Recovery of species richness and conservation of native Atlantic forest trees in the cacao plantations of southern Bahia in Brazil

The Atlantic forests of southern Bahia in Brazil present great species richness and a high degree of endemism. A large part of these native forests were transformed into cacao plantations in an agroforestry system known locally as cabrucas, where native trees were culled and cacao was planted under the shade of remaining trees. The present study analyzed the influence of time of implantation (age) and time of abandonment of management practices on tree species diversity of cabruca plantations to evaluate the capacity for conservation and recovery of species richness of native Atlantic Forest trees in cabrucas. Phytosociological surveys were conducted in five cabrucas with different conditions of age and state of abandonment. All trees, including hemiepiphytes and excluding the cacao plants, with a minimum stem diameter of 10 cm at breast height, were surveyed within a 3-ha sampling area in each plantation. A total of 2514 individual trees belonging to 293 species and 52 families were recorded in the five cabrucas. The Shannon diversity index varied from 3.31 to 4.22 among the cabrucas and was positively correlated with the time of abandonment (r = 0.97). The new cabrucas showed the highest values of estimated total richness (Chao) and the highest proportion of late successional species than the old ones. All areas preserved a very high proportion of native forest species while the three old cabrucas showed a higher proportion of exotic species than the two new ones. Thus the exotic species seem to replace more of the native species in the long run because of management practices and local preferences. The cabrucas presented also a high capacity for the regeneration of tree species richness after abandonment. Simple alterations in management practices could improve the recruitment of late successional species in these areas. Economic incentives may be necessary for the farmers to adopt management practices to retain native species which bring no economic returns.     

Facilitation and competition influence succession in a subtropical old field

The effects of neighboring vegetation and soil fertility on the establishment, survival and growth of tree species were studied in a subtropical old-field area in south Brazil. Seed damage, germination and seedling establishment of four tree species plus growth and survival of two transplanted tree species were monitored under factorial combinations of the following treatments: (1) pioneer vegetation (presence and absence); (2) soil fertility (addition of NPK and control). Facilitation was the main process affecting plant performance. The presence of pioneer vegetation significantly improved germination, establishment, growth and survival of most study species. Around 90% of sown seeds were damaged and the removal of pioneer vegetation significantly increased seed damage for all species studied, decreased germination in three out of four species, and decreased establishment in one species. Moreover a significantly higher seedling growth rate of Inga virescens was found after the first year of the experiment in plots where vegetation was present. The presence of vegetation significantly increased seedling survival of I. virescens by protecting seedlings from leaf loss due to winter frosts. Competition was detected by the second year when a higher growth of transplanted seedlings of the species Araucaria angustifolia occurred in plots where vegetation was present and fertilizer were applied. A lower growth rate was detected in plots where vegetation was present but fertilizer was not applied. These results indicate a balance between competition for soil nutrients and protection by neighbor vegetation. Damage of seedlings by leaf cutter ants was an important barrier for plant survival. Damage occurred in 80% of the A. angustifolia seedlings and 58% of these damaged seedlings died. The presence of neighboring vegetation tended to protect seedlings from ant damage. Although competition occurred, facilitation seems to be the main process driving early successional changes in this subtropical old field. This was mainly due to the improvement of local microclimatic conditions and protection against herbivores by neighboring vegetation. Facilitation occurred during establishment and growth phases in a subtropical area that is considered a productive, low stress environment. Our results indicate that facilitation may be more frequent in productive environments than previously thought.     

Seed dispersal, plant recruitment and spatial distribution of Bactris acanthocarpa Martius (Arecaceae) in a remnant of Atlantic forest in northeast Brazil

Seed dispersal ecology of Bactris acanthocarpa Mart. (Arecaceae), an Atlantic forest understory palm, was investigated during two years as an attempt to test the following predictions: (i) seeds of Bactris are dispersed by mammals and large-gaped birds; (ii) Bactris benefits from seed dispersal in terms of seed predation avoidance, improvement of seed germination and seedling survival; and (iii) spatial distribution of adults is related to patterns of seed dispersal. The study was conducted at Dois Irmãos Reserve, a 387.4-ha reserve of Atlantic forest in northeastern Brazil (8º S–35º W). Black–rumped agoutis (Dasyprocta prymnolopha) and Guianan squirrels (Sciurus aestuans) were identified as the seed dispersers/predators, moving seeds short distances (< 4 m from parents) and at low rates (0.04-0.05 diaspore/palm/day). Pyrene burial prevented seed predation by vertebrates and reduced by half seed infestation by Scolytidae beetles. Only buried pyrenes germinated. Pyrene predation was not correlated with distance from conspecific adults. In contrast, early seedling mortality was higher near conspecific adults. Most adults (64%) had their nearest conspecific adult neighbour > 4 m away in contrast to 96% of seedlings that occurred concentrated within 4 m from adults (77% under the palm crowns). Here, we present evidence that spatial distribution of B. acanthocarpa is partly due to low rates of seed removal, short-distance seed dispersal by agoutis and squirrels, and early seedling mortality associated with presence of seedlings under palm crowns.     

Bird diversity and abundance in forest fragments and Eucalyptus plantations around an Atlantic forest reserve, Brazil

Little of Brazil's remaining Atlantic forest is protected, so it is important to assess how well the region's wildlife can persist in areas/habitats outside reserves. We studied bird diversity and abundance during 546 point counts in the Sooretama/Linhares reserve, 200 point counts in 31 forest fragments (10–150 h), and 50 point counts in <30-year-old Eucalyptus plantations, within 7 km of the reserve. Only eight bird species were recorded in Eucalyptus, and this impoverishment, as compared to some Eucalyptus plantations elsewhere in Brazil may be a result of intensive clearance of understory vegetation. Species diversity in forest fragments was significantly lower than in the reserve. Twelve, mostly non-forest or edge species, were significantly commoner in the fragments, but nineteen species were frequent in the reserve but rare or absent in forest fragments. These included two Pyrrhura parakeets, a Brotogeris parakeet, a trogon Trogon, a jacamar Galbula, woodpeckers Piculus and Campephilus, Myrmotherula antwrens, and Hemithraupus and Tachyphonus tanagers. Bird species richness at points in forest fragments did not decline with fragment size, distance from the reserve, or forest quality. However, forest in fragments was more heavily degraded than forest within the reserve and poor forest quality may be the cause of declines in some species. Whilst protection of forest within reserves is a priority, management of forest fragments may aid conservation of some threatened species.     

Effect of plant growth regulators on the cellular growth and levels of intracellular protein,starch and polyamines in embryogenic suspension cultures of Pinus taeda

Somatic embryogenesis is the most important in vitro culture system for conifer propagation. However, Pinus taeda has been considered recalcitrant to somatic embryogenesis in commercial scale-up. The study of biochemical and physiological aspects of cell growth could lead to a better understanding of somatic embryogenesis in this species. In the present work, we investigated the cell growth dynamics, intracellular levels of proteins, starch and polyamines in suspension cultures of Pinus taeda established in plant growth regulator-free medium (BM0) and in medium supplemented with 2 M 2,4-dichlorophenoxyacetic acid, 0.5 M 6-benzylaminopurine and 0.5 M Kinetin (BM2). Cell cultures growing in BM0 medium showed an increase in the sedimented cell volume from 3.77 to 17.73 ml after 24 days of culture. Those cultured in BM2 medium showed an increase in the sedimented cell volume from 4.23 to 25.17 ml after 20 days of culture. Intracellular proteins levels increased during the exponential growth phase and starch levels decreased until the exponential phase, followed by a synthesis up to the stationary phase, in both BM0 and BM2 media. Highest putrescine levels occurred in cultures growing in BM0 medium and this was associated with the low cellular growth.     

Effects of microhabitat on palm seed predation in two forest fragments in southeast Brazil

The establishment of plants depends crucially on where seeds are deposited in the environment. Some authors suggest that in forest understory seed predation is lower than in gaps, and higher than at the forest edge. However, most studies have been carried out in large forest patches and very little is known about the effects of microhabitat conditions on seed predation in forest fragments. We evaluated the effects of three microhabitats (gaps, forest edge, and understory) on seed predation of two palm species (Euterpe edulis and Syagrus romanzoffiana) in two semi-deciduous forest fragments (230 and 2100 ha) in southeast Brazil. Our objective was to test two hypotheses: (1) Low rodent abundance in small fragments as a result of meso-predator action levels leads to lower seed predation in small fragments. (2) Most mammal species in small fragments are generalists with respect to diet and habitat, so that seed predation is similar in different microhabitats (gaps, forest edge and understory) in the small fragment, but not in the larger one. The study community of small fragments is usually composed of generalist species (in diet and habitat aspects), so we expected the same rate of seed predation among microhabitats (gaps, forest edge and understory) in the tested smaller fragment. The experiment was carried out in the dry season (for E. edulis) and in the wet season (for S. romanzoffiana) in 1999. We conclude that post-dispersal seed predation in forest fragments can be directly connected with mammal communities, reflecting their historical and ecological aspects.     

Slow death of Atlantic forest trees in cocoa agroforestry in southeastern Brazil

Cocoa (Theobroma cacao) is cultivated in the states of Bahia and Espírito Santo in eastern Brazil under the so-called cabruca system, where the understorey of native Atlantic forest is cleared and the canopy is thinned out to provide adequate shading for the cocoa trees. Apart from its economic and social role, the cabruca system is said to be important for the conservation of Atlantic forest biodiversity. In this paper we studied tree species richness and forest structure of cabrucas to examine the demographic health of these forests and discuss their long-term survival. Data were collected in 20 farms located alongside a 30km track of the northern margin of the Rio Doce, in northern Espírito Santo. All trees 5cm DBH were identified and their diameter was measured in 80 plots (600m²), totalling 4.8ha of sampled area. Recorded trees were also allocated to four different regeneration phases (pioneers, early secondary, late secondary and climax). The inventory resulted in 507 trees belonging to 105 species in 39 families. This species richness is much lower than in less disturbed forests located in the region. Pioneers and early secondary species dominate the cabruca forest in terms of number of species (56.2%), density (71.0%) and basal area (72.3%). The distribution of diameter frequency showed an imbalance in tree regeneration. Most trees in the range of 5–30cm DBH were pioneers (40.7%), or early secondary species (32.6%), while late secondary and climax trees were less frequent (10.2 and 16.5% of the sampled trees, respectively). The dominance of species of early regeneration phases was also observed for trees >30cm DBH (69.0% of pioneers or early secondary and 31.0% of late secondary or climax species). The results indicated that the cabruca forests are not only less diverse and less dense than secondary or primary forests of the region, but also, and more importantly, their natural succession and gap dynamics are being severely impaired. As a consequence, cabrucas present a structure where tree species of late successional phases are becoming increasingly rare while pioneers and early secondary species are becoming dominant. If current management practices of thinning and clearing of native trees are not improved, the long-term survival of these forests is questionable and their role in maintaining biodiversity in the long run is limited.     

Ficus seed shadows in a Bornean rain forest

Due to their copious seed production and numerous dispersers, rain forest fig trees have been assumed to produce extensive and dense seed shadows. To test this idea, patterns of seed dispersal of two species of large hemiepiphytic fig tree were measured in a Bornean rain forest. The sample included four Ficus stupenda and three F. subtecta trees with crop sizes ranging from 2,000 to 40,000 figs (400,000 to 13,000,000 seeds). Seed rain out to a distance of 60 m from each study tree was quantified using arrays of seed traps deployed in the understory. These trees showed a strongly leptokurtic pattern of dispersal, as expected, but all individuals had measurable seed rain at 60 m, ranging from 0.2 to 5.0 seeds/m². A regression of In-transformed seed rain density against distance gave a significant fit to all seven trees' dispersal patterns, indicating that the data could be fitted to the negative exponential distribution most commonly fitted to seed shadows. However, for six of seven trees, an improved fit was obtained for regressions in which distance was also In-transformed. This transformation corresponds to an inverse power distribution, indicating that for vertebrate-dispersed Ficus seeds, the tail of the seed rain distribution does not drop off as rapidly as in the exponential distribution typically associated with wind dispersed seed shadows. Over 50% of the seed crop was estimated to fall below each fig tree's crown. Up to 22% of the seed crop was dispersed beyond the crown edge, but within 60 m of the tree. Estimates of the maximum numbers of seeds which could have been transported beyond 60 m were 45% for the two largest crops of figs, but were under 24% for the trees with smaller crops. Seed traps positioned where they had an upper canopy layer above them were associated with higher probabilities of being hit by seeds, suggesting that vertebrate dispersal agents are likely to perch or travel through forest layers at the same level as the fig crown and could concentrate seeds in such areas to some degree. The probability of a safe site at 60 m from the fig tree being hit by seeds is calculated to be on the order of 0.01 per fruiting episode. Fig trees do not appear to saturate safe sites with seeds despite their large seed crops. If we in addition consider the rarity of quality establishment sites and post-dispersal factors reducing successful seedling establishment, hemiepiphytic fig trees appear to face severe obstacles to seedling recruitment.     

Feeding behavior of Geoffroy's marmoset (Callithrix geoffroyi) in an Atlantic forest fragment of South-eastern Brazil

We report on the diet and feeding behaviour of a group of Geoffroy's marmosets (Callithrix geoffroyi) in an Atlantic forest fragment in south-eastern Brazil, studied during the period February 1993 to Januaray 1994. Major food categories were gums (68.6%) fruits (15%), and small animal prey (invertebrates 14.6% and vertebrates 0.8%). Dietary changes were observed between the wet and dry seasons, although they were not statistically significant. Insects and gums were consumed throughout the year but fruits were more prevalent in the diet in the wet season. Plant species exploited for their gums includedInga stipularis, followed byAcacia paniculata, Paulinia carpopodia, andBauhinia angulosa. Chemical analysis of the gum of the four species most used all presented high values for carbohydrates and crude proteins. Fruits of an unidentified species of Myrtaceae (sp. 2) were consumed the most. Animal prey consisted mainly of insects, arachnids, snails, and in three cases, lizards and frogs, with orthopterans being the most prevalent insect prey. This study demonstrates thatC. geoffroyi efficiently exploits resources typically available in secondary and disturbed forest habitat. The main threat to the species is forest destruction, degradation, and fragmentation, but the management of small forest fragments may be an effective corservation strategy.     

Landscape and farm scale management to enhance biodiversity conservation in the cocoa producing region of southern Bahia, Brazil

In southern Bahia, Brazil, large land areas are used for the production of cocoa (Theobroma cacao), which is predominantly grown under the shade of native trees in an agroforestry system locally known as cabruca. As a dominant forest-like landscape element of the cocoa region, the cabrucas play an important role in the conservation of the region’s biodiversity. The purpose of this review is to provide the scientific basis for an action plan to reconcile cocoa production and biodiversity conservation in southern Bahia. The available research collectively highlights the diversity of responses of different species and biological groups to both the habitat quality of the cabrucas themselves and to the general characteristics of the landscape, such as the relative extent and spatial configuration of different vegetation types within the landscape mosaic. We identify factors that influence directly or indirectly the occurrence of native species in the cabrucas and the wider landscape of the cocoa region and develop recommendations for their conservation management. We show that the current scientific knowledge already provides a good basis for a biodiversity friendly management of the cocoa region of southern Bahia, although more work is needed to refine some management recommendations, especially on shade canopy composition and density, and verify their economic viability. The implementation of our recommendations should be accompanied by appropriate biological and socioeconomic monitoring and the findings should inform a broad program of adaptive management of the cabrucas and the wider cocoa landscape.     

Phenological changes in rate of respiration and annual carbon balance in a perennial herbaceous plant, <Emphasis Type="Italic">Primula sieboldii</Emphasis>

Primula sieboldii E. Morren is a clonal herbaceous species with a short foliar period from spring to early summer. We have studied the temperature-dependence of the rate of respiration at the whole-ramet level throughout the phenological stages of P. sieboldii to reveal its photosynthate-utilization strategy. P. sieboldii ramets were grown in a chamber enabling simulation of seasonal changes in temperature. Rates of respiration were measured at three phenological stages—the foliar period, the before-chilling defoliated (BCD) period, and the after-chilling defoliated (ACD) period. In the foliar period the rate of respiration, on a biomass basis at 20°C (R ₂₀), of the above-ground plant parts was 2.5 times that of the below-ground parts. The R ₂₀ of the below-ground parts in the foliar period was 6.5 times that in the BCD period and 1.6 times that in the ACD period. Estimation of the ramet carbon balance under these growth conditions showed that ramets respired 87% of total photosynthate production during the experimental period (8.5 months). Respiratory consumption in the foliar period accounted for 70% of the yearly total, whereas 24 and 6% were consumed in the BCD and ACD periods, respectively. An extremely low rate of respiration during the long defoliated period led to a positive net annual carbon balance for P. sieboldii ramets.     

The decomposition of mangrove litter in a subtropical mangrove forest

Decomposition of Avicennia marina (Forsk.) Vierh. leaf and woody litter (twigs) was studied using litter bag experiments in a subtropical mangrove forest at two tidal levels (high and low) with different inundation regimes and during two seasons (summer and winter). Losses in dry weight were best described by a single exponential model which showed loss rates of both leaves and twigs were significantly higher low down on the shoreline (greater inundation) and in summer. The time (days) required for the loss of half of the initial dry weight (t ₅₀) was summer: high 59, low 44; winter: high 98, low 78. For twigs the values (days) were summer: high 383, low 179; winter; high 1327, low 1207. There is an exponential relationship between leaf litter t ₅₀'s and latitude which indicates the importance of temperature and therefore season, to the dynamics of organic cycling and export in mangrove systems.     

Biomass and production of a naturally regenerated oak forest in southern Korea

Biomass and production of two stands with Quercus variabilis Bl. as the dominant species (stands 1 and 3) and one with Q. mongolica Fisch. as the dominant species (stand 2) were investigated in southern Korea. Stands 1 and 3 naturally occurred on sites with southerly aspects while stand 2 naturally occurred on northerly aspects; stand ages were similar for the three stands (36–38 years old). Total above- and belowground biomass including understory vegetation (Mg ha^–1) was 108.4 for stand 1, 115.6 for stand 2, and 132.0 for stand 3, respectively. Understory vegetation constituted 17.4% of the total biomass in stand 1 but only 3.7–4.5% in stand 2 and stand 3. Roots constituted 20.1–24.6% of the biomass of the overstory vegetation. Although stand 3 showed the highest total biomass, net production was highest in stand 2 at 12.6 (Mg ha^–1 year^–1); net production levels for stands 1 and 3 were 11.7 and 11.1 (Mg ha^–1 year^–1), respectively. It appeared that the differences in site conditions related to aspect influenced the distribution of naturally regenerated oak species within a relatively small area and resulted in differences in biomass and production among the stands.     

Distyly and sequential pollinators of Psychotria nuda (Rubiaceae) in the Atlantic rain forest, Brazil

Reciprocal herkogamy is known to promote intermorph-pollination in distylous species, which are generally self- and intramorph-incompatible. Legitimate crossing rates are also influenced by pollinator foraging strategy, which determines most of the pollen flow in natural populations. This study reports on the floral morphology, compatibility relations and pollination biology of Psychotria nuda (Cham. and Schltdl.) Wawra in the Atlantic rain forest, southeastern Brazil, based on floral measurements, hand-pollination experiments and focal observation of floral visitors. It was found that P. nuda does not present an exactly reciprocal herkogamy, however, this morphological trait in P. nuda is as good as in most of Rubiaceae. Reproductive experiments showed compatibility relations similar to those frequently observed in distylous species. The hummingbirds Ramphodon naevius and females of Thalurania glaucopis were the main pollinators of P. nuda, presenting the traplining foraging strategy, which seems to maximize intermorph crosses in P. nuda flowers. These hummingbirds pollinated P. nuda flowers sequentially throughout the flowering period.     

Primary production and phytoplankton in two small,polyhumic forest lakes in southern Finland

Primary production and phytoplankton in polyhumic lakes showed a very distinct seasonal succession. A vigorous spring maximum produced by Chlamydomonas green algae at the beginning of the growing season and two summer maxima composed mainly of Mallomonas caudata Iwanoff were typical. The annual primary production was ca. 6 g org. C · m^–2 in both lakes. The mean epilimnetic biomass was 1.1 in the first lake and 2.2 g · m^–2 (ww) in the second one. The maximum phytoplankton biomass, 14 g · m^–2, was observed during the vernal peak in May.     

Leaf age and the timing of leaf abscission in two tropical dry forest trees

We used experimental defoliations to examine the effect of leaf age on the timing of leaf shedding in two tropical dry forest trees. Trees of the deciduous Bombacopsis quinata (bombacaceae, a.k.a. Pachira quinata) and the brevi-deciduous Astronium graveolens (anacardiaceae) were manually defoliated for three times during the rainy season. All trees started to produce a new crown of leaves 2 weeks after defoliation, and continued expanding leaves throughout the rainy season. At the transition to the dry season, the experimental groups consisted of trees with known differences in maximum leaf age. Defoliations resulted in declines in stem growth but did not affect the mineral content or water relations of the leaves subsequently produced. There was no effect of leaf age on the timing of leaf abscission in B. quinata. In A. graveolens, the initiation of leaf shedding followed in rank order, the maximum leaf age of the four treatments, but there was substantial coherence among treatments in the major period of leaf abscission such that trees completed leaf shedding at the same time. In the two species, leaf water potential (Ψ_L) and stomatal conducantce (g _S) declined with the onset of the dry season, reaching minimum values of –0.9 MPa in P. quinata and <–2.0 MPa in A. graveolens. Within each species, leaves of different age exhibited similar Ψ_L and g _S at the onset of drought, and then decreased at a similar rate as the dry season progressed. Overall, our study suggests that the environmental factors were more important than leaf age in controlling the timing of leaf shedding.     

Diet of a muriqui group (<Emphasis Type="Italic">Brachyteles arachnoides</Emphasis>) in continuous primary forest

The feeding behaviour of a group of free-ranging muriquis (Brachyteles arachnoides) was monitored in the 380 km² Carlos Botelho State Park (PECB), between February 1992 and November 1993. Scan sample data indicated that 59.1% of feeding time was devoted to fruit, 33.2% to leaves, 4.1% to flowers, and 3.6% to other items (twigs, stem and bark). Little seasonal variation was recorded. These results contrast with those of all previous studies of Brachyteles, in which the consumption of leaves was generally double that of fruit. One key difference in comparison with previous studies is that the PECB is part of the largest remaining continuous area of primary Atlantic forest, in the Serra do Mar coastal range. Such intraspecific differences in ecology may have important implications for the conservation of the species.     

Croton campanulatus (Euphorbiaceae s.s.), a new species from the Brazilian Atlantic rain forest

Croton campanulatus, a new species from southeastern Brazil in the states of Minas Gerais and Rio de Janeiro, is here described and illustrated. Morphological data indicate that this species belongs to Croton section Cleodora based on its arborescent habit, pistillate flowers with imbricate sepals, reduced petals, and multifid styles that are fused at the base.