Relationship between annual rainfall and tree mortality in a tropical dry forest: Results of a 19-year study at Mudumalai, southern India

Variability in rainfall is known to be a major influence on the dynamics of tropical forests, especially rates and patterns of tree mortality. In tropical dry forests a number of contributing factors to tree mortality, including dry season fire and herbivory by large herbivorous mammals, could be related to rainfall patterns, while loss of water potential in trees during the dry season or a wet season drought could also result in enhanced rates of death. While tree mortality as influenced by severe drought has been examined in tropical wet forests there is insufficient understanding of this process in tropical dry forests. We examined these causal factors in relation to inter-annual differences in rainfall in causing tree mortality within a 50-ha Forest Dynamics Plot located in the tropical dry deciduous forests of Mudumalai, southern India, that has been monitored annually since 1988. Over a 19-year period (1988-2007) mean annual mortality rate of all stems >1 cm dbh was 6.9 +/- 4.6% (range = 1.5-17.5%); mortality rates broadly declined from the smaller to the larger size classes with the rates in stems >30 cm dbh being among the lowest recorded in tropical forest globally. Fire was the main agent of mortality in stems 1-5 cm dbh, elephant-herbivory in stems 5-10 cm dbh, and other natural causes in stems > 10 cm dbh. Elephant-related mortality did not show any relationship to rainfall. On the other hand, fire-related mortality was significantly negatively correlated to quantity of rainfall during the preceding year. Mortality due to other causes in the larger stem sizes was significantly negatively correlated to rainfall with a 2-3-year lag, suggesting that water deficit from mild or prolonged drought enhanced the risk of death but only with a time lag that was greater than similar lags in tree mortality observed in other forest types. In this respect, tropical dry forests growing in regions of high rainfall variability may have evolved greater resistance to rainfall deficit as compared to tropical moist or temperate forests but are still vulnerable to drought-related mortality.

Reproductive phenology of a tropical dry forest in Mudumalai, Southern India

1 Flowering and fruiting phenologies of a tropical dry forest in Mudumalai, southern India, were studied between April 1988 and August 1990. Two sites, a wetter site I receiving 1100mm and a drier site II receiving 600mm of rainfall annually, are compared. A total of 286 trees from 38 species at site I and 167 trees from 27 species at site II was marked for phenological observations. There were 11 species common to the two sites. Several hypotheses relating to the evolution of reproductive phenology are tested. 2 Frequency of species flowering attained a peak at site I during the dry season but at site II, where soil moisture may be limiting during the dry months, the peak was during the wet season. At both sites a majority of species flushed leaves and flowered simultaneously. Among various guilds, the bird-pollinated guild showed distinct dry season flowering, which may be related to better advertisement of large flowers to pollinators during the leafless dry phase. The wind-pollinated guild flowered mainly during the wet season, when wind speeds are highest and favourable for pollen transport. The insect-pollinated guild showed no seasonality in flowering in site I but a wet season flowering in site II. 3 Fruiting frequency attained a peak in site I during the late wet season extending into the early dry season; a time-lag correlation showed that fruiting followed rainfall with a lag of about two months. Site II showed a similar fruiting pattern but this was not statistically significant. The dispersal guilds (animal, wind, and explosive passively-dispersed) did not show any clear seasonality in fruiting, except for the animal-dispersed guild which showed wet season fruiting in site I. 4 Hurlbert's overlap index was also calculated in order to look at synchrony in flowering and fruiting irrespective of climatic (dry and wet month) seasonality. In general, overlap in flowering and fruiting guilds was high because of seasonal aggregation. Among the exceptions, at site II the wind-pollinated flowering guild did not show significant overlap between species although flowering aggregated in the wet season. This could be due to the need to avoid heterospecific pollen transfer. 5 Rarer species tended to flower earlier in the dry season and this again could be an adaptation to avoid the risk of heterospecific pollen transfer or competition for pollinators. The more abundant species flowered mainly during the wet season. Species which flower earlier have larger flowers and, having invested more energy in flowers, also have shorter flower to fruit durations.

Woody plant seedling distribution under invasive Lantana camara thickets in a dry-forest plot in Mudumalai, southern India

Lantana camara, a shrub of Central and South American origin, has become invasive across dry forests worldwide. The effect of the thicket-forming habit of L. camara as a dispersal and recruitment barrier in a community of native woody seedlings was examined in a 50-ha permanent plot located in the seasonally dry forest of Mudumalai, southern India. Sixty 100-m(2) plots were enumerated for native woody seedlings between 10-100 cm in height. Of these, 30 plots had no L. camara thickets, while the other 30 had dense thickets. The frequency of occurrence and abundance of seedlings were modelled as a function of dispersal mode (mammal, bird or mechanical) and affinities to forest habitats (dry forest, moist forest or ubiquitous) as well as presence or absence of dense L. camara thickets. Furthermore, frequency of occurrence and abundance of individual species were also compared between thickets and no L. camara. At the community level, L. camara density, dispersal mode and forest habitat affinities of species determined both frequency of occurrence and abundance of seedlings, with the abundance of dry-forest mammal-dispersed species and ubiquitous mechanically dispersed species being significantly lower under L. camara thickets. Phyllanthus emblica and Kydia calycina were found to be significantly less abundant under L. camara, whereas most other species were not affected by the presence of thickets. It was inferred that, by affecting the establishment of native tree seedlings, L. camara thickets could eventually alter the community composition of such forests.

Nest site characterization of sympatric hornbills in a tropical dry forest

Hornbills, among the largest and most threatened tropical frugivores, provide important seed dispersal services. Hornbill nest site characteristics are known primarily from wet tropical forests. Nests of the Indian grey hornbill Ocyceros birostris and Oriental pied hornbill Anthracoceros albirostris were characterized in a tropical dry forest. Despite A. albirostris being twice the size of O. birostris, few of the nest cavity attributes were different. A. albirostris nests were surrounded by higher proportion of mixed forest and lower sal forest compared to O. birostris. In this landscape, the larger A. albirostris may prefer to nest in sites with more food plants compared to the smaller O. birostris.

Differentiation between Neotropical rainforest, dry forest and savannah ecosystems by their modern pollen spectra and implications for the fossil pollen record

Accurate differentiation between tropical forest and savannah ecosystems in the fossil pollen record is hampered by the combination of: i) poor taxonomic resolution in pollen identification, and ii) the high species diversity of many lowland tropical families, i.e. with many different growth forms living in numerous environmental settings. These barriers to interpreting the fossil record hinder our understanding of the past distributions of different Neotropical ecosystems and consequently cloud our knowledge of past climatic, biodiversity and carbon storage patterns. Modern pollen studies facilitate an improved understanding of how ecosystems are represented by the pollen their plants produce and therefore aid interpretation of fossil pollen records. To understand how to differentiate ecosystems palynologically, it is essential that a consistent sampling method is used across ecosystems. However, to date, modern pollen studies from tropical South America have employed a variety of methodologies (e.g. pollen traps, moss polsters, soil samples). In this paper, we present the first modern pollen study from the Neotropics to examine the modern pollen rain from moist evergreen tropical forest (METF), semi-deciduous dry tropical forest (SDTF) and wooded savannah (cerradão) using a consistent sampling methodology (pollen traps). Pollen rain was sampled annually in September for the years 1999–2001 from within permanent vegetation study plots in, or near, the Noel Kempff Mercado National Park (NKMNP), Bolivia. Comparison of the modern pollen rain within these plots with detailed floristic inventories allowed estimates of the relative pollen productivity and dispersal for individual taxa to be made (% pollen/% vegetation or ‘p/v’). The applicability of these data to interpreting fossil records from lake sediments was then explored by comparison with pollen assemblages obtained from five lake surface samples.

Forest natural regeneration and biomass production after slash and burn in a seasonally dry forest in the Southern Brazilian Amazon

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

A new species of Symbra (Hymenoptera, Eurytomidae, Heimbrinae) from dry forest in Brazil and new occurrence records for other Heimbrinae

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Fluctuations in Richness and Abundance of Social Wasps During the Dry and Wet Seasons in Three Phyto-Physiognomies at the Tropical Dry Forest of Brazil

The social wasp nests were quantified in three different plant physiognomies (forested Caatinga, shrubby Caatinga, and agricultural systems) to analyze the effect of environmental seasonality and plant physiognomy on the richness, nest abundance, and species composition of social wasps in the region of tropical dry forest of Brazil. The forested Caatinga physiognomy had the greatest richness of species (S = 16), followed by shrubby Caatinga (S 13) and by agricultural system (S = 12). The first axis of detrended correspondence analysis (DCA) explained 67.8% of the variability and shows a gradient of the fauna from agricultural system and shrubby Caatinga to forested Caatinga. In the first axis, wet season scores were much higher than those for the dry season in forested Caatinga. The second axis explained 18.7% of the variability and shows a separation of samples collected during the wet or the dry periods in shrubby Caatinga. This separation was less evident in the agricultural system. Variations in nest abundance were more intense in arbustive caatinga (45% decrease in number of active nests in the dry period), moderate in forested Caatinga (24% decrease in number of active nests in the dry period), and low in agricultural systems (8% decrease in the dry period).

Preferential consumption of larger fruits of Piper arboreum (Piperaceae) by Carollia perspicillata (Phyllostomidae) in the Brazilian tropical dry forest

Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES)

Mating behaviour, nympho-imaginal development and description of a new Mesabolivar species (Araneae: Pholcidae) from the Brazilian dry forest

A new species of Mesabolivar is described from Brazilian forests: Mesabolivar delclaroi. The mating behaviour, postembryonic development, number of egg sacs and eggs, birth rate, number of instars, developmental time until adulthood, sex ratio and size of cephalothorax (per instar) were recorded. The sexual behaviour was described and categorized into four steps: courtship, pre-copulation, copulation and post-copulation. After hatching, individuals presented five instars until maturity. The mean number of eggs (42 +/- 16.6) and live births (31.5 +/- 3.4) of the first egg sac were significantly greater than that of a second one (23.8 +/- 3.8, and 19.25 +/- 3.9, respectively). The developmental time from birth to adulthood (130.8 +/- 9.6 days) did not differ significantly between egg sacs produced (128.61 +/- 11.1). The size of the cephalothorax did not differ among adults or between sexes. The sex ratio revealed a shift in favour of females (4: 3).

Assessing land use and land cover change in tropical dry forest or Northern Chinandega, Nicaragus from 1985 to 2011

The objective of this study is to gain a quantitative understanding of land use and land cover change (LULCC) that have occurred in a rural Nicaraguan municipality by analyzing Landsat 5 Thematic Mapper (TM) images. By comparing the potential extent of tropical dry forest (TDF) with Landsat 5 TM images, this study analyzes the loss of this forest type on a local level for the municipality of San Juan de Cinco Pinos (63.5 km2) in the Department of Chinandega. Change detection analysis shows where and how land use has changed from 1985 to the present. From 1985 to 2011, nearly 15% of the TDF in San Juan de Cinco Pinos was converted to other land uses. Of the 1434.2 ha of TDF that was present in 1985, 1223.64 ha remained in 2011. The deforestation is primarily a result of agricultural expansion and fuelwood extraction. If current rates of TDF deforestation continue, the municipality faces the prospect of losing its forest cover within the next few decades.

Regenerative morphological traits in a woody species community in Tumbesian tropical dry forest.

The study of functional morphological traits enables us to know fundamental aspects of the dynamics of plant communities in local and global habitats. Regenerative morphological traits play an important role in defining plant history and ecological behavior. Seed and fruit characteristics determine to a large extent the patterns for dispersal, germination, establishment and seedling recruitment a given species exhibits on its natural habitat. Despite their prominent role, seed and fruit traits have been poorly studied at the community level of woody plant species in neo-tropical dry forests. In the present study we aimed at i) evaluate the functional role of morphological traits of seeds, fruits and embryo in woody plant species; ii) determine which are the morphological patterns present in seeds collected from the community of woody species that occur in neo-tropical dry forests; and iii) compare woody plant species seed mass values comparatively between neo-tropical dry and tropical forests. To do so, mature seeds were collected from 79 plant species that occur in the Tumbesian forest of Southwest Ecuador. The studied species included the 42 and 37 most representative tree and shrubbery species of the Tumbesian forest respectively. A total of 18 morphological traits (seven quantitative and 11 qualitative) were measured and evaluated in the seeds, fruits and embryos of the selected species, and we compared the seeds mass with other forest types. Our results showed a huge heterogeneity among traits values in the studied species. Seed mass, volume and number were the traits that vary the most at the community level, i.e. seed length ranged from 1.3 to 39 mm, and seed width from 0.6 to 25 mm. Only six embryo types were found among the 79 plant species. In 40 % of the cases, fully developed inverted embryos with large and thick cotyledons to store considerable amount of nutrients were recorded. We concluded that highly variable and functionally complementary morphological traits occur among the studied woody plants of the dry Tumbesian forest. The latter favors a plethora of behavioral mechanisms to coexist among woody species of the dry forest in response to the environmental stress that is typical of arid areas.

Leaf Functional Traits and Forest Structure of Tropical Dry Forest Species Along a Rainfall Gradient in Florida and Puerto Rico

This study examined how different rainfall regimes affect a set of leaf functional traits related to plant stress and forest structure in tropical dry forest (TDF) species on limestone substrate. One hundred fifty eight individuals of four tree species were sampled in six ecological sites in south Florida and Puerto Rico, ranging in mean annual rainfall from 858 to 1933 mm yr-1. Leaf nitrogen content, specific leaf area (SLA), and N:P ratio of evergreen species, but not deciduous species, responded positively to increasing rainfall. Phosphorus content was unaffected in both groups. Canopy height and basal area reached maxima of 10.3 m and 31.4 m2 ha-1, respectively, at 1168 mm annual rainfall. Leaf traits reflected soil properties only to a small extent. This led us to the conclusion that water is a major limiting factor in TDF and some species that comprise TDF ecosystems are limited by nitrogen in limestone sites with less than ~1012 mm rainfall, but organismal, biological and/or abiotic forces other than rainfall control forest structure in moister sites.

Diversity of new root nodule bacteria from Erythrina velutina Willd., a native legume from the Caatinga dry forest (Northeastern Brazil).

This study aims to evaluate the phenotypical characteristics of bacterial isolates from mulungu (Erythrina velutina Willd.) nodules and determinate their Box-PCR fingerprinting. All bacteria were evaluated by the following phenotypic characteristics: growth rate, pH change, colony color and mucus production. The bacterial isolates able to re-nodulate the original host were also evaluated regarding its tolerance to increased salinity and different incubation temperatures, ability to growth using different carbon sources, intrinsic antibiotic resistance and ?in vitro? auxin biosynthesis. The molecular fingerprints were set up using the Box-PCR technique and the isolates were clustered by their profiles. Among the 22 bacterial isolates obtained, eight were able to re-nodulate the original host. Among the nodule inducing isolates, some were tolerant to 1% of NaCl and 39° C and all of them metabolized the maltose, fructose, glucose, sucrose and arabinose, were resistant to rifampicin and produced auxin. The bacteria showed low genetic similarity among them and reference strains, which indicates the great genetic variability of the isolates. The results of this work are the first reports about the bacterial isolates able to nodulate this species. A more deep study of these bacteria may reveal the existence of isolates tolerant to environmental stresses and suitable as a future mulungu inoculant.

Leaf flushing phenology and herbivory in a tropical dry deciduous forest, southern India

Patterns of leaf-flushing phenology of trees in relation to insect herbivore damage were studied at two sites in a seasonal tropical dry forest in Mudumalai, southern India, from April 1988 to August 1990. At both sites the trees began to flush leaves during the dry season, reaching a peak leaf-flushing phase before the onset of rains. Herbivorous insects emerged with the rains and attained a peak biomass during the wet months. Trees that flushed leaves later in the season suffered significantly higher damage by insects compared to those that flushed early or in synchrony during the peak flushing phase. Species whose leaves were endowed with physical defenses such as waxes suffered less damage than those not possessing such defenses. There was a positive association between the abundance of a species and leaf damage levels. These observations indicate that herbivory may have played a major role in moulding leaf flushing phenology in trees of the seasonal tropics.