Structure and dynamics of a tropical dry forest in Ghana

Forest under low rainfall (averaging 745 mm yr^-1) on the Shai Hills in S.E. Ghana has redeveloped following cessation of farming in the 1890s. Forest stature is low, with a canopy at about 11 m, principally of three species, Diospyros abyssinica, D. mespiliformis and Millettia thonningii. Drypetes parvifolia and Vepris heterophylla are common understorey trees. Twelve species of woody liane were recorded. Species of thicket vegetation in the area were also present at low density. Most species are evergreen.Tree mortality averaged 2.3% yr^-1 and exceeded recruitment (1.5% yr^-1). Differences between species in mortality and recruitment were pronounced: canopy species showed a small decline in density; understorey species increased markedly and the thicket species declined. Seed production was very variable, but seedling establishment was very poor for all species. Seedling mortality was high (11% yr^-1) especially for small seedlings. These population trends probably represent the latter stages of succession of forest regrowth after farming about 100 years ago.Compared with tropical rain forest, Shai Hills forest has similar relative tree diameter growth (1–3.5% yr^-1), mortality and recruitment rates, and small-litter fall (5.52 t ha^-1 yr^-1).Shai Hills forest differs from rain forest by its short stature, relatively few (evergreen) tree species, poor regeneration from seed, high soil nutrient status and low rainfall. Similar forests have been reported in east Africa and in parts of New Guinea.Abbreviations dbh diameter at breast height (1.3 m) - gbh girth at breast height died May 1984     

Non-structural carbohydrate pools in a tropical forest

The pool size of mobile, i.e. non-structural carbohydrates (NSC) in trees reflects the balance between net photosynthetic carbon uptake (source) and irreversible investments in structures or loss of carbon (sink). The seasonal variation of NSC concentration should reflect the sink/source relationship, provided all tissues from root to crown tops are considered. Using the Smithsonian canopy crane in Panama we studied NSC concentrations in a semi-deciduous tropical forest over 22 months. In the 9 most intensively studied species (out of the 17 investigated), we found higher NSC concentrations (starch, glucose, fructose, sucrose) across all species and organs in the dry season than in the wet season (NSC 7.2% vs 5.8% of dry matter in leaves, 8.8/6.0 in branches, 9.7/8.5 in stems, 8.3/6.4 in coarse and 3.9/2.2 in fine roots). Since this increase was due to starch only, we attribute this to drought-constrained growth (photosynthesis less affected by drought than sink activity). Species-specific phenological rhythms (leafing or fruiting) did not overturn these seasonal trends. Most of the stem volume (diameter at breast height around 40 cm) stores NSC. We present the first whole forest estimate of NSC pool size, assuming a 200 t ha^–1 forest biomass: 8% of this i.e. ca. 16 t ha^–1 is NSC, with ca. 13 t ha^–1 in stems and branches, ca. 0.5 and 2.8 t ha^–1 in leaves and roots. Starch alone (ca. 10.5 t ha^–1) accounts for far more C than would be needed to replace the total leaf canopy without additional photosynthesis. NSC never passed through a period of significant depletion. Leaf flushing did not draw heavily upon NSC pools. Overall, the data imply a high carbon supply status of this forest and that growth during the dry season is not carbon limited. Rather, water shortage seems to limit carbon investment (new tissue formation) directly, leaving little leeway for a direct CO₂ fertilization effects.     

Small-scale patterning in the recruitment of forest species during succession in tropical dry forest,Mozambique

Within a few years of field abandonment on the coastal dunes, a considerable number of species of trees and woody lianes of the climax evergreen forest enter the succession. At a relatively large-scale of analysis (a 100 m² plot), there is no evidence of soil development during the early succession, with no accumulation of soil organic matter nor nutrients being detected. At this scale there is, therefore, little support for the facilitation model of succession. However, when smaller scales of analysis are employed (e.g. a plot equivalent to the canopy area of a sapling) a very different picture emerges; most recruitment of forest woody species occurs beneath saplings found in early succession. Nutrient concentrations were higher in soils under sapling canopies than in soils beyond sapling canopies. With increasing sapling size, an increasing number of seedlings of forest woody species were found and there is progressive soil amelioration. Thus, during the early succession, there is considerable small-scale patterning in soil development and recruitment. This pattern is largely established through bird dispersal, with the intensity of recruitment being dependent on the distance to seed sources.     

Improved recruitment of a lemur-dispersed tree in Malagasy dry forests after the demise of vertebrates in forest fragments

The objective of this study was to examine how the processes of seed dispersal and seed predation were altered in forest fragments of the dry forest of Madagascar, where the usual seed dispersers and vertebrate seed predators were absent, using a lemur-dispersed tree species (Strychnos madagascariensis; Loganiaceae) as an example. We then assessed how the changes in vertebrate community composition alter the regeneration pattern and establishment of this tree species and thus, ultimately, the species composition of the forest fragments. By using size-selective exclosures, data from forest fragments were compared with results from continuous forest where vertebrate dispersers and predators were abundant. Visits to the exclosures by mammalian seed predators were monitored with hair traps. In the continuous forest up to 100% of the seeds were removed within the 7 days of the experiments. A substantial proportion of them was lost to seed predation by native rodents. In contrast, practically no predation took place in the forest fragments and almost all seeds removed were dispersed into the safety of ant nests by Aphaenogaster swammerdami, which improves chances of seedling establishment. In congruence with these findings, the abundance of S. madagascariensis in the forest fragments exceeded that of the continuous forest. Thus, the lack of vertebrate seed dispersers in these forest fragments did not lead to a decline in regeneration of this animal-dispersed tree species as would have been expected, but rather was counterbalanced by the concomitant demise of vertebrate seed predators and an increased activity of ants taking over the role of seed dispersers, and possibly even out-doing the original candidates. This study provides an example of a native vertebrate-dispersed species apparently profiting from fragmentation due to flexible animal-plant interactions in different facets, possibly resulting in an impoverished tree species community.     

Seedling interactions in a tropical forest in Panama

Competition is believed to be a central force limiting local diversity and controlling the structure of plant communities. However, it has been proposed that the stressed understory environment limits total understory plant density to such low levels that competitive exclusion cannot be an important factor limiting the local diversity of understory plants. To evaluate the importance of inter-seedling competition, we performed a seedling competition experiment with five shade-tolerant species in a tropical moist forest in Panama. Three-month-old seedlings were transplanted into the forest singly or with their roots intertwined with a single conspecific or heterospecific seedling in all pairwise species combinations. If competition is important, performance (survival, stem height, and number of leaves after one and six years) would be expected to be lowest with a conspecific neighbor and greatest without a neighbor. The experiment was replicated in five 0.24-m² plots at each of 20 sites in tall secondary forest. To test whether seedling performance differed among treatments we fitted linear mixed models (LMM) and generalized linear mixed models (GLMM), treating species identity and microsite (site and plot) as random effects. The five shade-tolerant study species all experienced good establishment with relatively high survival and growth rates. The neighbor treatment consistently affected seedling performance, but the effect was always very small, both in absolute terms and relative to the much stronger species and microsite effects. Seedlings with a conspecific neighbor consistently performed worse than seedlings with a heterospecific neighbor, but having no neighbor generally did not cause superior performance relative to the other treatments. We conclude that direct competitive interactions are relatively unimportant among understory plants in humid tropical forests.     

Seasonal patterns in a temperate intertidal fish community on the west coast of South Africa

Synopsis Seasonality was investigated in an intertidal ichthyofaunal community on the west coast of South Africa. Fishes were collected monthly from 12 to 16 intertidal pools between May 1992 and May 1993, using the ichthyocide rotenone. On each occasion sampling effort was divided equally between four biologically determined shore zones, namely the cochlear, lower and upper balanoid and littorina zones. A total of 5 409 fishes belonging to 20 species and five families was collected. All fishes were intertidal residents. No seasonal trend could be found in the total density of fishes during the year, the densities of individual species, species diversity or evenness. Thus species composition remained stable over the year. The proportion of mature individuals of each species decreased in summer as a result of recruitment of juveniles at this time. This almost complete lack of seasonality was attributed to the absence of transient species from this community.     

Propagule pressure and native species richness effects drive invasibility in tropical dry forest seedling layers

Understanding the factors that encourage or inhibit plant invasions is vital to focusing limited invasive control efforts within areas where they are most practical and cost-effective. To extend the range of contexts in which invasibility is studied and aid the development of practical strategies to limit damaging plant invasions, we set out to test the relative importance of native species richness, native seedling density, and invasive propagule pressure, on the invasibility of artificial assemblages of naturally occurring tropical woody seedling communities. Our greenhouse mesocosms included a species pool of twelve trees and woody shrubs native to South Florida's tropical hardwood hammocks, and an increasingly prevalent noxious woody invader of this system, Ardisia elliptica. We found that invader propagule pressure was the single most important factor determining community invasibility. We also revealed a positive relationship between invasibility and native species richness in our polyculture mesocosms. Because A. elliptica biomass production significantly differed among different native monocultures and was not related to overyielding in native polycultures, we suggest that the effect of species richness on invasibility in this experiment was the result of sampling effects rather than a true effect of diversity.Three broad findings hold potential for application in preventing and controlling plant invasions, especially in the seedling layers of tropical dry forests: (1) effective invasive control efforts will likely benefit from measures to minimize propagule pressure; (2) managers might do well to prioritize invasive monitoring and removal efforts on the most diverse habitats within a management region; and (3) while more data are necessary to further understand our finding of a lack of association between productivity and invasibility, management regimes aimed at maximizing primary productivity might not increase invasibility, and in fact, strategies for controlling invasive plants via the management of ecosystem productivity may be ineffective.     

Seedling recruitment patterns in a Belizean mangrove forest: effects of establishment ability and physico-chemical factors

A field study was conducted to evaluate the relative importance of factors affecting seedling establishment and survival on a mangrove-dominated island in Belize. An examination of spatial patterns of seedling relative densities in relation to reproductive adults and physico-chemical conditions provided correlative information on factors affecting mangrove regeneration patterns. Distance from reproductive adults explained 89–94% of the variation in relative density of Rhizophora mangle seedlings, whereas availability of resources (light and NH₄) explained 73–80% of variation in Avicennia germinans seedling relative density. Just after dispersal (December), 89% of the variation in Laguncularia racemosa seedling relative density was attributable to distance from reproductive adults, but 7 months later (July) 74% of the variation was explained by intensity of flooding- and salinity-related stresses. Survivorship (after 2.5 years) of propagules and seedlings of R. mangle and A. germinans transplanted to zones of contrasting physico-chemical conditions demonstrated that: (1) mortality was highest during the establishment phase and major causes were failure to strand before viability was lost, consumption by predators and desiccation; and (2) after establishment, differences in sensitivity to physicochemical stress factors such as flooding (A. germinans) and initial orientation of the seedling axis (R. mangle) exerted a further influence on seedling survival. The results indicate that seedling recruitment in these neotropical forests is strongly influenced by dispersal patterns, differential establishment abilities and effects of physico-chemical factors that vary with elevation and distance from the shoreline.     

Root biomass of a dry deciduous tropical forest in Mexico

The deciduous tropical dry forest at Chamela (Jalisco, Mexico) occurs in a seasonal climate with eight rainless (November through June) and four wet months (700 mm annual precipitation). The forest reaches a mean height of 10 m. Tree density in the research area was 4700 trees per ha with a basal area at breast height of 23 m² per ha. The above-and below-ground biomass of trees, shrubs, and lianas was 73.6 Mg ha^–1 and 31 Mg ha^–1, respectively. A root:shoot biomass ratio of 0.42 was calculated. Nearly two thirds of all roots occur in the 0–20 cm soil layer and 29% of all roots have a diameter of less than 5 mm.     

Grazing as a mediator for maintenance of offspring diversity: Sexual and clonal recruitment in alpine grassland communities

To understand the effects of grazing on grassland plants sexual and clonal recruitment, we conducted a demographic field investigation of species recruitment along a grazing gradient in the Tibetan alpine grassland. Grazing intensity had significant effects on quantity and diversity of sexual and clonal recruitment. Sexual recruitment increased significantly, but clonal offspring production decreased significantly with increased grazing intensity. Grazing intensity had different, significant effects on offspring recruitment of the various functional groups in the community, grasses (GG), sedges (SG), legumes (LG) and forbs (FG). Higher grazing intensity reduced offspring recruitment of GG and SG; it increased offspring recruitment of LG and FG. Seedlings were significantly more abundant in lightly grazed, moderately grazed and heavily grazed meadows than in non-grazed grasslands. Offspring diversity from sexual recruitment was significantly higher than that from clonal recruitment in grazed than in non-grazed grasslands. Our studies indicate that moderate grazing had positive effects on seedling recruitment and offspring diversity, but heavy gazing may alter community succession by affecting recruitment patterns among the four plant functional groups.     

The effect of mollusc grazing on seedling recruitment in artificially created grassland gaps

Two experiments conducted in spring and autumn 1992 examined the effect of mollusc grazing on seedling regeneration from natural grassland seedbanks by creating artificial gaps in plots in a grassland sward. Molluscs were excluded from half the gaps by application of molluscicide. Mollusc grazing in both the spring and autumn experiment significantly reduced seedling recruitment, though the intensity of grazing was greatest in autumn. Recruitment of five species was markedly influenced by molluscicide application. In spring, plots from which molluscs were excluded contained significantly more seedlings of Chenopodium polyspermum and Ranunculus acris. In the autumn, exclusion of molluscs resulted in increased populations of R. acris, Stellaria graminea and Rumex acetosa. Cerastium holosteoides populations were greatest in autumn grazed plots. Other species, notably the grasses Holcus lanatus and Agrostis capillaris and the legume Trifolium repens were unaffected by molluscicide application. Species diversity was significantly decreased by molluscicide application in the autumn. Gap size significantly affected the recruitment of two species. Ranunculus acris populations were significantly higher in small gaps in both spring and summer, while Chenopodium recruitment in the spring was greater in small gaps. Gap size also significantly influenced the risk of mollusc attack on Ranunculus as molluscs appeared to show an aggregative feeding response in the high seedling density small gaps. Selective grazing of vulnerable seedlings by molluscs may influence the eventual relative proportions of the species present and so provide a potent mechanism in shaping community composition in grasslands.     

Red spruce forest regeneration dynamics across a gradient from Acadian forest to old field in Greenwich,Prince Edward Island National Park,Canada

Red spruce forests have declined considerably throughout their range in the past decades. As agricultural fields are abandoned and land becomes available for reforestation, the possibility arises for red spruce forests to expand onto them. This study addresses the potential for red spruce forests to expand onto adjacent old fields in Greenwich, Prince Edward Island National Park, Canada. We examined red spruce distribution and abundance, plant species diversity and changes in community composition along a gradient from the interior of red spruce forests out towards the centre of adjacent old fields. Examining the patterns of red spruce distribution and abundance revealed that, where cultivation and logging have been abandoned recently in the fields and forests, regeneration is limited to the forest stands, but in the sites with older fields and forests, regeneration extends into and is more vigorous in the fields. Although species diversity varied from forest to field only for the tree and shrub layers, important changes occurred in the ground species composition. There is no evidence yet that the herbaceous species present in the forest stands will colonise the old fields. The results suggest that both environmental differences among sites and length of time since the fields were abandoned explain red spruce regeneration patterns. In order to more accurately assess the potential for red spruce regeneration in old fields, long-term monitoring of the production, dispersal and viability of red spruce seeds from adjacent forests and of the constraints to seedling establishment and survival in old fields will be needed.     

Albedo model for a tropical dry deciduous forest in western Mexico

A simple albedo model is presented for a tropical dry deciduous forest. The model is based on point observations of the solar radiation, leaf cover of the vegetation, precipitation and air temperature from 1981 to 1988 in western Mexico. Four main periods were noted: leafing, leafed, leaf-fall and leafless. During the leafed period the albedo was almost constant (0.16) but increased slowly in the leaf-fall period, at a rate of 0.0008/day, until its maximal values in the leafless period (0.24). During the leafing and leafed periods, the albedo decrease was a hyperbolic function of precipitation at a rate of 7.9 albedo percentage/mm. Albedo showed a linear regression on leaf cover and decreased at a rate of 0.119 albedo percentage per leaf cover percentage.     

Liana habitat associations and community structure in a Borneanlowland tropical forest

Lianas (woody vines) contribute substantially to the diversity and structure of most tropical forests, yet little is known about the importance of habitat specialization in maintaining tropical liana diversity and the causes of variation among forests in liana abundance and species composition. We examined habitat associations, species diversity, species composition, and community structure of lianas at Sepilok Forest Reserve, Sabah, Malaysia in northeastern Borneo among three soil types that give rise to three distinct forest types of lowland tropical rain forest: alluvial, sandstone hill, and kerangas (heath) forest. Alluvial soils are more nutrient rich and have higher soil moisture than sandstone soils, whereas kerangas soils are the most nutrient poor and drought prone. Lianas ≥0.5-cm in diameter were measured, tagged, and identified to species in three square 0.25-ha plots in each forest type. The number of lianas ≥0.5 cm did not differ significantly among forest types and averaged 1348 lianas ha⁻¹, but mean liana stem diameter, basal area, estimated biomass, species richness, and Fisher’s diversity index were all greater for plots in alluvial than sandstone or kerangas forests. Liana species composition also differed greatly among the three habitats, with 71% of species showing significant positive or negative habitat associations. Sandstone forests were intermediate to alluvial and kerangas forests in most aspects of liana community structure and composition, and fewer species showed significant habitat associations with this forest type. Ranking of forest types with respect to liana density, biomass, and diversity matches the ranking in soil fertility and water availability (alluvial > sandstone hill > kerangas). These results suggest that edaphic factors play an important role in maintaining liana species diversity and structuring liana communities.     

The influence of ants on patterns of colonization and establishment within a set of coexisting lycaenid butterflies in a south-east Asian tropical rain forest

In Peninsular Malaysia ten species of lycaenid butterflies use leaf flushes or inflorescences of the legume tree Saraca thaipingensis as larval hostplant. Resource partitioning among these species is regulated by a complex mixture of patterns of interaction with ants. Females of obligately myrmecophilous species lay their eggs exclusively on trees colonized by their specific host ants. On trees colonized by weaver ants, only specialist mutualists adapted to these territorial ants are able to survive, while larvae of other species are killed. The formicine ant Cladomyrma petalae, which inhabits hollow twigs of the myrmecophytic hostplant, likewise precludes oviposition by female butterflies. Lycaenid larvae confronted with this ant species never survive, but one concealed feeding species (Jamides caeruleus) escapes removal due to the cryptic life-habits of the larvae. Two facultative myrmecophiles associate in a mutualistic way with a wide and largely overlapping range of ant genera which forage at the extrafloral nectaries of leaf flushes. One species (Cheritra freja) is not myrmecophilous, but is tolerated by all but the most territorial ants. Ant-dependent hostplant selection and egg-clustering characterize the obligate mutualists, whereas facultative myrmecophiles and the non-myrmecophile distribute their eggs singly over appropriate hostplants. Signals mediating caterpillar-ant communication are highly specialized in one obligate myrmecophile (Drupadia theda), but rather unspecific in four other species tested. Altogether our observations indicate that colonization and establishment of lycaenid butterflies on S. thaipingensis trees are governed by specializations as well as opportunistic use of resources (ants and hostplant parts). Therefore, the diversity of this species assemblage is maintained by deterministic as well as stochastic factors.     

Nitrogen and phosphorus resorption in trees of a Mexican tropical dry forest

Resorption efficiency (RE) and proficiency, foliar nutrient concentrations, and relative soil nutrient availability were determined during 3 consecutive years in tree species growing under contrasting topographic positions (i.e., top vs. bottom and north vs. south aspect) in a tropical dry forest in Mexico. The sites differed in soil nutrient levels, soil water content, and potential radiation interception. Leaf mass per area (g m^–2) increased during the growing season in all species. Soil P availability and mean foliar P concentrations were generally higher at the bottom than at the top site during the 3 years of the study. Leaf N concentrations ranged from 45.4 to 31.4 mg g^–1. Leaf P varied from 2.3 to 1.8 mg g^–1. Mean N and P RE varied among species, occasionally between top and bottom sites, and were higher in the dry than in the wet years of study. Senesced-leaf nutrient concentrations (i.e., a measure of resorption proficiency) varied from 13.7 to 31.2 mg g^–1 (N) and 0.4 to 3.3 mg g^–1 (P) among the different species and were generally indicative of incomplete nutrient resorption. Phosphorus concentrations in senesced leaves were higher at the bottom than at the top site and decreased from the wettest to the the driest year. Soil N and P availability were significantly different in the north- and south-facing slopes, but neither nutrient concentrations of mature and senesced leaves nor RE differed between aspects. Our results suggest that water more than soil nutrient availability controls RE in the Chamela dry forest, while resorption proficiency may be interactively controlled by both nutrient and water availability.     

Seedling growth of climbing species from a southeast Brazilian tropical forest

Climbers are considered heliophytes. They are copious at the margins of forests and natural and man-made clearings. The objective of this paper was to study the initial growth of seedlings maintained under full sunlight and shaded conditions (under a vegetation canopy). The species studied were: Aristolochia galeata, Arrabidea triplinervia, Bidens brasiliensis, Canavalia parviflora, Chamissoa altissima, Cissus sicyoides, Dalechampia pentaphylla, Dicella bracteosa, Dioscorea sp., Gouania virgata, Mascagnia anisopetala, Mutisia coccinea, Oxypetalum molle, Pithecoctenium crucigerum, Rynchosia phaseoloides, Serjania multiflora and Solanum flaccidum. The initial growth of the seedlings was followed under two conditions: at the margin and under the canopy of a mesophyllous tropical forest (22° 4955S–47° 0633W). The climbers showed high rates of growth in sunlight when compared to those under canopy. Most of the species presented higher growth of the shoot than roots but in general no significative differences between root/shoot were found in both treatments. Bidens, Cissus, Mutisia and Pithecoctenium showed a very high mortality rate under canopy but, most of the studied species survived under deep shaded forest for approximately 100 days.