Interactive influence of rainfall manipulation and livestock grazing on species diversity of the herbaceous layer community in a humid savannah in Kenya

Changes in rainfall regime and grazing pressure affect vegetation composition and diversity with ecological implications for savannahs. The savannah in East Africa has experienced increased livestock grazing and rainfall variability but the impacts associated with those changes on the herbaceous layer have rarely been documented. We investigated the effect of livestock grazing, rainfall manipulation and their interaction on the composition and diversity of the herbaceous community in the savannah for two years in Lambwe, Kenya. Rainfall manipulation plots were set up for vegetation sampling;these plots received either 50% more or 50% less rainfall than control plots. Simpson’s diversity and Bergere Parker indices were used to determine diversity changes and dominance respectively. The frequency of species was used to compute their abundance and their life forms as determined from the literature. Grazing significantly increased species diversity through suppression of dominant species. Rainfall manipulation had no significant impact on plant diversity in fenced plots, but rainfall reduction significantly reduced diversity in grazed plots. In contrast, rainfall manipulation had no impact on dominance in either fenced or grazed plots. The interaction of grazing and rainfall manipulation is complex and will require additional survey campaigns to create a complete picture of the implications for savannah structure and composition.     

Floristic diversity of Mexican seasonally dry tropical forests

Studies of the variation in tropical plant species diversity and itsrelationship with environmental factors are largely based on research intropical moist/wet forests. Seasonally dry tropical forests (SDTFs), incontrast, have been poorly investigated. In this paper we present data from 20Mexican SDTF sites sampled to describe the magnitude of floristic diversity inthese forests and to address the following questions: (i) to what extent isspecies diversity related to rainfall? (ii) Are there other climatic variablesthat explain variation in species diversity in SDTFs? (iii) How does speciesidentity vary spatially (species turnover) within the country? We found thatspecies diversity was consistently greater (a ca. twofold difference) than wouldbe expected according to the sites' precipitation. Rainfall did notsignificantly explain the variation in species diversity. Likewise, the numberof dry and wet months per year was unrelated to species diversity. In contrast,a simple measure of potential evapotranspiration (Thornthwaite's index)significantly explained the variation in species diversity. In addition to thegreat diversity of species per site (local diversity), species turnover wasconsiderable: of a total of 917 sampled species, 72% were present only in asingle site and the average similarity (Sorensen's index) among sites wasonly 9%. These aspects of floristic diversity and the high deforestation ratesof these forests in Mexico indicate that conservation efforts should be directedto tropical forests growing in locations of low and seasonal rainfall.     

Influence of grazing and soil conditions on secondary savanna vegetation in India

Abstract. Savanna vegetation and pertinent soil features were studied on 43 sites in a dry tropical forest region of India. Grazing intensity ranged from 0.68 to 0.98. Soil moisture was positively related to the proportion of fine soil particles (< 0.1 mm), and the latter decreased while the proportion of coarse particles (2.0-0.5 mm) increased with increasing grazing intensity. Canopy biomass ranged from 28 to 104 g/m² in grazed communities and from 230 to 337 g /m² in ungrazed communities and was positively related to vegetation cover which ranged between 30–72 % in grazed and 68 - 91 % in ungrazed communities. Vegetation cover was negatively related to grazing intensity. Species richness and diversity were highest at low grazing intensity. Using community coefficients and Detrended Correspondence Analysis, the grazed stands were clustered into six and the ungrazed ones into three communities. The grazed communities were recognised as degradation stages and the ungrazed ones as recovery stages. Only five grass species, in various combinations were able to dominate in one of the different stages. Evidently the harsh climatic conditions (high temperatures, high variability in rainfall and a long dry period) in the region permit only a few species already adapted to these conditions to participate in the succession.     

Stocks and dynamics of carbon in trees across a rainfall gradient in a tropical savanna

In this study, systematic variation in tree morphology across a rainfall gradient in Australia's tropical savanna biome and its implications for carbon stocks and dynamics were quantified. The aim was to support efforts to manage fire regimes to increase vegetative carbon stocks as a greenhouse gas mitigation strategy. The height of trees for a given trunk diameter declines with decreasing rainfall from 2000 to 300 mm and increasing dry season length across the Australian savanna biome. It is likely that increasing dry season length is the main driver of this decline rather declining rainfall per se. By taking account of the response of total basal area to rainfall and soil type, stand structure, and tree height and diameter relationships, the carbon stocks in live trees were estimated to decline from about 34 t ha^?1 in the wetter savannas to 6 t ha^?1 in the drier savannas. These values are broadly consistent with field‐based estimates. Because of the declining ratio of height to trunk diameter, trees of a given diameter in drier regions will be more likely to be killed by fires of a given intensity than trees in wetter regions. Thus single fires of given intensity are likely to have a greater proportionate impact on live tree carbon stock in drier savannas, but a much greater absolute impact in wetter savannas due to the greater total carbon stock. Projected decreases in early wet season rainfall under climate change scenarios, despite projections of little change in total precipitation in northern Australia, may lead to decreased carbon stock in live trees through two mechanisms: a reduction in total basal area and decreases in tree height for given trunk diameters.     

Ecology and diversity of leaf litter fungi during early-stage decomposition in a seasonally dry tropical forest

Leaf litter samples of 12 dicotyledonous tree species (belonging to eight families) growing in a dry tropical forest and in early stages of decomposition were studied for the presence of litter fungi. Equal-sized segments of the leaves incubated in moist chambers were observed every day for 30 d for the presence of fungi. Invariably, the fungal assemblage on the litter of each tree species was dominated by a given fungal species. The diversity of fungi present in the litter varied with the tree species although many species of fungi occurred in the litter of all 12 species. A Pestalotiopsis species dominated the litter fungal assemblage of five trees and was common in the litter of all tree species. The present study and earlier studies from our lab indicate that fungi have evolved traits such as thermotolerant spores, ability to utilize toxic furaldehydes, ability to produce cell wall destructuring enzymes and an endophyte-litter fungus life style to survive and establish themselves in fire-prone forests such as the one studied here. This study shows that in the dry tropical forest, the leaf litter fungal assemblage is governed more by the environment than by the plant species.     

Dynamics of viable nitrifier community, N-mineralization and nitrification in seasonally dry tropical forests and savanna

The study was conducted in Vindhyan region, to assess the N-mineralization, nitrification and size of viable community of ammonium- and nitrite-oxidizing bacteria as affected by different sites and seasons. Six different ecosystems (four forests and two savannas), which differ in terms of topography, vegetation and moisture status, were selected for the present study. The soils of the study sites differ significantly in its physico-chemical properties. The savanna site had significantly higher pH (7.2), bulk density (1.37 g cm(-3)) and silt content (67.80%) but lower water holding capacity (1.37%), total-C (16,356 microg g(-1) dry soil), N (1090 microg g(-1) dry soil) and P (213 microg g(-1) dry soil) than forest sites. The soil moisture content, N-mineralization, nitrification rates and numbers of ammonium- and nitrite-oxidizing bacteria were highest in the wet season and lowest in dry season, while the size of mineral-N (NH4(+)-N and NO3(-)-N) showed a reverse trend at the sites. The N-mineralization, nitrification and nitrifier population size differ significantly across the site and season. The numbers of free-living cells of ammonium- and nitrite-oxidizing bacteria were significantly related to each other and to N-mineralization, nitrification, soil moisture and mineral-N components. The N-mineralization, nitrification and the viable number of nitrifying cells were consistently higher for forest soils compared to savanna sites. It was concluded that soil microbial process (N-mineralization and nitrification) and nitrifier population size were dependent on site topography, vegetation cover and soil moisture status.     

Long-term dynamics of a tropical savanna bird community

Complete two-month censuses of diurnal raptors and strip transect counts of other bird species were compared between 1968–1972 and 1996 to assess changes in the bird community of a 2700ha guinean savanna in Ivory Coast. Among 117 non-raptor species recorded at least once on the transects, 25 mostly uncommon species were absent in 1996 (but only two of them were likely to be extinct in the study area), 4 were first seen in 1996, 14 decreased and 4 increased significantly. All six large forest raptors and resident savanna eagles disappeared before the mid-1980s. Among the 12 remaining breeding raptors, 4 species increased (including the 2 seasonal migrants), 4 species remained stable (including the 2 forest accipiters) and 4 species decreased, but not significantly (including the 2 largest taxa). All 14 non-breeding raptors (Palearctic and African migrants) remained stable or increased. With on average 63 breeding pairs per 1000ha, this area still supports one of the highest densities of diurnal raptors ever recorded. The geographic distributions, seasonal movements and ecological requirements of species involved suggest the most likely origin of specific changes observed. They were attributed, in roughly similar proportion, to: (i) decreasing rainfall (southward shifts leading to appearance or increase of some resident or migrant species); (ii) deforestation, agricultural development and hunting pressure around the reserve (extinction of larger species following habitat fragmentation or degradation, increase of some species); and (iii) habitat modifications inside the reserve (increasing tree cover, invasion of dense thickets, illegal exploitation of palms). The overall community of this small protected area, however, exhibited a remarkable stability during the last 30 years, in spite of dramatic habitat changes outside its limits.     

Plant functional assembly is mediated by rainfall and soil conditions in a seasonally dry tropical forest

Understanding how species assembly is influenced by the interplay of climate, local environmental conditions and human-caused disturbances remains a central question in ecology and conservation. Here, we assess how plant species abundance is determined by combinations of functional traits (ecological strategies) and interacting gradients of rainfall, soil conditions (fertility and field capacity) and chronic anthropogenic disturbance in a Caatinga dry tropical forest, Brazil. We tested for trait–environment relationships using multivariate methods (RLQ) accounting for groups of species sharing similar responses to gradients and similar expression of multiple traits (i.e. response groups). Overall, species’ abundances changed predictably in response to rainfall and soil fertility, and were mediated by functional traits, i.e. species with particular trait combinations tended to respond similarly to multifactorial conditions. Briefly, three ecological strategies emerged: species with low wood density and soft (i.e. lower dry matter content), thick leaves converged into a trait syndrome characterizing a drought-avoidance strategy through water storage. They were particularly abundant under extremely low precipitation and relatively high soil field capacity. Under conditions of increasing rainfall and decreasing soil field capacity, species with high wood density were favored, consistent with a drought-tolerance strategy. However, these species fell into two groups relative to leaf-investment: more conservative leaves (low SLA) on relatively fertile soils vs. thinner and softer (i.e. high SLA) leaves on unfertile soils. In seasonally dry tropical forests, low SLA on relatively fertile soils may represent a water conservation strategy. Unexpectedly, no ecological strategy emerged in response to disturbance. The patterns we uncovered help to understand the interplay between precipitation, soil fertility and anthropogenic disturbance in plant species filtering in seasonally dry tropical forests. Moreover, our results underline that impacts of future climate change will depend on how rainfall patterns covary with finer-scale environmental factors such as soil fertility and field capacity.     

The Mexican tropical deciduous forest of Baja California Sur: a floristic and structural approach

A quantitative study is presented of the tropical deciduous forest located in the Sierra de La Laguna in the southern part of the peninsula of Baja California, Mexico with data on structure, species composition, diversity, density, and abundance of perennial plants.4 study plots were selected to represent the predominant geomorphologic units, and to include topographic and climatic variations reflected by the distribution of this vegetation on the lowlands of slopes facing the Gulf of California and the Pacific Ocean.25 families containing 67 perennial species were found on the lowlands, with Leguminosae, Cactaceae, and Euphorbiaceae best represented. A high family diversity was found in the plots, but there was a low number of species per family. Dissimilarities between sites were found to be reflected significantly in growth-form abundances as well as in structural features and species diversity. Results show that the xeric environment, the low number of species, and the high incidence of dominant shrub species confer the vegetation of the lowlands simpler structural traits than those described for other tropical dry forests.     

Herbivory effects on saplings are influenced by nutrients and grass competition in a humid South African savanna

Woody plant encroachment is a common consequence of disturbance in savannas. Grazers and browsers interfere with sapling establishment dynamics by direct consumption of plant tissue, changing soil nutrient status (through fertilization and trampling) and grass competition. Studies evaluating the effects of herbivory on sapling establishment have mostly been extrapolated from single species. In a controlled field experiment, we studied the effects of clipping (simulating grazing and browsing), nutrients, grass competition, and their interactive effects on sapling survival and growth of four dominant humid and four dominant mesic savanna species. We conducted this experiment in a humid South African savanna. We found no effects on sapling survival by the treatments provided. However, clipped saplings of all species increased their investment in relative growth rate of stem length (RGR_L). Clipping had a greater negative impact on relative growth rate of more humid than mesic species in terms of stem diameter (RGR_D), total dry biomass and proportion of leaf biomass. Nutrients had a positive effect on the RGR_L and sapling biomass of three mesic species. Positive effects of nutrients on RGR_L of one humid and two mesic species were observed in their clipped saplings only. Grass competition had a strong negative impact on all growth parameters measured. Clipped saplings of one humid and two mesic species had lower RGR_L with grass competition whereas intact saplings showed no significant response. After clipping, humid savanna species were more vulnerable to grass competition than mesic species, with reduced ability to use nutrients. In conclusion, herbivory increases sapling vulnerability to grass competition, with humid species being more susceptible than mesic species, indicating that woody-plant control strategies are more likely to be effective in humid savannas.     

Cultivation impacts soil microbial dynamics in dry tropical forest ecosystem in India

We studied for two years the seasonal changes in plant available nitrate and ammonium nitrogen (N), nitrification, N-mineralization, microbial biomass carbon (MBC), nitrogen (MBN) and phosphorus (MBP) in two forest and three cropland sites, derived from a tropical forest ecosystem of India. Results indicated that seasonal values of nitrate N, ammonium N and phosphate P ranged from 7.33–12.99, 5.1–10.22 and 4.0–7.8 μg g^?1 in forest and 4.13–9.26, 9.35–14.46 and 2.8–5.8 μg g^?1 in cropland ecosystems, respectively, with maximum values in summer and minimum in rainy seasons. Nitrification and N-mineralization values varied from 6–28 and 4–26 μg g^?1 mo^?1 in forest and 3–14 μg g^?1 mo^?1 and 4–17 μg g^?1 mo^?1 in cropland, with maximum values in rainy season and minimum in summer season.MBC, MBN MBP ranged from 393–753, 34–80 and 16–36 μg g^?1 in forests and 186–414, 21–41 and 11–22 μg g^?1 in croplands, being maximum in summer and minimum in rainy seasons. There was gradual increase in the values of inorganic N, nitrification, N-mineralization and MBC, MBN and MBP along the age of cropland. Analysis of variance indicated significant difference in the concentration of inorganic N, nitrification and N-mineralization and MBC, MBN and MBP due to sites and seasons.Cultivation caused decline in the mean annual organic C, N and P by 42%, 29% and 13%. The values of nitrate N were decreased by 23–38%, while ammonium N was increased by 39–74%. Nitrification and N-mineralization values were reduced by 39–63% and 40–60%, respectively. Microbial C, N and P were reduced by 44–54%, 41–50% and 28–44%, respectively. Nonetheless, the contribution of soil microbial biomass reflected in total N was enhanced from 4.76% in forest to 5.03% in cropland ecosystem. Enhancement of plant available ammonium-N and microbial contribution in total N are an indicator of natural conserving mechanism to check the nitrogen loss from the nutrient poor agro-ecosystem.     

Structure and seasonal dynamics of humid tropical grasslands in Meghalaya,India

Abstract. Four humid grassland communities at three different locations in Meghalaya, India were analysed during 1988 and 1989 for species and life-form composition, diversity and dominance in relation to altitude, soil and prevailing disturbances. Due to the adverse interactive influences of exceptionally high annual rainfall (> 10 000 mm), topography and human interference on soil fertility, the grassland at Cherrapunji, at 1300 m altitude, had a low species diversity (H'= 1.74) and was dominated by three perennial grass species. Similar grasslands, at both higher and lower altitudes on fertile soil and with lower rainfall (ca. 2000 mm), showed higher diversity values (H'= 2.28 at Burnihat and 2.31 at Upper Shillong). The proportion of perennial species and chamaephytes increased with elevation. At the high altitude site a grassland under short-term protection from fires and grazing had a higher species richness, density and basal cover than an unprotected grassland. All grasslands show a clear seasonality, albeit with different patterns, with a maximum in density and basal cover in August. The differences in structure and seasonality are discussed in terms of different levels of stress.     

Decadal dynamics of tree cover in an Australian tropical savanna

Spatio‐temporal variation in tropical savanna tree cover remains poorly understood. We aimed to quantify the drivers of tree cover in tropical mesic savannas in Kakadu National Park by relating changes in tree cover over 40 years to: mean annual rainfall, fire activity, initial tree cover and prior changes in tree cover. Aerial photography, acquired in 1964, 1984 and 2004, was obtained for fifty sites in Kakadu that spanned a rainfall gradient from approximately 1200 to 1600 mm. The remotely sensed estimates of tree cover were validated via field survey. Linear mixed effects modelling and multi‐model inference were used to assess the strength and form of the relationships between tree cover and predictor variables. Over the 40 years, tree cover across these savannas increased on average by 4.94 ± 0.88%, but was spatio‐temporally variable. Tree cover showed a positive albeit weak trend across the rainfall gradient. The strength of this positive relationship varied over the three measurement times, and this suggests that other factors are important in controlling tree cover. Tree cover was positively related to prior tree cover, and negatively correlated with fire activity. Over 20 years tree cover was more likely to increase if (i) tree cover was initially low or (ii) had decreased in the previous 20‐year interval or (iii) there had been fewer fires. Across the examined rainfall gradient, the greater variability in fire activity and inherently higher average tree cover at the wetter latitudes resulted in greater dynamism of tree cover compared with the drier latitudes. This is consistent with savanna tree cover being determined by interactions between mean annual rainfall, tree competition and frequent fire in these tropical mesic savannas.     

Water uptake and transport in lianas and co-occurring trees of a seasonally dry tropical forest

Water uptake and transport were studied in eight liana species in a seasonally dry tropical forest on Barro Colorado Island, Panama. Stable hydrogen isotope composition (D) of xylem and soil water, soil volumetric water content (_v), and basal sap flow were measured during the 1997 and 1998 dry seasons. Sap flow of several neighboring trees was measured to assess differences between lianas and trees in magnitudes and patterns of daily sap flow. Little seasonal change in _v was observed at 90–120 cm depth in both years. Mean soil water D during the dry season was –19 at 0–30 cm, –34 at 30–60 cm, and –50 at 90–120 cm. Average values of xylem D among the liana species ranged from –28 to –44 during the middle of the dry season, suggesting that water uptake was restricted to intermediate soil layers (30–60 cm). By the end of the dry season, all species exhibited more negative xylem D values (–41 to –62), suggesting that they shifted to deeper water sources. Maximum sap flux density in co-occurring lianas and trees were comparable at similar stem diameter (DBH). Furthermore, lianas and trees conformed to the same linear relationship between daily sap flow and DBH. Our observations that lianas tap shallow sources of soil water at the beginning of the dry season and that sap flow is similar in lianas and trees of equivalent stem diameter do not support the common assumptions that lianas rely primarily on deep soil water and that they have higher rates of sap flow than co-occurring trees of similar stem size.     

Influence of anthropogenic disturbance on bird communities in a tropical dry forest: role of vegetation structure

A study was carried out in Sariska Tiger Reserve in India to investigate the effects of anthropogenic disturbance caused by biomass extraction on the bird communities of tropical dry forests. The study was based on comparisons of the avifaunal community as well as vegetation structure between strictly protected ('undisturbed') and intensively used ('disturbed') sites that were demarcated a priori on the basis of disturbance indicators. There was no significant difference in the number of recorded species and bird abundance between disturbed and undisturbed sites. However, bird species diversity was significantly lower in disturbed sites. Bird species composition was found to differ significantly between disturbed and undisturbed sites and was associated with the measured disturbance indicators. Changes in bird species composition occurred because of seven of 26 locally abundant bird species (26.9%) responding significantly to the disturbance regime. All the affected bird species are primarily insectivorous. Bird species composition was significantly related to six vegetation structural variables, including two that were significantly altered by disturbance. Changes in vegetation structure accounted for all the changes in bird species composition caused by disturbance. However, vegetation structure had additional effects on bird species composition besides those caused simply by disturbance. Thus, our study indicates that forest use in the form of chronic biomass extraction can have significant effects upon bird diversity and species composition of tropical dry forest. There is a need to retain a proportion of natural ecosystems as inviolate if the full complement of biodiversity is to be conserved.     

Germinable soil seed banks in a tropical savanna: seasonal dynamics and effects of fire.

Abstract The germinable soil seed bank of a tropical eucalypt savanna of north‐eastern Australia was found to be dominated by grasses and forbs, with seed bank density ranging from 58 to 792 seeds per square metre, from a total of 53 species. Late dry season fires and the fire‐related cues, heat shock and smoke, broke the seed dormancy of a range of tropical savanna species. Heat shock promoted the germination of the species groups natives, exotics, subshrubs, ephemeral and twining perennial forbs, and the common species Indigofera hirsuta, Pycnospora lutescens and Triumfetta rhomboidea. Exposure to smoke at ambient temperature promoted germination from the soil seed bank of the species groups combined natives, upright perennial forbs and grasses, as well as the common grasses Digitaria breviglumis and Heteropogon triticeus. The germinable soil seed bank varied seasonally, increasing from the mid wet season (February) and early dry season (May) to a maximum in the late dry season (October). The effect of recent fire history on soil seed bank dynamics was limited to the immediate release of some seed from dormancy; a reduction in seed densities of subshrubs and monocots, other than grasses, in recently burnt savanna; and enhanced seed density of the ephemeral I. hirsuta in the year following fire. The seed banks of most savanna species were replenished in the year following burning.     

Biomass and carbon accumulation in a fire chronosequence of a seasonally dry tropical forest

Seasonally dry tropical forests (SDTF) are a widely distributed vegetation type in the tropics, characterized by seasonal rainfall with several months of drought when they are subject to fire. This study is one of the first attempts to quantify above- and belowground biomass (AGB and BGB) and above- and belowground carbon (AGC and BGC) pools to calculate their recovery after fire, using a chronosequence approach (six forests that ranged form 1 to 29 years after fire and mature forest). We quantified AGB and AGC pools of trees, lianas, palms, and seedlings, and BGB and BGC pools (Oi, Oe, Oa soil horizons, and fine roots). Total AGC ranged from 0.05 to nearly 72 Mg C ha⁻¹, BGC from 21.6 to nearly 85 Mg C ha⁻¹, and total ecosystem carbon from 21.7 to 153.5 Mg C ha⁻¹; all these pools increased with forest age. Nearly 50% of the total ecosystem carbon was stored in the Oa horizon of mature forests, and up to 90% was stored in the Oa-horizon of early successional SDTF stands. The soils were shallow with a depth of <20 cm at the study site. To recover values similar to mature forests, BGC and BGB required <19 years with accumulation rates greater than 20 Mg C ha⁻¹ yr⁻¹, while AGB required 80 years with accumulation rates nearly 2.5 Mg C ha⁻¹ yr⁻¹. Total ecosystem biomass and carbon required 70 and 50 years, respectively, to recover values similar to mature forests. When belowground pools are not included in the calculation of total ecosystem biomass or carbon recovery, we estimated an overestimation of 10 and 30 years, respectively.