Distribution and vegetation dynamics of humid savannas in Africa and Asia

Abstract. A review is presented on the literature about the distribution of savannas in humid climates in Africa and Asia and their vegetation dynamics. Sections are devoted to African lowland and montane savannas (the latter divided into southern, eastern, western and northern African), Madagascar, Indian subcontinent, SE Asia and New Guinea. It is concluded that the extension of savannas under humid climatic conditions and the relation to the distribution of forests is a function of cultivation, grazing by domestic and wild animals, present and previous climate, geomorphology and soil characteristics. Once established, savannas are often maintained by fires, both natural and man-made. Montane savannas are generally brought about by man's clearing, cultivation and burning. Fire is a stochastic variable; it creates an ecotone sensu stricto (an environmentally stochastic stress zone) at the forest/savanna border. On the other hand, if geomorphology and soil are the determinants, the transition between forest and savanna would have the character of an ecocline (a gradient zone) with fundamentally different conditions. In humid African lowland climates forests expand into savannas if the latter are not maintained by man. Whether forests also expand in less humid climates is disputed. In montane areas forest expansion may be delayed on degraded soils and when diaspores are lacking.     

Comparing and Combining Landsat Satellite Imagery and Participatory Data to Assess Land-Use and Land-Cover Changes in a Coastal Village in Papua New Guinea

In regions lacking socio-economic data, pairing satellite imagery with participatory information is essential for accurate land-use/cover (LULC) change assessments. At the village scale in Papua New Guinea we compare swidden LULC classifications using remote sensing analyses alone and analyses that combine participatory information and remotely sensed data. These participatory remote sensing (PRS) methods include participatory land-use mapping, household surveys, and validation of image analysis in combination with remotely sensed data. The classifications of the swidden area made using only remote sensing analysis show swidden areas are, on average, two and a half times larger than land managers reported for 1999 and 2011. Classifications made using only remote sensing analysis are homogeneous and lack discrimination among swidden plots, fallow land, and non-swidden vegetation. The information derived from PRS methods allows us to amend the remote sensing analysis and as a result swidden areas are more similar to actual swidden area found when ground-truthing. We conclude that PRS methods are needed to understand swidden system LULC complexities.     

Persistence of Swidden Cultivation in the Face of Globalization: A Case Study from Communities in Calakmul, Mexico

Over the last decades, political, economic and environmental pressures have encouraged changes from swidden to more intensive agricultural practices, resulting in the hypothesis that swidden cultivation systems are disappearing. In Calakmul, southeastern Mexico, communities decreased the area under milpa, the traditional maize swidden system, but a collapse did not occur. To document and explain the persistence of swidden we employ a variety of data: (1) 59 standardized household surveys from 2003 and 2010 in five villages, (2) in-depth interviews in one village, and (3) coupled human–environmental timelines in this same village. Droughts, hurricanes, and remittances were important drivers of decreases in milpa cultivation. Market crop profitability and conservation programs were also reported to affect the area under milpa. Off-farm employment and governmental transfers have tended to stabilize household economies and decrease dependency on agricultural production, but have also allowed households to maintain their milpas for subsistence and cultural reproduction. Findings in Calakmul point to the need to consider swidden as an evolving and active response to changing policy, economic, and environmental conditions.     

Vegetation history of the savanna corridor between the Guinean and the Congolian rain forest during the last 150,000 years

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

Guinean biodiversity at the edge: Rodents in forest patches of southern Mali

Southern Mali mainly belongs to the Sudanian savanna bioclimatic zone, but forest patches showing botanical affinities with Guinean humid forest remain as gallery forests or ravine forests. To characterize the rodent diversity of this area and check for the presence of some species of Guinean affinities in this group, rodent assemblages were sampled in four regions of southern Mali, using trapping and observational data in forest and surrounding habitats. Twenty-four species were recorded, comprising a representative sample of the expected overall diversity in this group according to rarefaction curves. Praomys rostratus was the dominant species in the most humid, closed lowland forest. Praomys daltoni was also present in this habitat type, being all the more abundant as habitat degradation was apparent. It became the dominant species in ravine forest on rocky substrate where P. rostratus was completely absent. In Sudanian savanna habitats and in herbaceous and cultivated areas, Mastomys erythroleucus dominated a diverse rodent community. A few species were found that testified for Guinean affinities of the most humid forest patches, especially in the extreme southeast of the country (region of Sikasso). Rodent assemblages of the Bafing and Mts Mandingues areas, in the western part of the study area, showed the highest similarity, in relation with environmental characteristics of this region representing an extension of the Fouta Djallon plateau in Guinea. The results obtained highlight the high biodiversity value of this forest-savanna mosaic, and provide new arguments in favour of the preservation of West African forest patches and their surrounding habitats.     

Beta diversity of frogs in the forests of New Guinea, Amazonia and Europe: contrasting tropical and temperate communities

Aim To test the hypothesis that animal communities within environmentally relatively uniform lowland forests are characterized by low beta diversity, both in tropical and in temperate areas. Location Lowland forests in the basins of the Sepik and Ramu rivers in New Guinea, the Amazon river in Bolivia, and the Elbe and Dyje rivers in the Czech Republic. Methods A network of 5–6 study sites spanning distances from 20–80 to 300–500 km in each study area was systematically surveyed for all frogs, using visual detection and call tracking. The community data were analysed for alpha and beta diversity. Results Local (alpha) diversity of frog communities was similar in the two tropical areas, New Guinea (mean ± SE of 22 ± 1.4 species per site) and Amazonia (24 ± 1.7 species), but was significantly lower in Europe (8 ± 0.8 species). In Amazonia, 36 of the total of 70 species were recorded from single sites. In contrast, widespread species dominated in Europe, whereas New Guinea exhibited an intermediate pattern with both local and widespread species well represented. The rate of species accumulation across different sites was lowest in Europe, intermediate in New Guinea and highest in Amazonia. The regional species diversity, expressed as the combined number of species from five study sites, was 1.5 times higher than the local species diversity at a single site in Europe, 2.0 times higher in New Guinea and 2.7 times higher in Amazonia. The proportion of species shared between communities decreased with geographic distance in New Guinea and Europe, but not in Amazonia. Main conclusions Frog communities in the lowland tropical rain forests of New Guinea and Amazonia had similar numbers of species, but differed in their beta diversity. More species in Amazonia had restricted distributions than in New Guinea. Both tropical areas had markedly higher alpha and beta diversity than the temperate area in Europe.     

Successful invasion of the neotropical species Piper aduncum in rain forests in Papua New Guinea

Abstract. Piper aduncum is a neotropical invasive species which has spread throughout Papua New Guinea over the past three decades. It has become a most successful alien woody plant in New Guinea, occurring from sea level up to 2000 m a.s.l. The species prefers initial stages of forest succession and is particularly common in recently abandoned gardens representative of a system of swidden agriculture. It often attains high cover, suppresses other pioneer species and becomes the absolute dominant species in these habitats. The species is now also spreading into naturally disturbed habitats far from direct human influence, such as natural tree‐fall gaps, landslides and frequently flooded stream banks. It has, however, never been found in a closed primary forest. The species germinates from faeces of mammal and bird species, and we conclude that dispersal through endozoochory contributes to this species’ extraordinary success in Papua New Guinea. A similar invasion behaviour has been documented over a large geographic area, from Malaysia to Fiji. Piper aduncum has attributes which are common amongst successful invasive species: (1) a large native geographic range; (2) aggressively colonizing disturbed habitats in its native area; (3) relatively small seeds; (4) a short juvenile period; (5) a large seed production every year.     

Contrasting phylogeographic signatures in two Australo‐Papuan bowerbird species complexes (Aves: Ailuroedus)

The Australo‐Papuan catbird genus Ailuroedus has a complex distribution and a contested taxonomy. Here, we integrate phylogenetic analysis of DNA data and morphology to study the group's biogeography and to re‐examine its taxonomy. We couple phylogeographic and abiotic data to examine differences between the major groups defined in our phylogenetic analysis. Our results are consistent with Ailuroedus catbirds being divided into two species complexes, one distributed in humid forests in the lowlands on New Guinea and another in comparably drier and colder forests mainly in mid‐mountains on New Guinea and Australia. Vicariant events during the Pliocene are surmised to have been the major force in shaping the contemporary phylogeographical signature of this genus. Several previously suggested vicariant events, such as fragmentation of xeric forests in Australia and the uplift of the central mountain range on New Guinea, are reinforced as important Pliocene barriers for tropical forest taxa in this region. Interaction between Pleistocene climatic fluctuations and differences in habitat requirements may explain a higher and more recent population structures in the mid‐mountain catbird complex and the lack of representatives from the lowland clade in the comparably drier Australia. Phylogeographical patterns in both catbird complexes, respectively, both comply and deviate from other lowland and mid‐mountain taxa in the region. This highlights that taxon‐specific properties, such as their historical spatial and ecological distributions, capacity to disperse and tolerance to habitat changes, affect the phylogeographical histories of organisms. Within both species complexes, the genetic differentiation between several geographically isolated populations was found to exceed those commonly observed for avian sister species. As these genetically distinct taxa also were found to be morphological diagnosable, we suggest a revised classification of the genus Ailuroedus, where we recognize three species within the lowland complex and seven species within the mid‐mountain complex.     

The Last Swiddens of Sarawak, Malaysia

Swidden cultivation was observed to be under pressure but still persisting in many areas in Sarawak, Malaysia around 2000–2003. Since then rapid development of smallholder oil palm and rubber, continuing urbanization and rural to urban migration appear to have considerably reduced the area under swidden cultivation. With the aim of understanding the extent and impacts of this development, 55 households in three communities that were interviewed in 2002–2003 were re-interviewed in 2011. In an area with rapid oil palm development, the households engaged in smallholder oil palm production have experienced considerable improvements in income and wealth whereas the other households have experienced more limited wealth increases or even a decline in income. Many households have decreased or abandoned cultivation of upland rice, which used to be the core of the swidden cultivation system, and the upland soils in the area are now dominated by oil palm. In another area, where no oil palm development has taken place because of a hydroelectric dam, upland rice fields under swidden cultivation have also decreased and new high yielding rubber is now being planted because of the favorable rubber prices. Moreover, tourism has in this area gained further economic importance and overtaken agriculture as the main economic activity of households. The demise of swidden has not yet occurred in Sarawak, but a continued decline has been observed. However, there is a possibility that the new smallholder oil palm and rubber may provide an opportunity for a new type of ‘productive fallow’ that will allow continued cultivation of upland rice on a small scale.     

Environmental Consequences of the Demise in Swidden Cultivation in Southeast Asia: Carbon Storage and Soil Quality

The effects of swidden cultivation on carbon storage and soil quality are outlined and compared to the effects of the intensified production systems that swidden systems of Southeast Asia transform into. Time-averaged aboveground carbon stocks decline by about 90% if the long fallow periods of traditional swidden cultivation are reduced to 4 years and by about 60% if swidden cultivation is converted to oil palm plantations. Stocks of soil organic carbon (SOC) in tree plantations are 0–40% lower than stocks in swidden cultivation, with the largest losses found in mechanically established oil palm plantations. Impacts of tree plantations on soil quality are to a large extent determined by management. Conversion of swiddening to continuous annual cropping systems brings about substantial losses of time-averaged aboveground carbon stocks, reductions of SOC stocks and generally leads to declining soil quality. Knowledge of carbon storage in belowground biomass of tree based systems of the tropics is sparse but failure to include this pool in carbon inventories may significantly underestimate the total biomass of the systems. Moreover, studies that consider the ecological reasons behind farmers’ land use decisions as well as spatial variability in biogeophysical and edaphological parameters are needed to evaluate the effects of the ongoing land use transitions in Southeast Asia.     

Periodicity of Wood Formation in Twigs of some Tropical Trees in Nigeria

The periodicity of wood formation was studied in twigs of treesgrowing in three ecological areas in Nigeria, namely lowlandrainforest, Southern Guinea savanna and mangrove swamp. Initiationof cambial activity was correlated with bud break, and wheremultiple vegetative bud formation occurred in the lowland rainforestand the Southern Guinea savanna, the number of growth ringscorresponded with the number of times the vegetative buds opened.In the mangrove swamp, although there may be a clear correlationbetween vegetative bud growth and wood formation in 1 to 2 year-oldtwigs of Drepanocarpus lunatus and Ormacarpum verrucosum therewas no such correlation in Dalbergia ecastaphyllum. Short axillarybranches and inflorescences which developed irregularly in plantsof the mangrove swamp induced multiple growth ring formation.In D. ecastaphyllum, the pattern of unfolding of young leavesseems to play a major role in the formation of multiple growthrings. In each of the areas multiple growth ring formation couldbe induced by injury. In Daniellia oliveri there was periodicdeposition of prismatic crystals. The crystals were depositedeach time wood formation stopped. In plants in which flowersopened before the opening of the vegetative buds, there wasinitiation of cambial activity by the opened flowers in thetwigs on which the flowers were borne.     

Lessons Learned about Research and Management: A Case Study from a Midwest Lowland Savanna,U.S.A.

Integrating research and practice is a fundamental challenge for restoration and conservation. Scientists recognize restoration as an opportunity to test basic ecological principles, and that core scientific practices such as establishing control plots, replication, monitoring, and data management are fundamental to learning from restoration and management. However, practical constraints inherent to many projects often work against fully integrating research and management. In 2005, we had the opportunity to evaluate a lowland savanna that had been undergoing restoration by The Nature Conservancy since 1997. We used the data generated as a case study to evaluate how the restoration process might have changed had scientists been involved from the outset. Prescribed fire is the primary tool used to restore upland savanna and was chosen by The Nature Conservancy to manage the lowland savanna. In open areas dominated by Reed canary grass (Phalaris arundinacea), fire was associated with reduced cover of this highly invasive species and an increase in native species richness. However, other results suggested that management options beyond the use of fire are needed to restore this lowland savanna. In retrospect, an approach that integrated the skills of researchers and managers at the outset of the study would have altered many of the restoration goals and processes. Pretreatment data, greater replication, and systematically varying treatments could have provided a more robust assessment of treatment effects. We recognize, however, that impediments in staffing, funding, and time were barriers to following this course of action.     

Local versus regional species richness in tropical insects: one lowland site compared with the island of New Guinea

1. The overlap in species composition of Cercopoidea (Aphrophoridae, Cercopidae, and Machaerotidae), Flatidae, and Ricaniidae between two data sets, an almost exhaustive census from 13 Ficus species and a sample from diverse vegetation in the same area, led to the estimate for local species richness of 111 (SE 11.5) species (45 species of Cercopoidea, 36 species of Flatidae, and 30 species of Ricaniidae) at a lowland rainforest site in New Guinea. 2. Samples restricted to 13 species of Ficus contained 66 species, i.e. 59% of the estimated local species richness. This high proportion probably results from the high proportion of polyphagous and tourist (transient) species in the Cercopoidea, Flatidae, and Ricaniidae. 3. The two largest museum collections of New Guinean insects contained 327 species of Cercopoidea from New Guinea, including 23 of the 34 species collected in the field samples. This overlap led to the estimate of 483 (SE 97.2) species of Cercopoidea present in New Guinea. 4. The species found in the field samples were also 2.6 times more likely to be found in the museum collection than other species. This sampling bias can be due to a positive correlation between species local abundance and geographic distribution and/or similar patterns of species abundance at different sites. 5. The estimate of species richness of Cercopoidea in New Guinea increased to 1222 species when corrected for this sampling bias. Thus, only 4% of the New Guinean species were present locally, in the study area. Such high beta diversity is probably a consequence of the exceptional habitat and vegetation diversity in New Guinea, as well as its complex geological history.     

Who Counts? Demography of Swidden Cultivators in Southeast Asia

Swidden cultivators are often found as a distinct category of farmers in the literature, but rarely appear in population censuses or other national and regional classifications. This has led to a worldwide confusion on how many people are dependent on this form of agriculture. The most often cited number of 200–300 million dates back to the early 1970s, but the source is obscure. We assess available, published data from nine countries in Southeast Asia and conclude that on this basis it is not possible to provide a firm estimate of the number of swidden cultivators in the region. A conservative range of 14–34 million people engaged in swidden cultivation in the region is suggested, however. We argue that along with improved knowledge of swidden livelihoods, there is an urgent need to develop techniques that will allow for better estimates of swidden populations in order to secure appropriate rural development and poverty reduction in swidden areas.     

Restoration of Midwestern U.S. Savannas: One Size Does Not Fit All

Lowland savannas are a rare variant of Midwestern United States savanna occurring on alluvial soils, for which reference information is sparse. To evaluate the appropriateness of using upland savanna as a surrogate source of reference information for lowland savanna, we studied a pre‐Euro‐American lowland savanna using original U.S. Public Land Survey data and other historical records. Historical vegetation was reconstructed and compared among upland savannas, lowland savannas, and lowland forests; we also evaluated potential disturbance dynamics maintaining these systems. We found that all three communities were dominated by members of the genus Quercus but also had extensive representation by many other tree species, especially notable for savannas in this region. There were no clear size–density relationships for species in the genus Quercus, indicating that these historical savannas were not characterized exclusively by large, scattered oak trees but rather by trees of many oak species and nonoak species in a wide range of size classes. Both upland and lowland savannas also contained a substantial shrub component. We found no evidence that lowland savannas were maintained by flooding, although the uneven‐aged canopy structure suggested that periodic disturbance occurred. Restoration of lowland savanna in this region should include provisions for maintaining nonoak species and shrubs, with disturbance timed to maintain an uneven‐aged canopy structure. Although the appropriateness of historical data in the face of climate change may be questionable, in this region, a warmer climate may actually help promote the “oak parkland” that was present from 8,000 BP up to Euro‐American settlement.     

Dynamics of energy and nutrient concentration and construction cost in a native and two alien C4 grasses from two neotropical savannas

In Venezuela, the alien grasses Melinis minutiflora Beauv. and Hyparrhenia rufa (Nees.) Stapf tend to displace the native savanna plant community dominated by Trachypogon plumosus (Humb. and Bonpl.) Nees. This occurs in either relatively wetter and fertile highland savannas or in drier and less fertile lowland savannas. Although the native and aliens are perennial C₄ grasses, higher net assimilation leaf biomass per plant and germination rate of the latter are some causes for their higher growth rates and for their competitive success. The objective of this study is to compare seasonal tissue energy, N, P and K concentrations and the calculated construction costs (CC) between the native grass and either one of the alien grasses from lowland and highland savannas. We predict that, in order to out-compete native plants, alien grasses should be more efficient in resource use as evidenced by lower tissue energy and nutrient concentrations and CC.Tissue energy and nutrient concentration were measured throughout the year and compared between M. minutiflora and the co-occurring local population of T. plumosus in a highland savanna and between H. rufa and its neighbor local population of T. plumosus in a lowland savanna. CC was calculated from energy, N and ash concentrations considering ammonium as the sole N source. Differences between co-occurring species, T. plumosus populations, seasons, and organs were analyzed with ANOVA.Highland and lowland grasses differed in concentration and allocation of energy and nutrients whereas the differences between alien and native grasses were specific for each pair considered. Highland grasses had higher energy, N, P and CC than lowland grasses. These variables were always lowest in the culms. In the more stressed lowland site, tissue energy and nutrient concentrations decreased significantly during the dry season except in the roots of both grasses which had the highest energy and nutrients concentrations during the drought. This seasonal response was more marked in the local lowland population of T. plumosus in which maximum CC alternated seasonally between leaves and roots. Energy and nutrient concentrations and CC were the lowest in H. rufa. In the lowland savannas, the higher efficiency of resource use in the invader grass contributes to its higher competitive success through increased growth rate. In the highlands, overall tissue energy concentration and CC, but not N nor P concentration, were lower in the fast growing M. minutiflora but seasonal differences were lacking. The higher leaf CC in T. plumosus can be attributed to the higher proportion of sclerenchyma tissue which is more expensive to construct. Considering CC, both fast growing alien grasses are more efficient in resource use than the co-occurring native grass. However, the role of CC explaining the competitive success of the former, through higher growth rates, is more evident in the more stressful environment of the lowland savanna.     

Saving Slash-and-Burn to Save Biodiversity

Several prominent articles have recently revived the debate on how to advance and reconcile two pressing global issues: conservation of biodiversity, and food production for an increasing human population. These discussions contrast a ‘land-sparing/intensive agriculture’ strategy with a ‘biodiversity-friendly’ agriculture approach. We propose that swidden or shifting cultivation should be an important component of the latter approach in the tropics because many swidden systems maintain very high levels of biodiversity while providing livelihood for populations in tropical forest areas worldwide. We suggest further that when many swidden systems are viewed without prejudice and in broader spatial and longer temporal perspectives, the conservationist aspects of the systems become evident.     

Climate, life forms and species distribution on the Loma Montane grassland, Sierra Leone

The Loma Mountains (2090 m), situated in lowland south Guinea savanna in Sierra Leone, are shown to have a cooler and more mesic climate than the savanna region. A temperature lapse rate of 7.5^o C/1000 m is calculated using means for the dry, dry/wet, wet and wet/dry seasons of the year. In the montane grassland the life forms of the species are predominantly hemicryptophytic, and the biological spectrum is closer to that of temperate grasslands than to the savanna type, although the adverse conditions are still dry-season drought and fires. The phytogeographical distribution of the flora indicates a 44% widespread, truly montane to montane/savanna element and a 43% West African endemic element. In the Guinean highlands, the Loma/Tingi group shows about 10% disjunction of its grassland flora with other African mountains outside West Africa.     

Agricultural intensification in a Philippine frontier community: Impact on labor efficiency and farm diversity

There continues to be much debate in anthropology concerning the mechanism by which agricultural intensification takes place and its impact on labor efficiency, farm diversity, and quality of diet. A major reason for this lack of consensus is the paucity of data from case studies that focus on specific agricultural systems at the point of transition from extensive to intensive methods of cultivation. Research in a frontier community in the Philippines, where farmers are making the shift from swidden cultivation to small-scale irrigated rice production, indicates that intensification does not necessarily result in lower efficiency or a decline in dietary standards. Rather, farmers faced with growing population pressure and an unproductive short fallow swidden system have been motivated to adopt irrigation because it increases the efficiency of their labor while maintaining a reliable and diverse farming system.     

Environmental Consequences of the Demise in Swidden Cultivation in Montane Mainland Southeast Asia: Hydrology and Geomorphology

The hydrological and geomorphological impacts of traditional swidden cultivation in Montane Mainland Southeast Asia are virtually inconsequential, whereas the impacts associated with intensified replacement agricultural systems are often much more substantial. Negative perceptions toward swiddening in general by governments in the region beginning half a decade ago have largely been based on cases of forest conversion and land degradation associated with (a) intensified swidden systems, characterized by shortened fallow and extended cropping periods and/or (b) the widespread cultivation of opium for cash after the Second World War. Neither of these practices should be viewed as traditional, subsistence-based swiddening. Other types of intensive agriculture systems are now replacing swiddening throughout the region, including semi-permanent and permanent cash cropping, monoculture plantations, and greenhouse complexes. The negative impacts associated with these systems include changes in streamflow response, increased surface erosion, a higher probability of landslides, and the declination in stream water quality. Unlike the case for traditional swiddening, these impacts result because of several factors: (1) large portions of upland catchments are cultivated simultaneously; (2) accelerated hydraulic and tillage erosion occurs on plots that are cultivated repetitively with limited or no fallowing to allow recovery of key soil properties, including infiltration; (3) concentrated overland flow and erosion sources are often directly connected with the stream network; (4) root strength is reduced on permanently converted hillslopes; (5) surface and ground water extraction is frequently used for irrigation; and (6) and pesticides and herbicides are used. Furthermore, the commercial success of these systems relies on the existence of dense networks of roads, which are linear landscape features renowned for disrupting hydrological and geomorphological systems. A new conservation focus is needed to reduce the impacts of these intensified upland agricultural practices.