Elephant herbivory in Majete Wildlife Reserve,Malawi

Elephants have a major influence on vegetation structure, composition and ecosystem processes, and are primary agents of habitat change in Africa. At moderate‐to‐high population densities, elephants can damage vegetation, especially when enclosed in protected areas. This study examines the effects of elephant browsing on woody trees in Majete Wildlife Reserve (WR), south‐western Malawi. Regression analysis is used to assess the associations of six factors known to drive elephant browsing in other areas and determine which ones have the most influence on browsing at Majete WR. Twenty‐four per cent of tagged trees had been subject to elephant browsing. The model with vegetation type, stem diameter and distance from permanent water correctly predicted browsing for 80% of the observations. Elephants mostly favoured riparian woodlands, followed by Acacia‐dominated woodland and Brachystegia‐dominated woodland. Browsing occurrence was negatively related to distance from permanent water and diameter at breast height(DBH). A larger number of trees, sampled at random and covering a larger portion of the reserve would provide more reliable estimates of browsing and related factors. Knowledge of time‐ and site‐specific factors affecting elephant browsing can be used to forecast future habitat transformations and manipulate the range of the elephants within the reserve.     

Observations of tree growth, woodland structure and elephant damage on Colophospermum mopane in Luangwa Valley, Zambia

Six samples of tagged Colophospermum mopane were monitored for five years in locations with varying soil characteristics but with similar elephant densities. Physiognomic variation among the samples was related to soil differences, which also correlated with different browsing habits by elephants. The impact of elephant browsing further influenced both the physiognomy and demography of C. mopane. Results from this study suggest that the influence of soils and elephants on C. mopane alter successional transitions from grassland to woodland. Soils that promote coppicing of C. mopane yield less stable woodlands when associated with elephants than soils promoting woodlands with large bolus, non-coppicing trees. The dynamics of the latter are determined more by tree recruitment as influenced by such agents as other browsers or frequency and seasonality of bush fires. Implications for forest/elephant management are discussed.     

Changes in the structure and composition of miombo woodlands mediated by elephants (Loxodonta africana) and fire over a 26-year period in north-western Zimbabwe

Changes in structure and composition of miombo woodlands mediated by elephants and fire were studied in 26-year-old permanent transects established in 1972 in north-western Zimbabwe. Elephants caused 48% decline in proportions of large trees (>11 cm diameter), significant reductions (30.9–90.9%) in tree heights, reductions in stem areas (43.5%) and densities (2.5%) of all trees. There were increases in proportions of small trees (64.8%), shrub canopy volumes (271%) and shrub densities (172%). These increases are attributed to natural recruitment because of longer fire-free periods and reduction of tree suppression effects on lower strata as a result of elephant-induced tree declines. Frequencies of occurrence of most species dropped by 28–89.6%. Brachystegia boehmii was replaced by Pseudolachnostylis maprouneifolia as the most dominant tree, largely because of high elephant preference for Brachystegia boehmii . A new suite of species, dominated by Combretaceae, increased in dominance resulting in local floristic changes. Reductions in old elephant (33.4%), old unknown (89.9%) and new elephant (13.7%) damage suggest that elephant occupancy of miombo woodlands has declined, possibly because of limited availability of preferred browse species. This study clearly shows that elephants and fire have contributed significantly to the changes in miombo woodlands in the area.     

Effects of large herbivores and fire on the regeneration of Acacia erioloba woodlands in Chobe National Park, Botswana

Acacia erioloba woodlands provide important forage and shade for wildlife in northern Botswana. Mortality of mature trees caused by browsing elephants has been well documented but the lack of regeneration of new trees has received little attention. Annual growth of new shoots and changes in height were measured to determine the influence of elephants and small ungulate browsers, rainfall and fire on the growth and survival of established A. erioloba seedlings from 1995 to 1997 in the Savuti area of Chobe National Park. All above‐ground vegetation was removed from 40% of established seedlings in 1995 and 28% in 1997 by browsing elephants, and the mean height of remaining seedlings decreased from >550 mm to <300 mm. When seedlings browsed by kudu, impala and steenbok but not elephants are considered, mean seedling height increased <50 mm per year, even though mean new shoot growth remaining at the end of the dry season was 100–200 mm. Fires burned portions of the study area in 1993 and 1997, killing above‐ground vegetation, but most established A. erioloba seedlings survived, producing coppice growth from roots. While elephants and fire caused the greatest reduction in established seedling height and number, small browsers suppressed growth, keeping seedlings vulnerable to fire and delaying growth to reproductive maturity.     

Termite Mounds as Nutrient–Rich Food Patches for Elephants

This study compared elephant use of woody vegetation on termite mounds with surrounding woodlands in western Zimbabwe. Twelve sites consisting of paired plots on termite mounds and in woodlands were selected. At each site, soil and vegetation samples (leaf and stem) were collected for chemical analysis. Both soil and plant samples were analyzed for calcium, magnesium, potassium, sodium, and phosphorus, and plant samples were also analyzed for crude protein concentration. Two indices of elephant feeding damage were computed: the median number of stems and branches removed per plant, and the mass of stems and branches removed by elephants per unit area. Termite mound soils had higher concentrations of all elements tested than soils from woodlands, and termite mounds differed from woodland plots in terms of plant species composition. Trees growing on termite mounds had higher concentrations of all nutrients except sodium and crude protein, and were subjected to more intense feeding by elephants than trees from the surrounding vegetation matrix. Termite mounds may play an important role in determining food availability and spatial feeding patterns by elephants and other herbivores.     

Freezing tolerance and avoidance in high tropical Andean plants: Is it equally represented in species with different plant height?

Summary CO₂ assimilation in relation to light intensity and the relationship between leaf nitrogen and phosphorus concentrations and CO₂ assimilation in 14 species of ecologically important Zimbabwean trees were examined. Eight of the species are members of the Fabaceae (Leguminosae). In the majority of Zimbabwean climax woodlands, the dominant trees are non-nodulating members of the sub-family Caesalpinioideae. The species examined have higher light saturation points (>700 mol m^–2 s^–1) than woody species from temperate areas; one species, Acacia nigrescens, did not reach saturation at photon fluxes greater than 1500 mol m^–2 sec^–1. Higher leaf nitrogen content was found to correlate positively with higher CO₂ assimilation rates (r=0.85; P0.0003); there was no correlation between leaf phosphorus content and CO₂ uptake rates. There were no significant differences between sites in terms of leaf nitrogen or phosphorus content, but the mean photosynthetic rate at one of the sites (Chizedzi) was lower. Taxa from the nodulating legumes were found to have higher leaf nitrogen contents (309.1±SD 22 mmol m^–2) than those of the non-nodulating species (239±33); the lowest nitrogen contents were found in nonleguminous trees (179±42), with the exception of Ziziphus mucronata. This species may form an association with an N₂-fixing actinomycete.     

Vegetation changes on formerly overgrazed hill slopes in semi-arid central Tanzania

Abstract. Composition of hill slope vegetation was studied in a semi-arid part of upland Tanzania where all grazing had been banned for 12 yr. The hills had been severely overgrazed previously and suffered from heavy gully and sheet erosion. Eight vegetation types are described. Floristic gradients revealed by numerical ordination techniques were found to be related mainly to degree of erosion, soil type and succession. The more or less bare soil that prevailed after grazing had ceased is now covered by grassland, woodland and immature secondary forest. The grasslands are still characterized by early successional species and they will probably remain open grassland as long as frequent burning continues. Brachystegia woodlands may have developed during earlier periods when the field layer was sparse due to grazing. The grazing had reduced the frequency of fire which in turn promoted the establishment of Brachystegia spp. Secondary forests are believed to have developed mainly where fires were not frequent, particularly at higher altitudes.     

Plant species as indicators of ancient woodland in northwestern Germany

Abstract. 315 isolated semi-natural and natural stands in NW Germany were investigated floristically, 285 stands of ancient woodlands and 30 recent ones. In the study area nearly all semi-natural and natural ancient woodlands are found on mesotrophic or eutrophic sites and can be assigned to the Fagetalia. 54 selected herbaceous and five woody Fagetalia-species have been tested in their association to ancient woodlands. 21 of the 59 selected plant species show a highly significant association to ancient woodlands and seven species show a weak significance. 31 plant species are not significantly correlated with ancient woodland sites, but 25 of them have a low frequency. The result shows that investigations of the historical ecology can help to understand floristic composition of present-day woodlands. Restriction of many woodland species to ancient woodlands, especially rare species, emphasizes the importance of woodlands with a long continuous history for the preservation of endangered species. Irrespective of ecological conditions, the restriction of plant species to ancient woodlands seems to be mainly caused by their low ability to colonize recent woodlands, especially isolated stands. Differences in the association to ancient woodlands between European countries are assumed to be a function of both time and degree of isolation of woodlands, of the dispersal mode of the plant species and of the availability of suitable habitats.     

How selective are elephants as agents of forest tree damage in Bwindi Impenetrable National Park,Uganda?

Elephants are locally concentrated in Bwindi Impenetrable National Park. Vegetation damage attributable to elephants appears to be increasing and may result in the modification of the forest. We examined the implied selectivity of stem damage due to elephants. We followed 26.84 km of recent elephant trails and used 122 plots to document tree damage in relation to species, stem sizes and locations. Of 897 trees (DBH ≥2 cm), 542 (60.4%) were intact, 22 (2.5%) debarked, 274 (30.5%) toppled and 172 (19.2%) had broken branches. Small trees were more likely to be pushed over or have their branches broken, whereas large trees were more commonly debarked. The species most frequently selected for damage included mid‐successional species such as Newtonia buchananii, Myrianthus holstii and Chrysophyllum albidum. These species may be vulnerable to increasing elephant numbers. Our analyses using general linear models indicate that elephants are selective concerning where, how and what tree stems they damage. We found a higher incidence of elephant damage per‐tree stem in open areas than in more closed areas, suggesting feedback in which elephants maintain open habitats that may be conducive for other species such as mountain gorillas. More work is needed to better determine how changing elephant numbers may influence Bwindi's conservation values.     

The Seronera bull problem: The elephants

(1) The activities of bull elephants in the vicinity of the Serengeti National Park headquarters at Seronera was monitored from 1968 to 1971. (2) Elephant bulls congregate in small groups of ever–changing composition; a dominance heirarchy cuts across group integrity. (3) Occupancy of the drainage area of the Seronera river is predictable by rainfall; changes in occupancy level are a question of duration rather than amplitude. (4) The feeding behaviour of the bulls is described. It is argued that large Acacia trees are uprooted for nutritional rather than social reasons. The effects of the bulls on the Seronera woodlands will be analysed in Part II (Croze, 1974).     

Model highlights likely long‐term influences of mesobrowsers versus those of elephants on woodland dynamics

The potential long‐term influences of mesobrowsers versus those of savannah elephants on woodland dynamics have not been explored. This may be a critical omission especially in southern African savannahs, where efforts to preserve existing woodlands are typically directed at elephant management. We describe a simple browse–browser model, parameterized from an extensive review of the literature and our own data, including quantitative assessment of impala impact, from the study site, iMfolozi Park, South Africa. As there is a paucity of species‐specific demographic data on savannah woody species, we modelled, in a novel approach, functional groups of plant species typical of Acacia woodlands. Outputs suggest that over the long term (100 years), low‐to‐moderate densities of impala will have a similar impact on woodland structure, in terms of density of adult trees, as low‐to‐moderate densities of elephant. Further, the outputs highlight the apparently strong synergistic effect impala and elephant impacts combined have on woodland dynamics, suggesting that reduction or removal of either impala or elephant will radically reduce long‐term destruction of savannah woodlands. Recorded changes in adult tree numbers in iMfolozi broadly supported the model's outputs.     

Woodland changes in Ruaha National Park (Tanzania) between 1976 and 1982

Tree surveys made in 1976 and 1977 were repeated in 1982. Three species were measured: Commiphora ugogensis, Acacia albida, and Adansonia digitata. All three showed significant decreases due to elephant browsing. These results confirm the predictions of the earlier surveys: elephants are causing a rapid and severe change in the rift valley woodlands of the Park.     

Woody vegetation damage by the African elephant during severe drought at Pongola Game Reserve,South Africa

Elephants (Loxodonta africana) significantly alter ecosystem structure and composition through browsing (e.g. pollarding, debarking and toppling). Such browsing is predicted to intensify during severe drought which may become more common with climate change. Here, we make use of an elephant impact survey from 2012 to 2015 and during the El Nino drought of 2015–2016 at Pongola Game Reserve (107 km²), KwaZulu-Natal, to investigate how severe drought influenced damage severity of different tree heights and species by elephants in this small reserve. Contrary to expectations, damage to common species did not change with severe drought. Crown damage had the highest predicted probability across heights (29%–90%) and species (46%–75%) regardless of drought. However, we found severe drought increased the predicted probabilities of crown damage to smaller trees <4 m, mortality >6 m and severe damage at 4–6 m. Consequently, elephant damage during severe drought may alter vegetation structure by severely damaging or killing large trees (>4 m) and extensively damaging the crowns of trees <4 m. Long-term monitoring of elephant effects on woody vegetation is essential to enable science-based management in response to future drought and elephant damage (e.g. range expansion, beehive deterrents) to protect elephants and conserve woody vegetation.     

Effect of low rainfall and browsing by large herbivores on an enclosed savannah habitat in Kenya

Savannah ecosystems in East Africa are rarely stable and can experience rapid local changes from dense woodlands to open plains. In this 3‐year study there was a reduction of 16.3% in a height‐stratified sample of nearly 1000 individually marked Acacia drepanolobium trees. The study was carried out in an enclosed fire‐free wooded grassland habitat in the Laikipia region of Kenya. The trees were monitored from 1998 to 2001, a period that included 12 months when rainfall was 60% below average. Elephants were responsible for the loss of 40% of the trees, black rhinos 33% and 27% died from the effects of the drought. Low rainfall was correlated with increased damage as elephants switched diet from grass to trees. Heavy browsing by giraffes reduced tree growth rates and increased their susceptibility to drought. Hence the combination of low rainfall and heavy browsing by elephants, black rhinos and giraffes led to the rapid tree loss. These findings have implications for research into the causes of instability in savannah ecosystems and the management of enclosed reserves.     

Sleeping Parties and Nest Distribution of Chimpanzees in the Savanna Woodland, Ugalla, Tanzania

We conducted ecological studies of chimpanzees (Pan troglodytes) in the Ugalla area, western Tanzania. Ugalla is one of the driest habitats of chimpanzees and the Ugalla River is the eastern boundary of chimpanzee distribution. Most of Ugalla is occupied by savanna woodlands dominated by deciduous trees of Brachystegia and Julbernardia. Chimpanzees tended not to make nests in riverine forests in plains, but in small patchy forests dominated by Monopetalanthus richardsiae and valley forests dominated by Julbernardia unijugata on slopes in mountainous areas. We estimated population density of chimpanzees to be 7–9 × 10⁻² individuals/km² based on nest censuses, suggesting that 2–3 × 10² individuals inhabited the 3352 km² area of Ugalla. The size of the largest nest cluster (n=23) suggests that 1 unit group (community) comprised 30–35 individuals. In the daytime, chimpanzees formed small feeding parties (mean 2.0 individuals), but larger ones in the evening (mean 4.8 individuals and 5.2 individuals based on fresh nest clusters). The pattern might reduce the predation risk from large nocturnal carnivores such as lions and leopards. The sleeping sites may function as both a safe sleeping site and a meeting point for chimpanzees with a huge home range that may have difficulty in finding other members of their unit group.     

Effects of European colonization on indigenous ecosystems: post-settlement changes in tree stand structures in Eucalyptus–Callitris woodlands in central New South Wales, Australia

Aim There has been considerable debate about pre‐settlement stand structures in temperate woodlands in south‐eastern Australia. Traditional histories assumed massive tree losses across the region, whereas a number of recent histories propose that woodlands were originally open and trees regenerated densely after settlement. To reconcile these conflicting models, we gathered quantitative data on pre‐settlement stand structures in Eucalyptus–Callitris woodlands in central New South Wales Australia, including: (1) tree density, composition, basal area and canopy cover at the time of European settlement; and (2) post‐settlement changes in these attributes. Location Woodlands dominated by Eucalyptus species and Callitris glaucophylla, which originally occupied approximately 100,000 km² in central New South Wales, Australia. Methods We recorded all evidence of pre‐settlement trees, including stumps, stags and veteran trees, from 39 relatively undisturbed 1‐ha stands within 16 State Forests evenly distributed across the region. Current trees were recorded in a nested 900 m² quadrat at each site. Allometric relationships were used to estimate girth over bark at breast height, tree basal area, and crown diameter from the girth of cut stumps. A post‐settlement disturbance index was developed to assess correlations between post‐settlement disturbance and attributes of pre‐settlement stands. Results The densities of all large trees (> 60 cm girth over bark at breast height) were significantly greater in current stands than at the time of European settlement (198 vs. 39 trees ha^?1). Pre‐settlement and current stands did not differ in basal area. However, the proportional representation of Eucalyptus and Callitris changed completely. At the time of settlement, stands were dominated by Eucalyptus (78% of basal area), whereas current stands are dominated by Callitris (74%). On average, Eucalyptus afforded 83% of crown cover at the time of settlement. Moreover, the estimated density, basal area and crown cover of Eucalyptus at the time of settlement were significantly negatively correlated with post‐settlement disturbance, which suggests that these results underestimate pre‐settlement Eucalyptus representation in the most disturbed stands. Main conclusions These results incorporate elements of traditional and recent vegetation histories. Since European settlement, State Forests have been transformed from Eucalyptus to Callitris dominance as a result of the widespread clearance of pre‐settlement Eucalyptus and dense post‐settlement recruitment of Callitris. Tree densities did increase greatly after European settlement, but most stands were much denser at the time of settlement than recent histories suggest. The original degree of dominance by Eucalyptus was unexpected, and has been consistently underestimated in the past. This study has greatly refined our understanding of post‐settlement changes in woodland stand structures, and will strengthen the foundation for management policies that incorporate historical benchmarks of landscape vegetation changes.     

The effect of woodland area on avian community composition in a fragmented southern UK landscape and associated management recommendations

ABSTRACT

Capsule: Smaller woodlands not only support fewer species but also show different avian community composition due to loss of woodland interior and an increase in edge habitat.

Aims: To use observed community composition changes, rather than traditional total species richness-area relationships, to make area-specific management recommendations for optimizing woodland habitat for avian communities in fragmented landscapes.

Methods: 17 woodlands were selected in Oxfordshire, UK, with areas between 0.2 and 120 ha. Three dawn area searches were conducted in each woodland between 1st April and 28th May 2016 to record encounter rates for each species. The impact of internal habitat variation on woodland comparability was assessed using habitat surveys.

Results: Woodlands with area less than 3.6 ha showed a significant positive relationship between total avian species richness and woodland area. Woodlands with area over 3.6 ha were all consistent with a mean (± se) total richness of 25.4?±?0.6 species, however the number of woodland specialists continued to increase with woodland area. Woodland generalists dominated the total encounter rate across the area range, however the fractional contribution of woodland specialists showed a significant positive correlation with woodland area, while the fractional contribution of non-woodland species significantly decreased. Non-woodland species numbers peaked in mid-sized woodlands with enhanced habitat heterogeneity.

Conclusions: Community composition analysis enabled more targeted recommendations than total species richness analysis, specifically: large woodlands (over 25?ha) in southern UK should focus conservation efforts on providing the specific internal habitats required by woodland specialists; medium-sized woodlands (between approximately 4 and 25?ha) should focus on promoting internal habitat variety, which can benefit both woodland species and non-woodland species of conservation concern in the surrounding landscape; small woodlands (under 4?ha) should focus on providing nesting opportunities for non-woodland species and on improving connectivity to maximize habitat for woodland generalists and facilitate movement of woodland specialists.     

Impact of sudden oak death on tree mortality in the Big Sur ecoregion of California

The Big Sur ecoregion in coastal California is a botanically and ecologically diverse area that has recently experienced substantial mortality of oak (Quercus spp.) and tanoak (Lithocarpus densiflorus) trees due to the emerging forest disease sudden oak death, caused by the invasive pathogen Phytophthora ramorum. In response to the urgent need to examine environmental impacts and create management response strategies, we quantified the impact of P. ramorum invasion on tree mortality across the Big Sur ecoregion using high-resolution aircraft imagery and field data. Using the imagery, we mapped all detectable oak and tanoak trees possibly killed by P. ramorum infection within redwood-tanoak forests and mixed oak woodlands. To validate and improve our remote assessment, we quantified the number, size, and infection status of host trees in 77 field plots (0.25 ha). The field data showed that our remote assessment underestimated mortality due to the occurrence of dead trees in the forest understory. For each forest type, we developed regression models that adjusted our remote assessments of tree mortality in relation to field observations of mortality and local habitat variables. The models significantly improved remote assessment of oak mortality, but relationships were stronger for mixed oak woodlands (r ² = 0.77) than redwood-tanoak forests (r ² = 0.66). Using the field data, we also modeled the amount of dead tree basal area (m²) in relation to the density of mapped dead trees in mixed oak woodlands (r ² = 0.73) and redwood-tanoak forests (r ² = 0.54). Application of the regression models in a GIS estimated 235,678 standing dead trees in 2005 and 12,650 m² of tree basal area removed from the ecoregion, with 63% of mortality occurring in redwood-tanoak forests and 37% in mixed oak woodlands. Integration of the remote assessment with population estimates of host abundance, obtained from an independent network of 175 field plots (0.05 ha each), indicated similar prevalence of mortality in redwood-tanoak forests (20.0%) and mixed oak woodlands (20.5%) at this time. This is the first study to quantify a realistic number of dead trees impacted by P. ramorum over a defined ecological region. Ecosystem impacts of such widespread mortality will likely be significant.

Detecting invisible growth rings of trees in seasonally dry forests in Thailand: isotopic and wood anatomical approaches

We measured radial variation of carbon isotope composition and vessel traits in tree species in seasonally dry forests of Northeast Thailand to explore a more reliable and amenable method of tropical dendrochronology for trees that lack visually detectable and consistent growth rings. Six Dipterocarpaceae species (3 Shorea, 2 Dipterocarpus, and 1 Hopea species) with indistinct or irregular growth rings and teak (Tectona grandis), a species which forms distinct growth rings, were examined. The δ¹³C value variations in all species showed annual cyclicity. Dipterocarpaceae species usually marked the lowest values of δ¹³C in the middle of the growing season, whereas teak had the lowest values at nearly the end of the growing season. Since the growing season of the species examined almost corresponds to the rainy season in the study area, the δ¹³C variation was likely caused by the change in moisture availability. The different variation pattern of teak was attributable to its stronger dependence on ¹³C-enriched reserved material early in the growing season. Changes in tree vessel traits for all species examined also showed annual cyclicity. Dipterocarpaceae species showed significant correlation between δ¹³C values and vessel measurements. Vessel lumen (mean area, tangential and radial diameter, and proportion of total area) had a negative correlation, whereas vessel frequency showed a positive correlation. The correlations indicated that changes in vessel traits were caused by the seasonal variation of moisture available to the trees. Thus, we concluded that methods using wood anatomy, as well as δ¹³C, have great potential for use as tools in tropical dendrochronology within the context of seasonal climate.     

Bird communities ofBrachystegia andAcacia woodlands in Zambia

Summary The bird communities of fourBrachystegia and twoAcacia woodlands near Lusaka, Republic of Zambia, were quantitatively assessed in the rainy season (February 1976). The number of species and the species diversity values were rather similar in all areas, and so was also the total bird density (with one exception) in spite of a sevenfold difference of foliage quantity between the poorest and richest habitats investigated. Foliage-gleaning birds exploited foliage-richer habitats less intensively than foliage-poorer ones. The density of Palaearctic migrants, among whichPhylloscopus trochilus andMusicapa striata were most abundant, was low throughout; in fact practically none was recorded in relatively untouched habitats, but in much disturbed areas, for example, thinned-outBrachystegia or heavily grazedAcacia woodlands, they were locally more numerous. Everywhere they made up only a minor portion of all birds present, and this holds also if their density is related to the guild of ecologically corresponding African species; for example, the density ofPh. trochilus in comparison with that of the whole guild of foliage-gleaning passerines.The total bird density of the Zambian habitats was compared to that of temperate broadleaved woodlands, with which especially theBrachystegia woods bear a superficial resemblance, and was found to be about similar. This was unexpected considering the twice higher primary production measured for tropicalBrachystegia. The different degree of seasonality is pointed out as an important circumstance in this context. It was also unexpected that the total bird density was about the same in theBrachystegia andAcacia habitats, respectively, since the primary production, to judge from the difference in foliage volume, was much higher in the former.The species richness and diversity were much higher in the Zambian study plots than in temperate areas of the same size. Diversity indices were calculated and compared to expected ones on the assumption of MacArthur's broken-stick distribution and the canonical lognormal distribution, respectively. It was found that the abundance distributions of the Zambian communities fell in the range between the values expected for these models; perhaps somewhat closer to the broken-stick distribution. This accords to expectation for closely integrated bird communities.The Palaearctic migrants in the study areas occurred in much lower densities than they do in moderately favourable habitats in their European breeding areas. At leastPhylloscopus trochilus, and maybe also the other Palaearctic species encountered, probably inhabit other areas in tropical Africa in higher density than that found by us. That Palaearctic migrants are more abundant in modified habitats than in more undisturbed ones has been asserted before on qualitative grounds and was quantitatively confirmed by the present study. Their capacity to exploit modified habitats may be a significant factor for their future destiny, considering the profound landscape alteration that has commenced and is now accelerating in much of Africa.

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Vogelgesellschaften derBrachystegia- undAcacia-Savannen in Zambia

Zusammenfassung Die Vogelgesellschaften auf 4 Kontrollflächen derBrachystegia- und auf 2 derAcacia-Baumsavanne wurden während der Regenzeit (Februar 1976) in der Umgebung von Lusaka, Zambia, quantitativ erfaßt. Artenzahl und Artendiversität waren auf allen 6 Flächen fast gleich, obwohl das Laubvolumen der am reichsten strukturierten Fläche etwa den siebenfachen Betrag jenes der ärmsten ausmachte. In der Laubschicht nahrungssuchende Vögel beuteten das Gebiet mit dem geringsten Laubvolumen am stärksten aus. Die Dichte der paläarktischen Zugvögel, unter denen Fitis und Grauschnäpper am häufigsten waren, war niedrig; fast keine wurden im unberührten Wald angetroffen. In den vom Menschen stärker beeinflußten Abschnitten kamen sie dagegen häufiger vor. Veränderungen der Baumsavanne treten durch Ausholzung der Wälder oder auch durch intensives Beweiden der offenen Savannenflächen ein. Paläarktische Zugvögel machten stets nur einen sehr geringen Teil der Vogelgesellschaften aus, auch innerhalb des relativ begrenzten Spektrums afrikanischer Vögel gleichen Nahrungsverhaltens.Die Vogeldichte der zambischen Wälder ist im Vergleich zu Laubwaldbiotopen der gemäßigten Zone eher niedriger, obwohl z. B. die Primärproduktion derBrachystegia-Wälder ungefähr zweimal so hoch ist. In diesem Zusammenhang ist natürlich der stärkere Jahreswechsel in der gemäßigten Zone von Bedeutung. Auch fällt auf, daß die Vogeldichte in allen unseren Untersuchungen ziemlich ähnlich war, ungeachtet offensichtlich stark unterschiedlicher Primärproduktion ausgedrückt durch die Laubmasse.Artendichte und -diversität waren in den zambischen Untersuchungsgebieten beträchtlich höher als in entsprechend großen Flächen der gemäßigten Laubwälder. Verschiedene Diversitätsindices wurden berechnet, die verglichen mit der Artenzahl dafür sprechen, daß die zambischen Vogelgesellschaften zwischen einer broken-stick- bzw. lognormalen Verteilung der Arten liegen. Die erstere ist bei gut integrierten und stabileren Gesellschaften zu erwarten, letztere bei instabileren.Die paläarktischen Zugvögel traten in viel geringerer Dichte auf als in mäßig günstigen Brutbiotopen ihrer Heimat. Wahrscheinlich besitzen die festgestellten Arten, vor allem der Fitis, andere Winterquartiere. Die Tatsache, daß Paläarkten in vom Menschen beeinflußten Biotopen in beträchtlich größerer Dichte auftreten als in weniger gestörten, war zwar schon früher aufgrund qualitativer Beobachtungen behauptet worden, wurde aber hier erstmals quantitativ bestätigt. Daß Zugvögel sich besser in der Kulturlandschaft im weitesten Sinne behaupten können als im unberührten Wald, kann für ihr weiteres Schicksal sehr bedeutsam sein, da zunehmend größere Teile Afrikas einer Landschaftsumgestaltung verbunden mit Zerstörung ursprünglicher Vegetationseinheiten unterworfen werden.