A species-poor tropical sedge community: Afrotrilepis pilosa mats on inselbergs in West Africa

The mat-forming poikilohydric sedge Afrotrilepis pilosa (Cyperaceae) occurs as the dominant member of characteristic stands on West African inselbergs. The species composition of this community has been examined in the Ivory Coast along a latitudinal gradient from the savanna to the rainforest region. Inselbergs consisting of Precambrian granites and gneisses, are prominent landscape elements characterized by large areas of exposed rock. Monocotyledonous mats are one of the most conspicuous communities of this ecosystem. Floristically, the mats are poor in higher plant species with a total of only 15 species (mostly therophytes) recorded in 110 relevés. Species diversity is likewise low, indicating extremely stressful environmental conditions. A major factor affecting plant colonization is the pronounced short-term variation in water-availability, even in the more humid regions in the south of the country. Afrotrilepis pilosa is highly competitive under these conditions, and the dense stands leave only little space for other species to become established. Longevity (attaining an age of several hundred years) and the ability, to spread radially to form almost monospecific stands are further characteristics which render this species a superior competitor. In marked contrast with most other tropical vegetation types, Afrotrilepis mats have therefore to be regarded as an example of a naturally occurring, extremely species-poor community, in which competitive displacement and the low degree of disturbance prevent higher diversity.     

Biodiversity and vegetation of small-sized inselbergs in a West African rain forest (Taı, Ivory Coast)

The vegetation of small granitic rock outcrops (geomorphologically small-sized inselbergs) which do not reach the canopy was studied in the Taı rain forest (southwestern Ivory Coast) under aspects of species diversity and phytogeographical affinities. Rock outcrops form edaphically arid (due to absent or very sparse soil cover) and microclimatologically xeric (i.e. low air humidity, temperature regularly exceeding 50°C) islands with cryptogamic crusts, succulents and poikilohydric vascular plants as characteristic elements of their vegetation which differs totally from the surrounding forest. Altogether sixty-six species of vascular plants out of twenty-nine families occur, the number of species correlates positively with inselberg size. Compared with large inselbergs the microclimatic attributes of small-sized rock outcrops are less pronounced. This is accompanied by a decrease of typical inselberg taxa (i.e. species mainly occurring on inselbergs). Low beta diversity between inselbergs indicates deterministic influences as important regulators of species composition. Annual Poaceae and Cyperaceae are richly represented. It can be hypothesized that inselbergs may represent natural growing sites of widely distributed tropical weeds today. Inselbergs might provide habitat resources for savanna elements in rain forest zones.     

Diversity and ecology of saxicolous vegetation mats on inselbergs in the Brazilian Atlantic rainforest

Inselbergs occur as mostly dome-shaped rock outcrops in all climatic and vegetational zones of the tropics. Consisting of Precambrian granites and gneisses, they form ancient and stable landscape elements. Due to harsh edaphic and microclimatic conditions, the vegetation of inselbergs differs markedly from those of the surroundings. Monocotyledonous mats form one of the most characteristic communities of this ecosystem. The floristic composition of this community was studied on six inselbergs located in the Brazilian Atlantic rainforest. Dominating are Bromeliaceae ( Alcantarea , Encholirium ), Velloziaceae ( Vellozia ), Cactaceae ( Coleocephalocereus ) and the Cyperaceae Trilepis . The alpha diversity of the mats was relatively uniform among the six outcrops. However, beta diversity varied considerably between the different sites. Beta diversity was highest at the most speciose locality indicating a high degree of stochasticity in colonization. In contrast to the low diversity mats on West African inselbergs, the Brazilian rock outcrops bear a floristically more diverse mat community rich in endemics. Thus the edaphically controlled inselberg vegetation reflects the outstanding diversity of the Mata Atlaântica. Possibly the high species richness of mats on East Brazilian inselbergs is a consequence of a large species-pool. The processes that regulate regional and local diversity in the Mata Atlaântica are not fully clear. It is assumed that historical (i.e. long-term stability) and biotic (i.e. evolutionary interchange of taxa between canopies and rock outcrops followed by differentiation of local populations) conditions have promoted high rates of speciation and their coexistence in isolated habitats. A danger to the unique vegetation of East Brazilian inselbergs is the establishment of invasive weeds.     

Seasonal Dynamics of Plant Diversity on lnselbergs in the Ivory Coast (West Africa)

Inselbergs are isolated monolithic outcrops which are characterized by large areas of exposed crystalline rock. Due to harsh edaphic and microclimatic conditions, inselbergs are completely differentiated from their surroundings. Consequently they host a very distinct vegetation which is being investigated on a global scale over a six year period. The seasonal dynamics of Selected plant communities (Afrotrilepis pilosa mat, shallow depression, ephemeral flush vegetation) on granitic inselbergs in the Comoe National Park (NE Ivory Coast) were studied during the rainy period from May to November 1991 by recording all vascular plant species at 12 intervals. For the habitats investigated, the seasonal vegetation dynamics were related to the rainfall pattern. Maximum values both in species diversity and richness were attained in the first third of the rainy period. Drought in August and September caused a decline in species number and diversity in the shallow depression and ephemeral flush vegetation, resulting in mortality of more than 20% of the species. The individual communities studied differed considerably in species diversity and richness. We conclude that ephemeral flush and shallow depression communities are more species rich than the mat community which is dominated by the highly competitive and specialized K-strategist Afrotrilepis pilosa (a poikilohydric Cyperaceae) due to stochastic climatic perturbations which allow the maintenance of species rich non-equilibrium assemblages with r-strategists as major components.     

Holocene boundary dynamics of a northern Australian monsoon rainforest patch inferred from isotopic analysis of carbon, (14C and δ13C) and nitrogen (δ15N) in soil organic matter

Abstract Soil organic matter (SOM) was sampled from lateritic soil profiles across an abrupt eucalypt savanna–monsoon rainforest boundary on the north coast of Croker Island, northern Australia. Accelerator mass spectrometry dating revealed that SOM that had accumulated at the base of these 1.5 m profiles had a radiocarbon age of about 5000 years. The mean carbon and nitrogen stable isotope composition of SOM from 10 cm deep layers from the surface, middle and base of three monsoon rainforest soil profiles was significantly different from the means for these layers in three adjacent savanna soil profiles, suggesting the isotopic ‘footprint’ of the vegetation boundary has been stable since the mid Holocene. Although there were no obvious environmental discontinuities associated with the boundary, the monsoon rainforest was found to occur on significantly more clay rich soils than the surrounding savanna. Tiny fragments of monsoon rainforest and abandoned ‘nests’ (large earthen mounds) of the orange‐footed scrubfowl, an obligate monsoon rainforest species, occurred in the savanna, signalling that the rainforest was once more extensive. Despite episodic disturbances, such as tropical storm damage and fires, the stability of the boundary is probably maintained because clay rich soils enable monsoon rainforest tree species to grow rapidly and achieve canopy closure, thereby excluding grass and reducing the risk of fire. Conversely, slower tree growth rates, grass competition and fire on the savanna soils would impede the expansion of the rainforest although high rainfall periods with shorter dry seasons may enable rainforest trees to grow sufficiently quickly to colonize the savanna successfully.     

Habitat diversity associated to island size and environmental filtering control the species richness of rock‐savanna plants in neotropical inselbergs

Disentangling the multiple factors controlling species diversity is a major challenge in ecology. Island biogeography and environmental filtering are two influential theories emphasizing respectively island size and isolation, and the abiotic environment, as key drivers of species richness. However, few attempts have been made to quantify their relative importance and investigate their mechanistic basis. Here, we applied structural equation modelling, a powerful method allowing test of complex hypotheses involving multiple and indirect effects, on an island‐like system of 22 French Guianan neotropical inselbergs covered with rock‐savanna. We separated the effects of size (rock‐savanna area), isolation (density of surrounding inselbergs), environmental filtering (rainfall, altitude) and dispersal filtering (forest‐matrix openness) on the species richness of all plants and of various ecological groups (terrestrial versus epiphytic, small‐scale versus large‐scale dispersal species). We showed that the species richness of all plants and terrestrial species was mainly explained by the size of rock‐savanna vegetation patches, with increasing richness associated with higher rock‐savanna area, while inselberg isolation and forest‐matrix openness had no measurable effect. This size effect was mediated by an increase in terrestrial‐habitat diversity, even after accounting for increased sampling effort. The richness of epiphytic species was mainly explained by environmental filtering, with a positive effect of rainfall and altitude, but also by a positive size effect mediated by enhanced woody‐plant species richness. Inselberg size and environmental filtering both explained the richness of small‐scale and large‐scale dispersal species, but these ecological groups responded in opposite directions to altitude and rainfall, that is positively for large‐scale and negatively for small‐scale dispersal species. Our study revealed both habitat diversity associated with island size and environmental filtering as major drivers of neotropical inselberg plant diversity and showed the importance of plant species growth form and dispersal ability to explain the relative importance of each driver.     

Diversity and distribution of murid rodent populations between forest and derived savanna sites within south western Nigeria

Southward encroachment of the derived savanna zone with its attendant denudation of the rainforest in Nigeria adversely affects the diversity and distribution of mammals, particularly murid rodents. This study was carried out to establish the identities of murid rodent populations and to compare their diversity between forest and derived savanna sites within south western Nigeria. Identification of captured specimens was carried out by multivariate statistics of body and skull measurements, and also by analysis of certain discrete characters. Six species were identified: Praomys tullbergi, Hylomyscus stella, Mastomys natalensis, Arvicanthis rufinus, Lemniscomys striatus and Lophuromys sikapusi. Forest sites possessed a higher Shannon’s diversity index (1.685) than the derived savanna sites (0.978), containing a higher number of taxa and larger abundance of murid rodents. This underscores the negative impact that encroachment of the derived savanna has upon diversity of animals in areas that were once forest.     

High species diversity and turnover in granite inselberg floras highlight the need for a conservation strategy protecting many outcrops

Determining patterns of plant diversity on granite inselbergs is an important task for conservation biogeography due to mounting threats. However, beyond the tropics there are relatively few quantitative studies of floristic diversity, or consideration of these patterns and their environmental, biogeographic, and historical correlates for conservation. We sought to contribute broader understanding of global patterns of species diversity on granite inselbergs and inform biodiversity conservation in the globally significant Southwest Australian Floristic Region (SWAFR). We surveyed floristics from 16 inselbergs (478 plots) across the climate gradient of the SWAFR stratified into three major habitats on each outcrop. We recorded 1,060 species from 92 families. At the plot level, local soil and topographic variables affecting aridity were correlated with species richness in herbaceous (HO) and woody vegetation (WO) of soil‐filled depressions, but not in woody vegetation on deeper soils at the base of outcrops (WOB). At the outcrop level, bioclimatic variables affecting aridity were correlated with species richness in two habitats (WO and WOB) but, contrary to predictions from island biogeography, were not correlated with inselberg area and isolation in any of the three habitats. Species turnover in each of the three habitats was also influenced by aridity, being correlated with bioclimatic variables and with interplot geographic distance, and for HO and WO habitats with local site variables. At the outcrop level, species replacement was the dominant component of species turnover in each of the three habitats, consistent with expectations for long‐term stable landscapes. Our results therefore highlight high species diversity and turnover associated with granite outcrop flora. Hence, effective conservation strategies will need to focus on protecting multiple inselbergs across the entire climate gradient of the region.     

Climate and land-use interactively shape butterfly diversity in tropical rainforest and savanna ecosystems of southwestern China

Human-induced habitat conversion and degradation, along with accelerating climatic change, have resulted in considerable global biodiversity loss. Nevertheless, how local ecological assemblages respond to the interplay between climate and land-use change remains poorly understood. Here, we examined the effects of climate and land-use interactions on butterfly diversity in different ecosystems of southwestern China. Specifically, we investigated variation in the alpha and beta diversities of butterflies in different landscapes along human-modified and climate gradients. We found that increasing land-use intensity not only caused a dramatic decrease in butterfly alpha diversity but also significantly simplified butterfly species composition in tropical rainforest and savanna ecosystems. These findings suggest that habitat modification by agricultural activities increases the importance of deterministic processes and leads to biotic homogenization. The land-use intensity model best explained species richness variation in the tropical rainforest, whereas the climate and land-use intensity interaction model best explained species richness variation in the savanna. These results indicate that climate modulates the effects of land-use intensity on butterfly alpha diversity in the savanna ecosystem. We also found that the response of species composition to climate varied between sites: specifically, species composition was strongly correlated with climatic distance in the tropical rainforest but not in the savanna. Taken together, our long-term butterfly monitoring data reveal that interactions between human-modified habitat change and climate change have shaped butterfly diversity in tropical rainforest and savanna. These findings also have important implications for biodiversity conservation under the current era of rapid human-induced habitat loss and climate change.     

Local and regional-scale responses of ant diversity to a semiarid biome transition

The locations of biome transitions and ecotones are frequently defined by the rapid shift from one form of dominant vegetation to another. The composition of animal taxa is predicted to shift in parallel with that of dominant plants and species diversity is predicted to he greater in transitional zones than in adjacent areas. We asked whether ant species diversity and composition supported these predictions across a biome transition between shortgrass steppe and Chihuahuan desert vegetation. Neither species richness nor diversity was highest at the biome transition region as a whole, or within habitats in the biome transition. The biome transition region was not intermediate in ant species composition or in the representation of different faunal complexes. The community similarity between matched habitats shared between the biome transition zone and adjacent regions was less than that between distinct habitats occurring within regions. A zoogeographic transition for ants may occur to the north of the phytogeographic transition and may be coincident with the northern limits of monsoonal precipitation patterns. In contrast, the phytogeographic transition may be related to less extreme climatic variation within the monsoonal region occurring further south.     

Spatial pattern of dry rainforest colonizing unburnt Eucalyptus savanna

Abstract The spatial pattern of dry rainforest and savanna tree species was analysed in a 1.56‐ha plot within an unburnt eucalypt savanna woodland in north Queensland, Australia. Rainforest colonization constituted only 1.3% of the basal area and mostly consisted of individuals less than 3 m high. The distribution of rainforest trees was highly clumped around the large savanna eucalypt trees. Ecological mechanisms generating the clumped distribution are discussed in light of evidence from this study and the literature. Herbaceous biomass was not reduced under trees, suggesting that relief from grass competition has not favoured rainforest colonization under tree crowns. Edaphic facilitation through nutrient enrichment under savanna tree crowns appears to be only minor on the moderate fertility soils of the area. The highly clumped pattern of colonizing dry rainforest may be a consequence of seeds dropped from birds roosting in savanna trees.     

高黎贡山自然保护区西坡垂直带蚂蚁群落研究

首次研究了高黎贡山自然保护区西坡垂直带蚂蚁群落及共物种多样性。在北段和中北段随海拔升高优势种数目增加,而在南段随海拔升高优势种数目递减。在北段和中北段海随海拔升高优势种所占百分比递减,而在南段随海拔升高优势种所占比例递增。在西坡４个地段均呈现随海拔升高种数目和密度递减的基本规律。北段和中北段随海拔升高优势度指数降低,中南段和南段随海拔升高优势度指数增大。北段随海拔升高多样性指数递增,中北段多样性指数缺乏规律性,中南段和南段随海拔升高多样性指数递减。北段、中北段和中南段均呈现随海拔升高无效度指数递增规律,而南段表现出随海拔升高均匀度指数递减的相反情况。西坡４个地段垂直带上,蚂蚁群落之间的相似系数几乎均在０．００－０．２５,处于极不相似水平,只有１个系数超过此范围,我们认为在原始植被状态下,高黎贡山自然保护区西坡蚂蚁群落的基本规律是随着海拔升高。优势种数目递减,优势种所占比 递增,物种数目递减,优势度指数递增,多样性指数递减,均匀度指数递减。山体中部和下部植被的毁坏导致了实际调查中出现的反常情况。     

Cyanobacteria and Cyanobacterial Lichens from Inselbergs of the Ivory Coast,Africa

This study focuses on the saxicolous lichens and cyanobacteria of the open, exposed rock surface of inselbergs. Twenty-three species of cyanobacteria and 17 cyanobacterial lichen species (“cyanolichens”) from several inselbergs and other rocky outcrops of three major climatic regions, savanna, transition zone and rain forest, are reported from the Ivory Coast. Inselbergs are isolated and frequently mountains consisting of Precambrian granites or gneisses that abruptly rise from the surrounding plains. Cyanobacteria were found to be the dominating organisms on all rock surfaces. The lichens found mainly belong to the family Peltulaceae and a few were present from the family Lichinaceae. Nine species of the cyanolichens and most of the cyanobacteria are new for the Ivory Coast. A gradient in total species number (cyanolichens and cyanobacteria) occurs from the savanna to the rain forest, with a decrease in species number towards the rain forest. Saxicolous cyanobacterial lichens reached a higher species number in the savanna type ecosystem (11) than on inselbergs in the rain forest (7). The cyanolichens and cyanobacteria found are characteristic for larger, light-exposed rock surfaces and species like P. congregate, P. lingulata, P. tortuosa and P. umbilicata preferentially occur on the granite or sandstone of inselbergs.     

Genetic structure of the xerophilous bromeliad Pitcairnia geyskesii on inselbergs in French Guiana – a test of the forest refuge hypothesis

Inselbergs are isolated granitic rock outcrops that provide distinctive ecological conditions. In northern South America they rise above the surrounding rainforest. Among inselberg specialists, Pitcairnia geyskesii (Bromeliaceae) is restricted to these habitats in French Guiana. We studied populations from 12 inselbergs using 7 microsatellite loci to give a "reverse image" of the reduction-expansion of the rainforest in the context of the refuge hypothesis. Our analyses showed that populations are fragmented with dispersal occurring only over very short distances. Genetic diversity was higher in northern French Guiana, whereas specific alleles were observed in the south. The results point to the occurrence of a dry corridor in the north, as hypothesized by Tardy (1998) based on charcoal analyses, whereas de Granville's (1982) hypothesis of a unique past refuge is not confirmed. Moreover, our data suggests the importance of Oyapock River as a pathway for range expansion, arguing against the potential role of the Inini-Camopi Mountains as a physical barrier. Finally, in spite of a strongly argued scenario in favour of a north-to-south migration history, a clear genetic isolation of P. geyskesii populations living on inselbergs of the Mitaraka archipelago suggests a distinct ancestry of the most southern populations.     

佛坪国家级自然保护区植被垂直带谱及其与邻近地区的比较

在首先确定垂直带划分的原则和将佛坪自然保护区植被划分为落叶阔叶林带、中山小叶林带和亚高山针叶林带３个垂直带,各垂直带植被物种组成的区系、生活型组成和物种多样性的差异证实了这种划分的合理性。与邻近地区植被垂直带谱的比较表明,佛坪自然保护区的植被垂直带谱与秦岭北坡有明显差异,表现为典型的暧温带与北亚热带过渡区域的植被景观,虽然基带以上各植被带暖温带特色很明显,但其植被属性应是北亚热带的。     

Provenance variation in morphology and leaflet anatomy of Parkia biglobosa and its relation to drought tolerance

 Measurements of morphological and anatomical characteristics made on 4-month-old seedlings from five provenances of Parkia biglobosa (Jacq.) Benth., grown in a tropical greenhouse under controlled conditions, showed significant differences between the northern provenances, which represented the Sudan savanna zone, and the southern provenances which represented the forest zone of the species distribution. Seedlings of provenances from the savanna zone showed more xerophytic characteristics than those from the forest zone. They were smaller in height, with a greater number of smaller leaflets per pinnae, a lower specific leaflet mass, smaller palisade and guard cells and a lower ratio of palisade:non-palisade tissue. There were also significant differences in the same characteristics, except palisade cell length, between provenances from the easterly and westerly extremes of the species range. Stomatal conductance measured after 2 weeks of droughting also showed that provenances from the savanna zone had higher values, indicating greater physiological activity and reduced stress, compared with provenances from the forest zone. Multivariate discriminant analysis of the morphological and anatomical characteristics showed that 98% of the seedlings were classified correctly with respect to provenance. The squared distances between provenances were closely related to the geographical distances between locations. Yet, not all variation in leaf anatomy could be explained by the difference in climatic conditions from north to south. Also drought tolerance was not always related to morphological and anatomical features. It is possible that clinal variation within the species from east to west may be interacting with climatic differences from north to south. Received: 2 June 1998 / Accepted: 29 June 1998     

Vegetation survey of the Wasgomuwa National Park: reconnaissance

Wasgomuwa National Park is situated south of Polonnaruwa and north of Mahiyangana in Sri Lanka and is bordered by two rivers Mahaweli Ganga and Amban Ganga. Bioclimatically most of the park lies in the dry zone of the country but the south west embraces the intermediate zone. The vegetation is broadly a dry monsoonal forest but forms an interesting mosaic of vegetation types from Semi evergreen forest to abandoned paddy fields, on a finer scale.The vegetation of the park was studied in detail on ground and also using aerial photographs. It was possible to classify the vegetation into many distinct types or phytosociological units, based on their cover, site characteristics, physiognomy and species composition.     

Anthropogenic changes to the landscape resulted in colonization of koalas in north‐east New South Wales,Australia

Significant changes in the distribution and persistence of species have been driven by Pleistocene cyclical climate changes and, more recently, by human modification of the environment. In eastern Australia, Pleistocene cyclical patterns in temperature and aridity led to the expansion and retraction of rainforest and likely affected the distribution of the koala (Phascolarctos cinereus, family Phascolarctidae), a species preferring dry or open woodland. More recently, anthropogenic landscape modification has led to a large‐scale change in distribution of the koala following the destruction of approx. 75 000 ha of subtropical rainforest in north‐east New South Wales termed the ‘Big Scrub’. Sharing of the control region haplotypes to the north and south of this region indicate historical connectivity of coastal koala populations. However, the majority (110/115) of sampled koalas from this region shared a single mitochondrial control region haplotype, suggesting that koalas did not persist in multiple refugial pockets within a heterogenous rainforest but expanded into the region following deforestation. Bayesian cluster analysis of microsatellite data consistently identified two clusters of koalas. One cluster, in the north of the area, had high microsatellite diversity (10 alleles per locus, He = 0.79) and clustered with koalas further to the north, thus suggesting a southerly expansion into the cleared area. To the south was a cluster with significantly lower diversity (six alleles per locus, He = 0.59, P < 0.001). It is possible there has been restricted or filtered movement of koalas between these clusters, which coincides with a cleared river valley and associated roads or immigration from populations both to the north and to the south. This study gives an insight into the timescale of changes in species distribution following rapid alterations to suitable habitat.     

A Preliminary study on the Vertical Belts of Vegetation of the Eastern Himalayas

The horizontal differentiation of vegetation is very conspicuous, on both the south and the north slopes of the Himalayas. The Eastern Himalayas are the most humid part of the mountain system, being mainly covered by forests. The tropieal forest stretches the south slope of the mountain as far north as 29˚ N, mueh beyond the ordinary boundary of tropieal forest in other parts of the world. On the north slope, the vegetation shows a transition from forest to steppe. In general, 4 vertical spectra of vegetation zones may be recognized in tile Eastern Himalayas with two on either slope. On the south the basic belts of the spectra are respeetively (1) tropical evergreen rainforest plus semievergreen rainforest (humid) and (2) tropical rainforest plus monsoon deciduous forest (subhumid). On the north, the respective basic belts are (3) montane needle and broad-leaf mixed forest (subhumid) and (4) montane shrubby steppe (semiarid). The 4 altogether indicate the regional differentiation of vegetation in the Eastern Himalayas, according to different climatic conditions.