GEOGRAPHICAL GENETIC STRUCTURING AND PHENOTYPIC VARIATION IN THE VELLOZIA HIRSUTA (VELLOZIACEAE) OCHLOSPECIES COMPLEX

Premise of the study: Vellozia hirsuta forms a complex presenting wide morphological and anatomical variation, resulting in five specific names and 14 morpho-anatomical patterns occurring in disjunct populations. We carried out a phylogeographical study to investigate the existence of correlation among the genetic and morphological patterns within this complex, and to determine whether it is composed of various species or should be treated as an ochlospecies, a species having widely polymorphic and weakly polytypic complex variation, with morphological characteristics varying independently. Methods: We carried out phylogeographical analyses using cpDNA rpl32F-trnL intergenic region. Key results: We found 20 haplotypes in 23 populations sampled. The populations are genetically structured (Phi(ST) = 0.818) into four phylogeographical groups demonstrating geographical structuring but with no correlation with morpho-anatomical patterns. Our analyses do not support recognizing any of the species now synonymized under Vellozia hirsuta. The northern populations were the most genetically differentiated and could be considered a distinct taxon, as they are also morphologically different. Conclusions: It is recommended that Vellozia hirsuta be considered a single enormously variable species. The patterns of variation within V. hirsuta probably are related to climatic changes that occurred during the Pleistocene Epoch in tropical Brazil when reductions in forest cover favored the expansion of V. hirsuta populations into extensive lowland areas. The expansion of forest cover at the end of the glaciations would have again restricted the occurrence of campos rupestres vegetation to high elevations, which constitute the current centers of diversity of this species.

Oleoresin glands in copaiba (Copaifera trapezifolia Hayne: Leguminosae), a Brazilian rainforest tree

Although studies have addressed the chemical analysis and the biological activity of oleoresin in species of Copaifera, the cellular mechanisms of oleoresin production, storage, and release have rarely been investigated. This study detailed the distribution, ontogeny, and ultrastructure of secretory cavities and canals distributed in leaf and stem, respectively, of Copaifera trapezifolia, a Brazilian species included in a plant group of great economic interest. Axillary vegetative buds, leaflets, and portions of stem in primary and secondary growth were collected and processed in order to study the anatomy, histolocalization of substances, and ultrastructure. Secretory cavities are observed in the foliar blade and secretory canals in the petiolule and stem. They are made up of a uniseriate epithelium delimiting an isodiametric or elongated lumen. Biseriate epithelium is rarely observed and is a novelty for Leguminosae. Cavities and canals originate from ground meristem cells and the lumen is formed by schizogenesis. The content of the cavities and canals of both stem and leaf is oily and resinous, which suggests that the oleoresin could be extracted from the leaf instead of the stem. Phenolic compounds are also detected in the epithelial cell cytoplasm. Cavities and canals in the beginning of developmental stages have polarized epithelial cells. The cytoplasm is rich in smooth and rough endoplasmic reticula connected to vesicles or plastids. Smooth and rough endoplasmic reticulum and plastids were found to be predominant in the epithelial cells of the secretory cavities and canals of C. trapezifolia. Such features testify the quantities of oleoresin found in the lumen and phenolic compounds in the epithelial cell cytoplasm of these glands. Other studies employing techniques such as correlative light electron microscopy could show the vesicle traffic and the compartmentalization of the produced substances in such glands.

Structure and inferred dynamics of a large grove of Microberlinia bisulcata trees in central African rain forest: the possible role of periods of multiple disturbance events

A 272-ha grove of dominant Microberlinia bisulcata (Caesalpinioideae) adult trees greater than or equal to 50 cm stem diameter was mapped in its entirety in the southern part of Korup National Park, Cameroon. The approach used an earlier-established 82.5-ha permanent plot with a new surrounding 50-m grid of transect lines. Tree diameters were available from the plot but trees on the grid were recorded as being greater than or equal to 50 cm. The grove consisted of 1028 trees in 2000. Other species occurred within the grove. including the associated subdominants Tetraberlinia bifoliolata and T. korupensis. Microberlinia bisulcata becomes adult at a stein diameter of c. 50 cm and at an estimated age of 50 y. Three oval-shaped subgroves with dimensions c. 8 50 in x 13 50 in (90 ha) were defined. For two of them (within the plot) tree diameters were available. Subgroves differed in their scales and intensities of spatial tree patterns, and in their size frequency distributions, these suggesting differing past dynamics. The modal scale of clumping was 40-50 m. Seed dispersal by pod ejection (to c. 50 in) was evident from the semi-circles of trees at the grove's edge and from the many internal circles (100-200 m diameter). The grove has the capacity. therefore, to increase at c. 100 m per century. To form its present extent and structure. it is inferred that it expanded and infilled from a possibly smaller area of lower adult-tree density. This possibly happened in three waves of recruitment, each one determined by a period of several intense disturbances. Climate records for Africa show that 1740-50 and 1820-30 were periods of drought, and that 1870-1895 was also regionally very dry. Canopy openings allow the light-demanding and fast-growing ectomycorrhizal M. bisulcata to establish, but successive releases are thought to be required to achieve effective recruitment. Nevertheless, in the last 50 y there were no major events and recruitment in the grove was very poor. This present study leads to a new hypothesis of the role of periods of multiple extreme events being the driving factor for the population dynamics of many large African tree species such as M. bisulcata.

Tree size and fecundity influence ballistic seed dispersal of two dominant mast-fruiting species in a tropical rain forest

Seed production, seed dispersal, and seedling recruitment are integral to forest dynamics, especially in masting species. Often these are studied separately, yet scarcely ever for species with ballistic dispersal even though this mode of dispersal is common in legume trees of tropical African rain forests. Here, we studied two dominant main-canopy tree species, Microberlinia bisulcata and Tetraberlinia bifoliolata (Caesalpinioideae), in 25 ha of primary rain forest at Korup, Cameroon, during two successive masting events (2007/2010). In the vicinity of c. 100 and 130 trees of each species, 476/580 traps caught dispersed seeds and beneath their crowns c. 57,000 pod valves per species were inspected to estimate tree-level fecundity. Seed production of trees increased non-linearly and asymptotically with increasing stem diameters. It was unequal within the two species’ populations, and differed strongly between years to foster both spatial and temporal patchiness in seed rain. The M. bisulcata trees could begin seeding at 42–44 cm diameter: at a much larger size than could T. bifoliolata (25 cm). Nevertheless, per capita life-time reproductive capacity was c. five times greater in M. bisulcata than T. bifoliolata owing to former’s larger adult stature, lower mortality rate (despite a shorter life-time) and smaller seed mass. The two species displayed strong differences in their dispersal capabilities. Inverse modelling (IM) revealed that dispersal of M. bisulcata was best described by a lognormal kernel. Most seeds landed at 10–15 m from stems, with 1% of them going beyond 80 m (<100 m). The direct estimates of fecundity significantly improved the models fitted. The lognormal also described well the seedling recruitment distribution of this species in 121 ground plots. By contrast, the lower intensity of masting and more limited dispersal of the heavier-seeded T. bifoliolata prevented reliable IM. For this species, seed density as function of distance to traps suggested a maximum dispersal distance of 40–50 m, and a correspondingly more aggregated seedling recruitment pattern ensued than for M. bisulcata. From this integrated field study, we conclude that the reproductive traits of M. bisulcata give it a considerable advantage over T. bifoliolata by better dispersing more seeds per capita to reach more suitable establishment sites, and combined with other key traits they explain its local dominance in the forest. Understanding the linkages between size at onset of maturity, individual fecundity, and dispersal capability can better inform the life-history strategies, and hence management, of co-occurring tree species in tropical forests.

Palynological record and radiocarbon dates for sediment core T89-16, Congo River mouth

We present a high-resolution reconstruction of tropical palaeoenvironmental changes for the last deglacial transition (18 to 9 cal. kyr BP) based on integrated oceanic and terrestrial proxies from a Congo fan core. Pollen, grass cuticle, Pediastrum and dinoflagellate cyst fluxes, sedimentation rates and planktonic foraminiferal d18O ratios, uK37 sea-surface temperature and alkane/alkenone ratio data highlight a series of abrupt changes in Congo River palaeodischarge. A major discharge pulse is registered at around 13.0 cal. kyr BP which we attribute to latitudinal migration of the Intertropical Convergence Zone (ITCZ) during deglaciation. The data indicate abrupt and short-lived changes in the equatorial precipitation regime within a system of monsoonal dynamics forced by precessional cycles. The phases of enhanced Congo discharge stimulated river-induced upwelling and enhanced productivity in the adjacent ocean.

Palynology of marine sediment cores from the West African coast

Seven sediment cores from the cruises of the "Meteor" and "Valdivia" were examined palynologically. The cores were retrieved from the lower continental slope in the area of between 33.5° N and 8° N, off the West African coast. Most of the cores contain sediments from the last Glacial and Interglacial period. In some cases, the Holocene sediments are missing. Some individual cores contain sediments also from earlier Glacial and Interglacial periods. The main reason for making this palynological study was to find out the differences between the vegetation of Glacial and Interglacial periods in those parts of West Africa which at present belong to the Mediterranean zone, the Sahara and the zones of the savannas and tropical forests. In today's Mediterranean vegetation zone at core 33.5° N, forests and deciduous forests in particular, are missing during Glacial conditions. Semi-deserts are found instead of these. In the early isotope stage 1, there is a very significant development of forests which contain evergreen oaks; this is the Mediterranean type of vegestation development. The Sahara type of vegetation development is shown in four cores from between 27° N and 19° N. The differences between Glacial and Interglacial periods are very small. It must be assumed therefore that in this latitudes, both Glacial and Interglacial conditions gave rise to desert generally. The results are in favour of a slightly more arid climate during Glacial and more humid one during Interglacial periods. The southern boundary of the Sahara and the adjacent savannas with grassland and tropical woods were situated more to the south during the Glacial periods than they were during the Interglacial ones. In front of today's savanna belt, it can be seen from the palynological results that there are considerable differences between the vegetation of Glacial and Interglacial periods. The woods are more important in Interglacial periods. During the Glacial periods these are replaced from north to south decreasingly by grassland (savanna and rainforest type of vegetation development). The southern limit of the Sahara during stage 2 was somewhat between 12° N and 8° N which is between 1.5 and 5 degrees in latitude further south than it i s today. Not only do these differences in climate and vegetation apply to the maximum of the last Glacial and for the Holocene, but they apparently apply also to the older Glacial and Interglacial periods, where they have been found in the profiles. The North African deset belt can be said to have expanded during Glacial times both towards the north and towards the south. All the available evidence of this study indicates that the grass land or the semi-desert of the Southern Europe cam einto connection with those of the N Africa; there could not have been any forest zone between them. The present study was also a good opportunity for investigating some of the basic marine palynological problems. The very well known overrepresentation of pollen grains of the genus Pinus in marine sediments can be traced as fa as 21° N. The present southern limit for the genus Pinus is on the Canaries and on the African continent as approximately 31° N. Highest values of Ephedra pollen grains even occur south of the main area of the present distribution of that genus. These does not seem to be any satisfactory explanation for this. In general, it would appear that the transport of pollen grains from the north is more important than transport from the south. The results so far, indicate strongly that further palynological studies are necessary. These should concentrate particularly on cores from between 33° N and 27° N as well as between 17° N and 10° N. It would also be useful to have a more detailed examination of sediments from the last Intergalcial period (substage 5 e). Absolute pollen counts and more general examination of surface samples would be desirable. Surface samples should be taken from the shelf down to the bottom of the continental slope in different latitudes.

Pollen distribution in marine sediment samples along the south-western African coast

The distribution of pollen in marine sediments is used to reconstruct pathways of terrigenous input to the oceans and provides a record of vegetation change on adjacent continents. The wind transport routes of aeolian pollen is comprehensively illustrated by clusters of trajectories. Isobaric, 4-day backward trajectories are calculated using the modelled wind-field of ECHAM3, and are clustered on a seasonal basis to estimate the main pathways of aeolian particles to sites of marine cores in the south-eastern Atlantic. Trajectories and clusters based on the modelled wind-field of the Last Glacial Maximum hardly differ from those of the present-day. Trajectory clusters show three regional, and two seasonal patterns, determining the pathways of aeolian pollen transport into the south-eastern Atlantic ocean. Mainly, transport out of the continent occurs during austral fall and winter, when easterly and south-easterly winds prevail. South of 25°S, winds blow mostly from the west and southwest, and aeolian terrestrial input is very low. Generally, a good latitudinal correspondence exists between the distribution patterns of pollen in marine surface sediments and the occurrence of the source plants on the adjacent continent. The northern Angola Basin receives pollen and spores from the Congolian and Zambezian forests mainly through river discharge. The Zambezian vegetation zone is the main source area for wind-blown pollen in sediments of the Angola Basin, while the semi-desert and desert areas are the main sources for pollen in sediments of the Walvis Basin and on the Walvis Ridge. A transect of six marine pollen records along the south-western African coast indicates considerable changes in the vegetation of southern Africa between glacial and interglacial periods. Important changes in the vegetation are the decline of forests in equatorial Africa and the north of southern Africa and a northward shift of winter rain vegetation along the western escarpment.

Pollen analysis and age model of sediment core GIK-15856-2

Pollen and spores from a deep-sea core located west of the Niger Delta record an uninterrupted area of lowland rain forest in West Africa from Guinea to Cameroon during the last Interglacial and the early Holocene. During other periods of the last 150 ka, a savanna corridor between the western - Guinean - and the eastern - Congolian - part of the African lowland rain forest existed. This so-called Dahomey Gap had its largest extension during Glacial Stages 6, 4, 3, and 2. Reduced surface salinity in the eastern Gulf of Guinea as recorded by dinoflagellate cysts indicates sufficient precipitation for extensive forest growth during Stages 5 and 1. The large modern extension of dry forest and savanna in West Africa cannot be solely explained by climatic factors. Mangrove expansion in and west of the Niger Delta was largest during the phases of sea-level rise of Stages 5 and 1. During Stages 6, 4, 3, and 2, shelf areas were exposed and the area of the mangrove swamps was minimal.

Pollen record and dating of sediment core GIK16776-1 off Liberia

Based on pollen analysis of a sediment core from the Atlantic Ocean off Liberia the West African vegetation history for the last 400 ka is reconstructed. During the cold oxygen isotope stages 12, 10, 8, 6, 4, 3 and 2 an arid climate is indicated, resulting in a southward shifting of the southern border of the savanna. Late Pleistocene glacial stages were more arid than during the Middle Pleistocene. A persistence of the rain forest in the area, even during the glacial stages, is recorded. This suggests a glacial refuge of rain forest situated in the Guinean mountains. Afromontane forests with Podocarpus occurred in the Guinean mountains from the stages 12 to 2 and disappeared after. The tree expanded from higher to lower elevations twice in the warm oxygen isotope stage 11 (pollen subzones 11d, 11b) and at least twice during the warm stage 5 (pollen subzones 5d, 5a), indicating a relative cool but humid climate for these periods.

Pollen analysis of surface sediment off northwest Africa

Over 100 samples of recent surface sediments from the bottomn of the Atlantic Ocean offshore NW Africa between 34° and 6° N have been analysed palynologically. The objective of this study was to reveal the relation between source areas, transport systems, and resulting distribution patterns of pollen and spores in marine sediments off NW Africa, in order to lay a sound foundation for the interpretation of pollen records of marine cores from this area. The clear zonation of the NW-African vegetation (due to the distinct climatic gradient) is helpful in determining main source areas, and the presence of some major wind belts facilitates the registration of the average course of wind trajectories. The present circulation pattern is driven by the intertropical front (ITCZ) which shifts over the continent between c. 22° N (summer position) and c. 4° N (winter position) in the course of the year. Determination of the period of main pollen release and the average atmospheric circulation pattern effective at that time of the years is of prime importance. The distribution patterns in recent marine sediments of pollen of a series of genera and families appear to record climatological/ecological variables, such as the trajectory of the NE trade, January trades, African Easterly Jet (Saharan Air Layer), the northernmost and southernmost position of the intertropical convergence zone, and the extent and latitudinal situation of the NW-African vegetation belt. Pollen analysis of a series of dated deep-sea cores taken between c. 35° and the equator off NW African enable the construction of paleo-distribution maps for time slices of the past, forming a register of paleoclimatological/paleoecological information.