Differences between banded thickets (tiger bush) at two sites in West Africa

Abstract. This paper deals with the influence of edaphic conditions on the spatial structure of banded thickets or tiger bush (brousse tigrée). It is based on two sites in West Africa, with similar climatic conditions but located on contrasting substrates. The spatial structure was described with standardized characteristics including thicket spacing, thicket/inter‐thicket contrast, upslope/downslope asymmetry and species zonation throughout the vegetation band. Recruitment and senescence features of woody stands were emphasized in order to understand current dynamics. Data were collected on transects oriented perpendicular to the contours and so to the thickets as well. A standardized analytical procedure was applied to data from both sites to ensure consistent and thorough delineation of thickets. The overall periodicity of thickets, the woody flora and the dominant species Combretum micranthum were similar at the two sites. However, thicket spacing, thicket/inter‐thicket contrast and upslope/downslope floristic asymmetry of the thickets were higher in the less favourable site. Also seedlings were less abundant, with a greater dependence on pre‐existing thickets. Not all banded vegetation systems show sharp contrasts and are strongly asymmetric, since such characteristics are likely to be reinforced by adverse environmental conditions. As a consequence, current dynamics may be more diverse than expected. Quantified inter‐site comparisons can greatly help to classify African banded vegetation systems and to discuss potential dynamic outcomes.     

Rain forest expansion mediated by successional processes in vegetation thickets in the Western Ghats of India

Aim The objective of this study was to document succession from grassland thickets to rain forest, and to provide evidence for their potential as restoration tools. Location The Linganamakki region (State of Karnataka) of the Central Western Ghats of India. Method We selected thirty vegetation thickets ranging from 4 to 439 m² in area in the vicinity of rain forest. The area of each small thicket was estimated as an oval using its maximum length and its maximum width. When the shape was irregular (mostly in large thickets) the limits of the thicket were mapped and the area calculated from the map. Plant species were identified, the number of individuals was estimated and their heights measured. Results There was a progression in the thickets from early to late successional species. Small thickets were characterized by ecotone species and savanna trees such as Catunaregam dumetorum. Savanna trees served as a nucleus for thicket formation. Colonizing species were mostly bird‐dispersed. As succession proceeded in larger thickets, the proportion of evergreen, late‐successional rain forest trees increased. The species composition of the large thickets differed depending on the species composition of reproductive adults in the nearby forested areas. The species within small thickets were also found in the large thickets. The nestedness in species composition suggested that species turnover was deterministic based on thicket size. Human disturbance (leaf and wood collection by the local populations) affected the species composition and the species–area relationship of thickets. Main conclusions Vegetation thickets are nodal centres for rain forest colonization within grasslands. They expand and replace savanna. Early successional bird‐dispersed species established around savanna trees followed by late‐successional rain forest trees dispersed from the nearby forest by birds. Restoration programmes that reproduce natural successional processes such as those observed in thickets will be more successful and less expensive than the methods currently being employed, where trees are individually planted in grassland. Wood harvesting is the only factor that prevents thicket growth and coalescence and hampers forest expansion.     

Spatial and temporal growth dynamics of Barrier Island shrub thickets

Spatial and temporal dynamics of barrier island shrub thickets were investigated on Hog Island, a barrier island along the Eastern Shore peninsula of Virginia. The island thickets are dominated by the actinorhizal shrub, Myrica cerifera. On the accreting, northern half of the island, thickets expanded as swale area increased. Analysis of aerial photographs revealed that total thicket area increased by nearly 400% from 1949 to 1989, paralleling a substantial increase in area for the northern portion of the island. Thicket characteristics varied with the age and position of the three field sites on Hog Island. At the youngest site examined (20 yr), recruitment and shrub growth were high. Shrub growth remained high and recruitment continued in the stable central portions of the island (50 yr). However, at the oldest site near a bay side salt marsh (≈130 yr), recruitment was not evident, shrub growth was reduced, and mortality was most apparent. Several “life stages” for shrub thickets were identified relative to island accretion. After the stabilization of new land, seedlings establish and grow rapidly to form a thicket. Once established, a thicket rapidly expands in the relatively protected, mesic swale environment. Thicket decline follows, characterized by an increase in vine growth on and around the shrubs and the formation of gaps as the thicket breaks up. Spatial variation in M. cerifera thickets on Hog Island is related to length of time soils have been stabilized and, especially, to time since thicket establishment.     

Population dynamics of tundra voles in relation to configuration of willow thickets in southern arctic tundra

The areal extent and configuration of thickets of willow shrubs are currently changing in the Arctic both as an effect of global warming and changed browsing pressure of reindeer. These changes have been predicted to impact the distribution and abundance of wildlife species relying on willow thickets as habitat. We assessed the relation between variables quantifying willow thicket configuration and population dynamics of tundra voles (Microtus oeconomus) in three riparian regions in Finnmark, northern Norway, which were subject to intense browsing by semi-domesticated reindeer. The tundra vole, which exhibits 5-year population cycles in Finnmark, is the dominant small rodent species in riparian landscape elements in southern arctic tundra. In the course of a 4-year trapping study, tundra vole populations went through the cyclic phases of increase, peak and crash, however, with distinct differences between the three regions in the population dynamics. Within regions, the occupancy pattern during the increase phase was positively related to willow thicket configuration (in particular edge density and willow height) only in the region attaining the highest abundance and occupancy. However, local abundance was not clearly related to habitat features within any regions. The lack of consistency in the response of tundra vole populations to willow thicket configuration, as well as the positive relation between the degree of thicket shredding and tundra vole habitat occupancy in one of the regions, indicates that tundra voles will not be much affected by climate or browsing induced changes in the shrubbiness of the tundra in the future.     

Establishment of Broad‐leaved Thickets in Serengeti,Tanzania: The Influence of Fire,Browsers, Grass Competition,and Elephants1

The role of Euclea divinorum in the establishment of broad‐leaved thickets was investigated in Serengeti National Park, Tanzania. Thickets are declining due to frequent fires, but have not reestablished when fires have been removed. Seedlings of E. divinorum, a fire‐resistant tree, were found in grassland adjacent to thickets and as thicket canopy trees and may function to facilitate thicket establishment. Seedlings of thicket species were abundant under E. divinorum canopy trees but not in the grassland, indicating that E. divinorum can facilitate forest establishment. We examined E. divinorum establishment in grassland by measuring survival and growth of seedlings with respect to fire, browsers, elephants, and competition with grass. Seedling survival was reduced by fire (50%), browsers (70%), and competition with grass (50%), but not by elephants. Seedling growth rate was negative unless both fire and browsers, or grass was removed. Establishment of thickets via E. divinorum is not occurring under the current conditions in Serengeti of frequent fires, abundant browsers, and dense grass in riparian areas. Conditions that allowed establishment may have occurred in 1890–1920s during a rinderpest epizootic, and measurements of thicket canopy trees suggest they established at that time.     

Spatial/temporal variations in shrub thicket soil seed banks on an Atlantic Coast barrier island

Potential species replacement within low-diversity shrub thicket communities was investigated for a Virginia barrier island. Seed bank species composition was quantified in a glasshouse study using soil samples collected beneath closed Myrica cerifera thickets, as well as from thicket gaps. Samples were collected from productive and aging thickets, corresponding to differences in soil age. These data were compared to species presently occurring within the thickets and gaps. Seedbank species composition was not indicative of current community composition for either the intact thickets or the gaps. Seed banks resembled a more pioneer community. Thirteen families, 23 genera, and 25 species were identified from the seed bank beneath the M. cerifera thickets. Four species were woody. The within-gap seed bank included 19 families, 30 genera, and 34 species. Eight species were woody. The current community included 21 families, 33 genera, and 36 species beneath the intact thickets as well as within the thicket gaps. Eighteen species were woody. The species richness of gaps was more than three times that of intact thickets. For low-diversity shrub thickets, gaps enhance species richness.     

鄱阳湖沙地蔓荆灌丛沙堆形态特征及空间分布格局

目前灌丛沙堆研究主要集中在干旱半干旱区的沙质草原和沙漠边缘,对亚热带湿润区灌丛沙堆的形成、演变过程并不清楚。以鄱阳湖沙地为研究区,通过样方调查和地统计学的方法,研究不同沙化程度下蔓荆灌丛沙堆的形态特征及分布格局。结果表明:鄱阳湖沙地蔓荆沙堆的形态以盾形为主,其冠幅变化幅度为1.2—18.2 m~2,固定和半固定沙地显著高于流动沙地;对灌丛沙堆的形态参数来说,其长轴与短轴在固定和半固定沙地上呈极显著的线性相关关系,流动沙地上呈二次函数关系;半固定和流动沙地上沙堆底面积与沙堆高度呈二次函数关系(r0.6);3种类型沙地上灌丛底面积与沙堆体积之间极显著线性相关,其中半固定沙地线性函数的斜率最大;除固定沙地的沙堆高度和半固定沙地的灌丛高度外,3种沙地上蔓荆灌丛与沙堆的其他形态参数间均极显著相关,说明随着沙地的固定,蔓荆灌丛有利于沙堆水平尺度的增长;3种沙地上蔓荆沙堆均呈随机分布。     

Uganda kob prefer high-visibility leks and territories

In lekking species, where males provide estrous females withlittle more than sperm, it has been widely supposed that theonly possible benefits to females of mate choice are genetic.We studied female choice of leks and territories in a reduncineantelope, the Uganda kob (Kobus kob thomasi), and found thatfemales consistently preferred high-visibility mating sites.Leks were elevated and had shorter grass and fewer thicketsthan the surrounding areas. Changes in the number of male andfemale kob on 10 leks were correlated with changes in surroundinggrass height, and both females and males preferred leks withexperimentally reduced grass height over neighboring controls.Within a lek, territory popularity was the primary determinantof male daily mating success, and females preferred territoriesrelatively far from thickets, but removal of thickets did notaffect female territory preferences. Because lion hunting successon kob increases with grass height and thicket density, femalesmay benefit directly from these preferences by reducing therisk of predation.     

Tree species diversity in small, tropical riparian forest fragments in Belize, Central America

Tree species diversity was measured in a network of very small galleryforests within the Mountain Pine Ridge savanna in Belize. Research focussed onforest patches smaller than 1 ha in size (micro-forests) and linearstrips of trees along creeks lacking interior core zones with low understoreylight levels (tree thickets). Twenty-five micro-forests and 51 tree thicketsites were sampled throughout the savanna. A total of 144 morphospecies 5cm dbh (106 in micro-forests and 117 in tree thickets) werefound, which represents 1/5 of the approximately 700 native tree species in Belize.Most (85.3%) of the species encountered are typically found in tropical rainforests and few are restricted to savanna or riparian environments. Speciesaccumulated at a much faster rate in micro-forests than in tree thickets. Onlyone species, the palm Acoelorraphe wrightii, was extremelyabundant, accounting for almost 30% of all stems. Many of the species werepresent in very low densities: 19% of all species found in micro-forests and 42%of those found in tree thickets had on average one or fewer stems per hectare. Alarge proportion of species were also found infrequently across the landscape,being present in only 36% of micro-forests and at 52% of tree thicket sites. Theresults indicate that networks of very small forest patches can contain highnumbers of species and could therefore contribute to the maintenance of regionalbiodiversity.     

Shrub expansion stimulates soil C and N storage along a coastal soil chronosequence

Expansion of woody vegetation in grasslands is a worldwide phenomenon with implications for C and N cycling at local, regional and global scales. Although woody encroachment is often accompanied by increased annual net primary production (ANPP) and increased inputs of litter, mesic ecosystems may become sources for C after woody encroachment because stimulation of soil CO₂ efflux releases stored soil carbon. Our objective was to determine if young, sandy soils on a barrier island became a sink for C after encroachment of the nitrogen‐fixing shrub Morella cerifera, or if associated stimulation of soil CO₂ efflux mitigated increased litterfall. We monitored variations in litterfall in shrub thickets across a chronosequence of shrub expansion and compared those data to previous measurements of ANPP in adjacent grasslands. In the final year, we quantified standing litter C and N pools in shrub thickets and soil organic matter (SOM), soil organic carbon (SOC), soil total nitrogen (TN) and soil CO₂ efflux in shrub thickets and adjacent grasslands. Heavy litterfall resulted in a dense litter layer storing an average of 809 g C m^?2 and 36 g N m^?2. Although soil CO₂ efflux was stimulated by shrub encroachment in younger soils, soil CO₂ efflux did not vary between shrub thickets and grasslands in the oldest soils and increases in CO₂ efflux in shrub thickets did not offset contributions of increased litterfall to SOC. SOC was 3.6–9.8 times higher beneath shrub thickets than in grassland soils and soil TN was 2.5–7.7 times higher under shrub thickets. Accumulation rates of soil and litter C were highest in the youngest thicket at 101 g m^?2 yr^?1 and declined with increasing thicket age. Expansion of shrubs on barrier islands, which have low levels of soil carbon and high potential for ANPP, has the potential to significantly increase ecosystem C sequestration.     

Cross-Scale Patterns in Shrub Thicket Dynamics in the Virginia Barrier Complex

Abstract To interpret broad-scale erosion and accretion patterns and the expansion and contraction of shrub thickets in response to sea level rise for a coastal barrier system, we examined the fine-scale processes of shrub recruitment and mortality within the context of the influence of ocean current and sediment transport processes on variations in island size and location. We focused on Myrica cerifera shrub thickets, the dominant woody community on most barrier islands along the coastline of the southeastern USA. Observations suggest that M. cerifera, a salt-intolerant species, is increasing in cover throughout the Virginia barrier islands, yet rising sea level in response to climate change is increasing erosion and reducing island area. Our objective was to explain this apparent paradox using pattern–process relationships across a range of scales with a focus on ocean currents and sediment transport interacting with island characteristics at intermediate scales. Multi-decadal comparisons across scales showed a complex pattern. At the scale of the entire Virginia barrier complex, modest decreases in upland area were accompanied by large increases in shrub area. Responses were more variable for individual islands, reflecting inter-island variations in erosion and accretion due to differences in sediment transport via ocean currents. Several islands underwent dramatic shrub expansion. Only for within-island responses were there similarities in the pattern of change, with a lag-phase after initial shrub colonization followed by development of linear, closed canopy thickets. Understanding the fine-scale processes of shrub seedling establishment and thicket development, in conjunction with the influence of ocean currents and sediment transport, provides a framework for interpreting island accretion and erosion patterns and subsequent effects on shrub thicket expansion or contraction across scales of time and space.     

Kenyan endemic bird species at home in novel ecosystem

Riparian thickets of East Africa harbor a large number of endemic animal and plant species, but also provide important ecosystem services for the human being settling along streams. This creates a conflicting situation between nature conservation and land‐use activities. Today, most of this former pristine vegetation is highly degraded and became replaced by the invasive exotic Lantana camara shrub species. In this study, we analyze the movement behavior and habitat use of a diverse range of riparian bird species and model the habitat availability of each of these species. We selected the following four riparian bird species: Bare‐eyed Thrush Turdus tephronotus, Rufous Chatterer Turdoides rubiginosus, Zanzibar Sombre Greenbul Andropadus importunus insularis, and the Kenyan endemic Hinde′s Babbler Turdoides hindei. We collected telemetric data of 14 individuals during a 2 months radio‐tracking campaign along the Nzeeu River in southeast Kenya. We found that (1) all four species had similar home‐range sizes, all geographically restricted and nearby the river; (2) all species mainly use dense thicket, in particular the invasive L. camara; (3) human settlements were avoided by the bird individuals observed; (4) the birds' movement, indicating foraging behavior, was comparatively slow within thickets, but significantly faster over open, agricultural areas; and (5) habitat suitability models underline the relevance of L. camara as suitable surrogate habitat for all understoreyed bird species, but also show that the clearance of thickets has led to a vanishing of large and interconnected thickets and thus might have negative effects on the population viability in the long run.     

Shifts in litterfall and dominant nitrogen sources after expansion of shrub thickets

Woody encroachment into herbaceous ecosystems is emerging as an important ecological response to global change. A primary concern is alterations in C and N cycling and associated variations across a variety of ecosystems. We quantified seasonal variation in litterfall and litter N concentration in Morella cerifera shrub thickets to assess changes in litterfall and associated N input after shrub expansion on an Atlantic coast barrier island. We also used the natural abundance of ¹⁵N to estimate the proportion of litterfall N originating from symbiotic N fixation. Litterfall for shrub thickets ranged from 8,991 ± 247 to 3,810 ± 399 kg ha⁻¹ year⁻¹ and generally declined with increasing thicket age. Litterfall in three of the four thickets exceeded previous estimates of aboveground annual net primary production in adjacent grasslands by 300–400%. Leaf N concentration was also higher after shrub expansion and, coupled with low N resorption efficiency and high litterfall, resulted in a return of as much as 169 kg N ha⁻¹ year⁻¹ to the soil. We estimated that ∼70% of N returned to the soil was from symbiotic N fixation resulting in an ecosystem input of between 37 and 118 kg ha⁻¹ year⁻¹ of atmospheric N depending on site. Considering the extensive cover of shrub thickets on Virginia barrier islands, N fixation by shrubs is likely the largest single source of N to the system. The shift from grassland to shrub thicket on barrier islands results in a substantial increase in litterfall and foliar N concentration that will likely have a major impact on the size and cycling of ecosystem C and N pools. Increasing C and N availability in these nutrient-poor soils is likely to permanently reduce cover of native grasses and alter community structure by favoring species with greater N requirements.     

High-latitude<Emphasis Type="Italic"> Acropora cervicornis</Emphasis> thickets off Fort Lauderdale,Florida, USA

Baseline studies conducted in 1998 document the presence of robust, non-reef-building Acropora cervicornis thickets in shallow (3–7 m depth), near-shore waters off the coast of Fort Lauderdale, Florida, USA. These thickets thrive in a high-latitude environment, the northernmost in the continental USA, in the midst of potential anthropogenic stressors. Within thickets, the spatial variation in mean percent coral cover, macroalgal cover, scleractinian species richness, density of A. cervicornis juveniles, and the density and size of A. cervicornis colonies and fragments were recorded. Thicket size ranged between ~0.1 and 0.8 ha and mean coral cover varied between ~5 and 28%, with A. cervicornis accounting for ~87–97% of all scleractinians. Mean A. cervicornis colony and fragment densities per thicket were 1.3–3.3 colonies m^–2 and 0.7–2.8 fragments m^–2, respectively. Recruit densities varied between 0 and 1 ind. m^–2. White band disease was detected at all thickets, with a mean A. cervicornis colony surface area affected of 1.8%. For all thickets, densities of the corallivorous polychaete Hermodice carunculata ranged between ~18 and 86 ind. ha^–1, with predation scars affecting <0.2% of the A. cervicornis cover. These flourishing A. cervicornis thickets off Fort Lauderdale provide an interesting counterpoint to the declining and disease-stricken A. cervicornis populations reported in the Florida Keys and wider Caribbean.     

How Fast Can Carbon Be Sequestered When Restoring Degraded Subtropical Thicket?

Is the maximum rate of carbon sequestration reported for the CAM‐C₃ plant Portulacaria afra (spekboom), viz. 15.4 t CO₂ ha^?1 yr^?1, unusual in comparison with other plants with similar physioliogies, or could such rates be expected routinely in restoration with P. afra? Private sector investors in thicket restoration need an answer to this question in order to assess the feasibility of using carbon finance as the main income stream from their investments. A literature review showed that 15.4 t CO₂ ha^?1 yr^?1 is not an unusual rate of carbon sequestration for CAM plants in arid and semi‐arid environments, which suggests that investors in thicket restoration should not consider this an outlier value. The results also suggest that carbon finance could be used to fund restoration using other CAM plants in degraded xeric thickets in countries such as Argentina, Chile, Mexico, and Madagascar.     

Nightingales Luscinia megarhynchos in scrub habitats in the southern fens of East Anglia, England: associations with soil type and vegetation structure

The southeast corner of the East Anglian Fens supports a large concentration of Nightingale Luscinia megarhynchos territories. A total of 382 territories were located in extensive surveys in 1999 and 2000, probably representing over 5% of the English population. Transect counts revealed that the Fenland population is restricted in distribution and is associated with localized thickets of scrub. The highest densities are found on humus‐rich soils, suggesting that soil type, in addition to habitat availability, may have a strong influence on the Nightingale's distribution in this region. This paper provides the first assessment of habitat requirements of the Nightingale in scrub, which now forms a principal habitat for the species in England. Detailed studies of the attributes of over 100 Nightingale territories revealed subtle differences in the vegetation structure of these thickets when compared with paired, unoccupied, but apparently similar thickets. The Nightingale territories tended to have a higher proportion of bare ground or short vegetation in the field layer under the canopy, whereas paired sites were more likely to have low field layer vegetation beneath the canopy. The bare ground within the thickets is a feature of shading beneath very dense foliage cover. Within Nightingale territories, low field layer volume and shrub twig volume at the thicket edges was higher than in unoccupied thickets. The differences detected in vegetation structure suggest that a dense and continuous canopy forming a shell over bare ground but with dense low foliage at thicket edges provides the ideal vegetation structure for Nightingales in scrub habitats. Our study suggests that Nightingales occupy scrub of a very specific structure, and specific stage in vegetation succession. This structure probably provides an optimal combination of foraging habitat, microclimate and cover from predators. It is suggested that humus‐rich soils may be preferred because they may support a particularly rich source of invertebrate food, but this remains to be tested empirically.     

The impact of beech thickets on biodiversity

Beech bark disease has dramatically altered hardwood forest structure and composition across northeastern North America. Extensive overstory mortality has resulted in prolific root-sprouting in some stands leading to the development of understory thickets of clonal small-stemmed beech. Beech thickets may impact local forest biodiversity, but this has not been adequately evaluated. We hypothesized significant differences in diversity of groundcover flora, craneflies, amphibians, and small mammals between plots with and without beech thickets. Paired plots were established in uneven-aged northern hardwood forest stands with no recent management history at two sites in the Adirondack Mountains of New York State. Groundcover plants, terrestrial craneflies, amphibians and small mammals were sampled on twenty paired plots. Discriminant analysis showed a significant difference between thicket and non-thicket (control) areas; significant variables in plot type separation were beech sapling abundance, leaf litter depth, and coarse woody debris volume. Groundcover plant cover, richness, and diversity were significantly lower in thicket compared to non-thicket plots, while beech sapling density explained 17–38 % in groundcover plant species diversity. There were no significant differences between the diversity of cranefly, amphibian and small mammal communities of each plot type. Beech thickets are important determinants of local biodiversity.     

Nitrogen-fixers <Emphasis Type="Italic">Alnus</Emphasis> and <Emphasis Type="Italic">Lupinus</Emphasis> influence soil characteristics but not colonization by later successional species in primary succession on Mount St. Helens

Changes to the primary successional environment caused by colonizing plants that present symbiotic associations with nitrogen-fixing bacteria were investigated at two areas on Mount St. Helens. One area was occupied by alder (Alnus viridis) thickets and old lupine (Lupinus lepidus) patches and the other area by young lupine patches and pumice barrens. Alder thicket soils had higher levels for a few soil nutrients and had greater cover by other pioneer species as compared to old lupine patches. Many soil nutrients, including nitrogen and soil organic matter, were below detection limits in old lupine patches but not in alder thicket soils. Young lupine patch soils were generally not different from barren site soils but had greater cover by other pioneer species. Below detection nitrogen and soil organic matter levels also occurred in many barren soil samples but not in young lupine patch soils. Barren soils were moister than were the other sites. The apparent increase in soil fertility has not led to invasion by later successional species, perhaps due to dry conditions or to other inhibitory factors. Seedbanks, composed of early successional species, appear to be developing in these areas.     

VAM association in the shrub Myrica cerifera on a Virginia,USA barrier island

A combined laboratory and field study examined the potential for a symbiotic association between the actinorhizal shrub Myrica cerifera and vesicular arbuscular mycorrhizal (VAM) fungi on a Virginia barrier island. M. cerifera seedlings and two test species, Zea mays and Strophostyles umbellata, were grown in an environmental chamber on soils collected from four sites differing in soil age (< 5 to over 130 years), salinity (1–35 g/g total soil chloride), and edaphic characteristics. Seedling root infection was significantly lower for all three species in the youngest soils from the beach where salinity was highest. Stained M. cerifera roots revealed all the components for a functional VAM association; however, there were significantly fewer arbuscules and vesicles relative to the test species. Among field-collected M. cerifera, infection was not detected in mature shrubs from the bay side of the island, where M. cerifera thickets were in a state of degeneration. Infection was highest in soils from the young, developing thickets, and in the most stable thickets of the island interior. Despite the dynamic nature of the barrier island environment, VAM associations with M. cerifera appear to be present, especially in seedlings and developing shrub thickets.     

Keywords: ornithochory; fleshy fruit; succession; species losses; cultural biotope.

Abandonment of traditional agricultural practices in fruit orchards on hillsides in Central Germany results in successive changes in vegetation. We examined three hypotheses relating to these changes: (1) thickets of fleshy-fruited plants develop around planted trees as a result of ornithochory and local soil and site amelioration, (2) woody plants have long-term effects on soil fertility, and (3) thicket development reduces the plant-species diversity of semi-dry grassland between the trees. Field observations and nursery experiments supported the first and second hypotheses. Increased soil fertility, a seed rain of fleshy-fruited shrubs, and shrub establishment occurred mainly around planted fruit trees. Soils from old tree-planting sites remained fertile after the trees had died and disappeared. The third hypothesis was rejected because no decrease in species' richness or diversity occurred during the initial stages of thicket formation. Nevertheless, herbaceous plant species characteristic of the Festuco-Brometea community were absent from the vegetation and seed bank of shaded, nutrient-enriched sites. Conservation of semi-dry grasslands following orchard abandonment will therefore require active control of woody plants.