearing. The number of plastoglobuli in the plastids also increases when nectaries change color from pale yellow to intense red. Pistillate and staminate flowers produce meager nectar rewards. • Conclusions: Changes in plastoglobuli number seem to be related to an increase in carotenes and color changes during anthesis. Carotenes can be linked to the protection of the plant against oxidative stress, which results from secretory activities. Nectary color has a limited role as a pollinator attractant. Floral rewards comprise small nectar droplets in both flower types, in addition to a few pollen grains in staminate flowers. These meager rewards are probably adapted to attract small generalist insects.     

Multiplexed systems of microsatellite markers for genetic analysis of mahogany,Swietenia macrophylla King (Meliaceae), a threatened neotropical timber species

Mahogany (Swietenia macrophylla King [Meliaceae]) is the most valuable hardwood species in the neotropics. Its conservation status has been the subject of increasing concern due to overexploitation and habitat destruction. In this work we report the development and characterization of 10 highly variable microsatellite loci for S. macrophylla. Twenty-nine percent of the 126 sequenced mahogany clones yielded useful microsatellite loci. Three high-throughput genotyping systems were developed based on polymerase chain reaction (PCR) multiplexing of these mahogany loci. We identified a total of 158 alleles in 121 adult individuals of S. macrophylla, with an average of 15.8 alleles (range 11-25) per locus. All loci showed Mendelian inheritance in open-pollinated half-sib families. The mean expected heterozygosity was 0.84 and the mean observed heterozygosity was 0.73. The combined probability of identity-the probability that two individuals selected at random from a population would have identical genotypes--was 7.0 x 10(-15), and combined probability of paternity exclusion was 0.999998 overall loci. These microsatellite loci permit precise estimates of parameters such as gene flow, mating system, and paternity, thus providing important insights into the population genetics and conservation of S. macrophylla.     

Restoration of genetic diversity in the dry forest tree Swietenia macrophylla (Meliaceae) after pasture abandonment in Costa Rica

We studied the levels of genetic diversity of Swietenia macrophylla (big leaf mahogany) in five successional plots in the Santa Rosa National Park, Guanacaste, Costa Rica. We selected sites with different lengths of time since the last major disturbance (typically fire): 6, 9, 15 and 20 years. In addition, we also included a patch of mature forest that had experienced selective logging and other human activity in the past 100 years. Genetic diversity was assessed using five polymorphic DNA microsatellite loci. We found a total of 21 alleles in the five loci examined, in which the number of alleles present varied among the five sites studied. Allelic diversity varied between sites ranging from 20 to 14 alleles, and our data revealed that earlier successional sites have more alleles than older sites. There was significant heterogeneity in allele frequencies between sites; however, genetic differentiation between populations was low (FST = 0.063) indicating that most of the variation was found within sites and extensive gene flow between sites. In addition, our analysis also showed that genetic diversity of adult trees does not solely determine the diversity of seedlings and saplings found around them, also supporting the existence of extensive gene flow. The impact of these findings for the design of conservation strategies for tropical dry forests trees is discussed.     

Ecology and management of mahogany (Swietenia macrophylla King) in the Chimanes Forest, Bed, Bolivia

Mahogany ( Swietenia macrophylla King) regenerates in areas of erosion on high terraces and in forest killed by flooding and deposition of alluvial sediments in the Chimanes Forest, Bolivia. These hydrological disturbances are patchy, and only one of five stands of mahogany that we inventoried was regenerating. Mahogany survives these disturbances significantly better than the common tree species. The long time between disturbances appears to favour late maturation. Mahogany trees allocate little photosynthates to reproduction until they are very large emergents, at least 80 cm in diameter. The episodic nature of the regeneration sites means that mahogany stands are composed of one or a few cohorts, which are vulnerable to overharvesting, particularly with the current use of a minimum cutting diameter to regulate harvest. The delayed onset of fecundity means that the small trees that escape harvest are not very fecund, resulting in minimal seed input to logged forest. Only 7–9% of the gaps created by logging contain natural regeneration after 20 + yr. A successful management plan for mahogany would entail a monocyclic harvest, with a rotation age of 100 + years, the estimated time that it takes for trees to achieve commercial size in natural forest. Since the number of seed trees that will be left is small, they should be concentrated in sites that are likely to be conducive to natural regeneration, such as near rivers and flood damaged forest. Seed production will be maximized for a given basal area (opportunity cost to loggers) if trees c. 110 cm dbh are selected as seed trees. The mahogany stocks in the Chimanes Forest are nearly exhausted, but the findings of this study could be used to help rebuild the mahogany populations, or to design management plans for the commercial species that have similar ecologies to mahogany.     

Systemic activity of plant extracts in Cedrela odorata (Meliaceae) seedlings and their biological activity on Hypsipyla grandella (Lepidoptera: Pyralidae) larvae

1 Deterrent and toxic substances known to affect Hypsipyla grandella larvae were bioassayed to determine their possible systemic activity under laboratory conditions in Turrialba, Costa Rica. 2 Leaflets from Spanish cedar Cedrela odorata seedlings grown in vitro on culture media treated with different plant extracts were excised from the seedlings and exposed to H. grandella larvae. 3 Both Azatin EC (derived from neem, Azadirachta indica) and plant extracts (from Quassia amara wood, Ruta chalepensis leaves and Sechium pittieri fruits) were able to translocate within Spanish cedar seedlings and express their biological activity against H. grandella larvae. 4 Rates of leaflet consumption were nil for seedlings treated with the commercial standard (carbofuran) and Azatin EC, both causing larval death. 5 Intermediate levels of consumption were reached with the three plant extracts, all of them causing deterrence, because they precluded feeding activity by larvae when exposed to treated leaflets.     

中国分类学文献中Swietenia mahagoni之订正

通过调查,对Swietenia mahagoni的后选模式、S.macrophylla的主模式与中国分类学文献(如《植物分类学报》、《广州植物志》、《中国高等植物图鉴》、《云南植物志》、《广东植物志》、《中国植物志》和《中国高等植物》)的S.mahagoni作比较,证实中国分类学文献将S.mahagoni和S.macrophylla相混,上述分类学文献中的S.mahagon是S.macrophylla King。桃花心木(S.mahagoni(L.)Jacq.)的羽状复叶及小叶较短,小叶长度绝不超过10 cm,果卵球形,长度绝不超过15 cm;而大叶桃花心木(S.macrophylla King)的羽状复叶及小叶较长,总有一部分小叶的长度超过10 cm,果长卵球形,长12~22 cm。     

Light Availability Influences Growth‐Defense Trade‐Offs in Big‐Leaf Mahogany (Swietenia macrophylla King)

We report on findings from a common garden experiment showing that the magnitude of change in the production of phenolic compounds in response to different amounts of light availability is tissue‐specific in saplings of big‐leaf mahogany (Swietenia macrophylla). Moreover, we show that trade‐offs between growth and the production of chemical defenses emerge only under light‐limited conditions. These findings emphasize the need for considering the specificity of plant defensive responses to resource availability, as well as the influence of the abiotic environment (via trade‐offs) on plant defense allocation patterns.     

Floral development in the tribe Cedreleae (Meliaceae, sub-family Swietenioideae): Cedrela and Toona

BACKGROUND AND AIMS: Floral development of Cedrela and Toona, the genera comprising the basal tribe Cedreleae of the sub-family Swietenioideae of Meliaceae, is described. The focus was on three endangered, ecologically and economically important species: Cedrela fissilis, Cedrela odorata and Toona ciliata. The aims of the study were to characterize the patterns of floral development in the tribe and to establish apomorphic and plesiomorphic floral characters in relation to other taxa within the family based on the current molecular phylogeny of Meliaceae. METHODS: A detailed floral structural and developmental study was completed using both scanning electron microscopy and visualization of microtome sections with a light microscope. KEY RESULTS: Twelve floral developmental stages were identified. The initial development of the pentamerous flowers of both Toona and Cedrela is strikingly similar. The morphological differences observed between them are due to differential patterns of organ elongation and adnation/connation occurring late in development. Additionally, the formation of functionally male and female flowers was found to occur at specific positions within the inflorescence. CONCLUSIONS: Due to the basal position of the tribe Cedreleae in the phylogeny of Meliaceae, functionally either male or female pentamerous flowers and the presence of (at least partially) free stamens may be considered plesiomorphic traits within the family. In contrast, sympetaly and the absence of nectaries in Cedrela species are synapomorphies.     

Catastrophic disturbance, logging and the ecology of mahogany (Swietenia macrophylla King): grounds for listing a major tropical timber species in CITES

Recent debate on whether or not mahogany ( Swietenia macrophylla King) is threatened by the international timber trade has focused on the breadth of its range and estimates of the remaining stock of mahogany trees. These data are inadequate to reveal the status of mahogany populations, both because they are incomplete in areal extent and because they do not reveal population parameters such as the existence or density of young trees smaller than commercial size. However, there is sufficient information on the regeneration ecology of mahogany to indicate that under natural conditions this species regenerates in essentially even-aged stands after catastrophic disturbances destroy many or most trees, and, in the case of fires and flooding, saplings and seedlings as well. Adult mahoganies tend to survive these events, and regenerate by shedding seed onto the resulting gaps or clearings. This ecological strategy makes mahogany vulnerable to logging, first because juvenile mahoganies are not found in the understorey, and secondly because logging operations shortcircuit mahogany regeneration processes by selectively removing almost all mahogany seed sources while leaving standing competing vegetation of other species. Listing of mahogany in CITES Appendix II could provide both a mechanism to fill in gaps in information and an incentive to change current practices in favour of silvicultural management to provide for regeneration of this valuable timber species in forests subjected to logging.     

Teleconnections and edaphoclimatic effects on tree growth of Cedrela odorata L. in a seasonally dry tropical forest in Brazil

The Seasonally Dry Tropical Forests (SDTF) present very high biodiversity and a number of tree species that are adapted to prolonged periods of water stress. Considering tree ring formation is mainly driven by seasonal variation in precipitation in tropical environments, tree-ring studies from STDF can provide important contributions to understanding how these forests are responding to climate variations. In the present study, we demonstrate the influence of edaphoclimatic variables (precipitation, air temperature and soil water deficit-SWD) and the ocean teleconnections (Tropical Southern Atlantic-TSA, Atlantic Multidecadal Oscillation-AMO, Western Hemisphere Warm Pool-WHWP and El Niño 3.4) on Cedrela odorata L. growth from a SDTF of northeastern Brazil. We used standard dendrochronological methods to develop an 89-year-long ring-width index chronology. The climate sensitivity of C. odorata was assessed through Pearson's correlation tests and linear regressions, which allowed to identify the determinant months (cause-effect) of each variable on the chronology. Tree growth was positively correlated with precipitation and negatively correlated with air temperature and SWD, particularly during the rainy season (March to August). In parallel, we identified that extremely dry years can contribute to missing rings, exposing the lack of growth in C. odorata caused by water stress. Among the oceanic variables, all of them showed a negative effect on radial growth of C. odorata, except for TSA, which had no significant effect. Tree growth is constrained in years with strong El Niño and high values of AMO index during the rainy months (May, June and October). However, the WHWP showed a more pronounced negative effect in the beginning of the dry season (September). Our findings add valuable information on C. odorata responses to hydrological seasonality from SDTF and the fluctuations in oceanic teleconnections, which in turn, influence the rainfall dynamics in northeastern Brazil.