Phytoclimatic characterization ofQuercus frainetto Ten. stands in peninsular Italy

The floristic differentiation of the deciduousQuercus frainetto forests along the climatic gradients of Apennine Italy has been studied. The ecological amplitude of this oak, and the bioclimatological relationships here assessed, suggest potentiality for the growth ofQ. frainetto-rich communities as broad zonal vegetation belt, ranging from NW to SE along the W side of peninsular Italy. Strong floristical and ecological similarities to the balcanic stands are described.The status of real vegetation belt for the ItalianQuercion frainetto s.l. is here emphasized.     

Hybrid Zones Between Two European Oaks: a Plant Community Approach

Phenomena of hybridization can affect the ecology and evolution of the species involved in the process, as well as their communities. Although numerous papers focus upon the problem of taxonomy, few of these have attempted to study hybrid zones in relation to the analysis of their communities. On the Iberian Peninsula, hybridization phenomena among different oak species are frequent. It is, however, between Quercus faginea Lam. and Quercus pubescens Willd. where the most noteworthy hybridization phenomenon occurs. In this respect, we are familiar with the existence of different introgression levels but we are unaware of whether these hybrids are the transitory result of the interspecific genetic flow or whether these are maintained by means of extrinsic selection processes. Study of plant communities’ flora and environment might shed light upon this issue. Comparison between plant communities dominated by one of the parental species and those dominated by individuals of hybrid origin might enable us to establish the presence or absence of an environment that is potentially selective in favour of the hybrids. Thus the possible existence of extrinsic selection. Furthermore, this information will help us to understand plant community distribution in an area␣that is difficult to interpret. To this purpose, we used multivariate ordination techniques (DCA and CCA) based upon a total of 395 floristic releves covering the whole range of the parental species on the Iberian Peninsula and upon climatic and edaphic variables for each of these releves. We also compared the groups obtained in relation to floristic similarity (Jaccard index), richness and diversity (Shannon–Weaver index). Forests associated with Quercus pubescens are related to heavy summer precipitation, whereas Quercus faginea forests correspond to lower values of this variable and higher ones for continentality. Between both formations, there is a broad hybrid zone, with diffused borders that are related to an environmental gradient of Mediterranean influence. In this region, two types of forest communities were distinguished, which enabled us to divide the hybrid zones into two territories. Our results allowed us to locate the hybrid zone in an ecotone. The differentiation between habitats appears to indicate models of ecological selection. These models require, by definition, the presence of an environmental gradient between the parental zones. We are, however, aware of the need for future experiments in order to establish whether the hybrids are better adapted than the parental species. Only with availability of all this information can intrinsic selection be rejected.     

Notes on Quercus ilex L. in Liguria (NW Italy)

The climatic characteristics of the Liguria region have been outlined by means of raw data and derived indices (water balance related to potential evapotraspiration, Rivas-Martinez's index of mediterraneity and thermicity, De Martonne's index of dryness, and Emberger's pluviothermic quotient and index of summer dryness). Their interpretation suggests that Liguria is a boundary region between two different climatic areas: the Mediterranean and that of Central Europe.The distribution of Quercus ilex communities in Liguria have suffered the heavy consequences of human activity on the coastal belt of the region. There is, however, sufficient evidence to show that their presence is closely linked to climatic conditions. Quercus ilex communities are absent from the western coastal belt where the annual water balance is below –50 mm and high mean temperatures occur. They are also absent from marly limestone dominated areas where annual water balance is below 300 mm. Quercus ilex woods show a preference for water balance values between 0 and 600, with the better developed forests being found in central and eastern coastal Liguria.The Ligurian Quercus ilex woods are ascribed to Quercetum ilicis Br.-Bl. 1915. Given that the presence within their floristic composition of a group of species of Querco-Fagetea (more species in eastern stands, few in western ones) is an almost constant characteristic, the subass. fraxino-ostryetosum Mariotti 1984 can be retained for many of them. Quercus ilex is not present in the initial succession stages of Mediterranean Liguria vegetation. It can be found in later stages when the vegetation is more fully developed towards a maquis type structure. In this case it tends to become dominant and the community develops (if no fire or other human interference occurs) into a Quercus ilex forest. Frequently, this succession takes place under an old tree layer cover (generally Pinus sp.), a remanant of man's past management of the vegetation in the area.     

Quantitative analysis of acorn transportation by rodents using magnetic locator

Transported distance and mortality of acorns scattered by rodents were investigated with magnets inserted into acorns (40 of Quercus serrata and 20 of Quercus acutissima) and a magnetic locator in a natural forest stand. All the treated acorns were transported, and 60% of them were discovered again with a magnetic locator from autumn to the next spring. Most transported acorns suffered predation within one month after the start of the experiment. Several acorns were rehoarded at least two or three times. Average transported distance of scattered acorns was 22.1±8.9 m (max=38.5 m) and the survival rate was 3.0%. The magnet method is one of the most effective methods for tracking acorns through the winter.     

Successional trends and apparent <Emphasis Type="Italic">Acer saccharum</Emphasis> regeneration failure in an oak-hickory forest in central Missouri,USA

Permanent vegetation plots established in 1968 at the Baskett Research and Education Area (BREA) were remeasured in 2004/2005 to assess trends in canopy composition, vegetation–environment relationships, and regeneration patterns. Quercus spp. [particularly Quercus alba (white oak)] remained dominant in BREA forests. However, Nonmetric Multidimensional Scaling Ordination of a dataset composed of tree importance values from both samples indicated the two most common community types (Dry Ridge and Slope; Mesic Slope) were shifted away from Quercus spp. abundance toward greater importance of Acer saccharum (sugar maple) in ordination space. Plots in glade-like environments shifted toward greater Juniperus virginiana (eastern redcedar) importance. Environmental analysis indicated that soil properties (pH, organic matter, Ca, Mg, and K) and slope position were highly correlated with ordination axes. Saplings of Quercus spp. exhibited greatly reduced abundance in the most recent sample and the reduction in Quercus spp. density was highest in plots with greatest canopy cover. While Acer saccharum saplings were still quite abundant in BREA forests, smaller sapling size classes were becoming notably depleted, reflecting growth of a cohort of plants established before the 1968 sample toward and into tree-sized individuals. Seedlings surveys emphasized that A. saccharum seedlings have become very rare in BREA forests, while Quercus spp. seedling densities are higher, but likely below those needed for eventual adequate recruitment to the canopy. Although a number of mechanisms to explain apparent A. saccharum regeneration failure can be developed, herbivory, recent climate trends (warming late-winter conditions) and increased litter layer depth are the most plausible explanations for the observed scarcity of A. saccharum seedlings. Although increasing importance of A. saccharum in the canopy of BREA forests appears inevitable in coming decades, longer-term trends are unclear given the shortage of seedling regeneration.     

Spectral composition of shade light in coastal-zone oak forests in SE Bulgaria,and relationships with leaf area index: a first overview

Major knowledge gaps exist with respect to light-quality regimes in the coastal-zone Strandzha Quercus frainetto (Q.f.) forest region adjoining the southern Bulgarian Black Sea. This paper presents preliminary results that help narrow these gaps. In conjunction with leaf area index (LAI) field campaigns we undertook measurements with an array of 7 broad-band (ca 40 nm) sensors covering the range 0.40–0.94 μm, plus 1 sensor for UVB (0.297 μm peak) and 1 for photosynthetically active radiation (PAR). Measurements focused on inside-forest shade conditions at sites 0 to ca 15 km from the Black Sea and at altitudes up to ca 120 m above sea level. Some of the sites were also studied using a high-resolution spectroradiometer. A sequential measuring strategy was necessary. This involves potentially large uncertainties, here addressed through estimations of the variability around the sinusoidal course of daylight. Light-quality regimes were found to be in general support of earlier studies of deciduous forests. Our data from the broad-band sensors and from the spectroradiometer are mutually supportive. They indicate a stronger red-shift below Q.f. canopies than below canopies in enclaves dominated by Fagus orientalis and Pinus sylvestris. Transmission in the range 0.50–0.55 μm increases beneath the three types of canopies, most pronounced in the Q.f. case. Analysis of relationships between the inside-forest to open-field irradiance ratio and LAI supports the use of Beer’s Law. We found a fairly strong relationship between the red (0.66 μm) to far-red (0.73 μm) irradiance ratios (R/FR) and LAI for the Q.f. forest. In quantitative terms, the result is new for this Q.f. region, and suggests further research to explore whether a two-sensor approach (0.66 and 0.73 μm) might offer possibilities for further low-cost mapping of the spatio-temporal patterns of R/FR and LAI in Strandzha. Such mapping would assist in further studies of the region’s forest biogeochemistry and vitality.     

Not only size matters: Acorn selection by the European jay (Garrulus glandarius)

A strong selection for acorn characteristics is expected to have evolved in the mutualistic relationship between the European jay (Garrulus glandarius) and the oak (Quercus spp.). Bossema's pioneer work suggested that jays do not select acorns randomly, but rather they preferentially select some size and species. Preference for some seeds over others may have implications on plant community dynamics by conferring advantages (or disadvantages) on the selected (avoided) seed characteristics. In this paper we test to what extent jays select acorns by species and/or by size and the relation between these two traits in Mediterranean oak species. The experiments consist of a set of field tests in which acorns from four different coexisting Mediterranean oak species (Quercus ilex, Quercus faginea, Quercus suber, and Quercus coccifera) were placed in artificial feeders accessible to wild jays. The acorns were previously measured to control individual acorn characteristics. Using video-recording techniques, we followed jay activity and the fate of each acorn (sequence of acorn selection and method of transport). Q. ilex acorns were preferred over other acorns, and Q. coccifera acorns were avoided when other acorns were available. Preference for Q. faginea and Q. suber acorns was intermediate, that is, they were preferred over Q. coccifera acorns but not over Q. ilex acorns. Large acorns were also preferred although acorn species selection was stronger than size selection. Jays selected species and size both by visual means and by using acorn area as an indicator of size. Acorns wider than 17–19 mm were carried in the bill because of throat limitation. Our results confirm Bossema's study on temperate oaks and extend it to Mediterranean oak species, revealing implications on mixed oak forest dynamics.     

Guild structures of seed insects in relation to acorn development in two oak species

We investigated patterns of acorn growth in Quercus variabilis Blume and Quercus serrata Thunb., seasonal trends in emergence of seed insects found in the acorns, oviposition periods of the insects and falling periods of insect-infested acorns. In Q. variabilis, two insect guilds were associated with acorn development: (i) the immature acorn feeding (IAF) guild [Curculionidae sp., Poecilips cardamomi (Schaufuss), and Characoma ruficirra (Hampson)]; and (ii) the mature acorn feeding (MAF) guild [Curculio robustus (Roelofs), Curculio sikkimensis (Heller), and Cydia glandicolana (Danilevsky)]. In Q. serrata, there were three guilds: (i) the pistillate flower feeding (PFF) guild (cynipid wasp); (ii) the IAF guild [sap absorption by Mechoris ursulus (Roelofs)]; and (iii) the MAF guild (M. ursulus, C. sikkimensis, Cydia danilevskyi (Kuznetzov), C. glandicolana and Autostichidae sp.). The succession of guilds during acorn development may be a consequence of the use by different species of the limited food resource. The lack of a PFF guild in Q. variabilis that was found in our field site would have a positive effect on IAF guilds in utilizing the acorns.     

A phenetic study of the taxonomic delimitation of Quercus acutifolia and Q. conspersa (Fagaceae)

A phenetic study ofQuercus acutifolia andQ. conspersa (Fagaceae) is carried out. The study was based on morphological characters, and included cluster analysis and principal component analysis. The results revealed that the characters employed so far to separateQuercus acutifolia andQ. conspersa seem to be quite unreliable. Nevertheless, both species can still be recognized as taxonomically different entities, although mostly based on characters of leaf margins. We propose that the individuals that have leaves with dentate-aristate margins correspond toQ. acutifolia, whereas those that have leaves with entire and not dentate-aristate margins belong toQ. conspersa.     

Riverscape-level patterns of riparian plant diversity along a successional gradient, Willamette river, Oregon

Riparian forest communities dominated by Populus balsamifera ssp. trichocarpa L. (Torr. and Gray ex Hook.) Brayshaw are important contributors to biodiversity in terrestrial and aquatic ecosystems of the Western United States. Species composition along a successional gradient from stand initiation to late-succession of P. balsamifera-dominated riparian forests was investigated along 145 km of the Willamette River, Oregon. There were 151 total species encountered across 28 stands and a mean species richness of 33.3 species per stand. Young stands were dominated by P. balsamifera and Salix tree spp. and opportunistic herbaceous species. Understory trees, shrubs, and herbaceous species as well as late-successional tree species established 12–15 years after stand initiation. Fraxinus latifolia Benth. was the dominant late-successional tree species. Vertical structural diversity, P. balsamifera mean diameter at breast height, large tree biomass, and stand age were strongly correlated with understory species presence and abundance based on non-metric multidimensional scaling (NMS) ordination. There were no young stands on mid and high terraces and this was reflected in geomorphic position being strongly correlated with the stand age gradient. Abundance of Phalaris arundinacea L. an invasive grass species, was also significantly correlated with plant species composition and abundance. This study indicates that Willamette River riparian forests are diverse and therefore important to the biodiversity of the Willamette River valley and that their presence as a mosaic of communities of different successional stages may be threatened by human interventions, including influences exerted by introduced plant species.     

Biochemical symptoms of stress in the mycorrhizal roots of Norway spruce (Picea abies)

Summary The mycorrhizal activity of spruce in a mixed-wood forest was monitored over 1 year by measuring biochemical characters in fine roots of six canopy trees and of a regrowth stand. The concentration of adenosine 5-triphosphate (ATP), a measure of living biomass, showed two peaks per year, one at bud break and one after main shoot growth. The concentration of storage polysaccharides in mycorrhizae showed the same cycles even more pronouncedly. It is proposed that these changes reflect growth and senescence of mycorrhizae and that the timing of the cycles is controlled by translocation of assimilates from the shoot. Differences between mycorrhizae collected from canopy trees and the regrowth stand were small and not significant. Characters known to be related to fungal activity of the mycorrhizal symbiosis (concentration of trehalose, glucose uptake, respiration) also varied little among the six canopy trees. Large differences among fine-root samples from different canopy trees, however, were detected in the concentrations of ATP and storage polysaccharides, measures which seemed to be physiologically integrated within trees. If low concentrations in roots precede losses of foliage from trees, these two symptoms could be used as early indicators of growth decline in individual spruce trees.     

纳塔栎和柳叶栎对铅锌矿区污染土壤的修复潜力分析:田间试验

通过野外调查、采样和实验室分析方法,研究优新景观树种纳塔栎（Quercus nuttallii）和柳叶栎（Quercus phellos）对湖南郴州Pb、Zn矿区复合污染土壤的适应性和修复潜力。试验设计A、B、E区种植1年生纳塔栎,C、D区种植1年生柳叶栎,1年后测量株高、地径、生物量等生长指标,随机采集植物全株及根际土壤,测试植物根际土壤和树木组织中的重金属含量。试验结果:Pb、Zn矿区土壤受到重金属Cd、Pb、Zn和As的复合污染,不同区域的土壤表现出污染异质性,采用单污染指数（Pi）和内梅罗指数（P_N）评价不同地块的污染程度,A区尾矿库（P_N=20.08）和B区（P_N=3.14）为重度复合污染,C区（P_N=2.43）为中度复合污染,D区（P_N=1.55）和E区（P_N=1.07）为轻微污染。纳塔栎和柳叶栎在以上不同污染地块均能正常生长,株高、地径和生物量与复合污染指数（内梅罗指数）及重金属含量呈负相关。其中纳塔栎对Cd的生物富集系数（BCF）在A、B、E区分别为6.27—8.37、3.67—4.38、42.93—52.75,高于C、D区柳叶栎对Cd的生物富集系数1.79—2.15、0.89—1.07。B—E区Zn的转运系数（TF）在1.79—2.28之间,A区Zn的转运系数为0.43。结论:纳塔栎和柳叶栎表现出较强的重金属耐性,对Cd具有较高的生物富集能力,对Zn具有较高的转运能力。其中纳塔栎对重金属积累能力较强,可作为亚热带地区铅锌矿区Cd、Pb、Zn、As复合污染土壤的植被恢复及生态修复候选树种。     

Rodent acorn selection in a Mediterranean oak landscape

Quercus suber, Quercus ilex and Quercus coccifera (Cork, Holm and Kermes oaks, respectively) are common evergreen oak species that coexist in the landscapes of the western part of the Mediterranean basin. Rodents are the main acorn predators and thus one of the main factors for understanding recruitment patterns in oaks. In this paper we analyse to what extent mice prefer acorns from one oak species over another in three oak species studied using acorn removal experiments and video tape recordings. Twenty labelled acorns from each of the three Quercus species (60 acorns) were placed in 40 cm×40 cm quadrats on each plot. Because selection might vary as a result of the vegetation context, we performed the trials in the five main vegetation types within the study area (four replicates in each vegetation type) in order to control for habitat influences on rodent acorn preferences (a total of 20 plots). The removal of 1,200 acorns occurred within 68 days. Mice removed 98.7% of the acorns. Q. ilex acorns were preferred over Q. suber and Q. coccifera in all vegetation types except in pine forest, where no acorn preferences were detected. Acorn removal rates differed with vegetation type, correlating positively with shrub cover. The distance at which acorns were displaced by rodents (mean =4.6 m±5.1 SD) did not differ between acorn species, but varied among vegetation types. Bigger acorns of Q. coccifera were selected only after Q. ilex and Q. suber acorns were depleted, while no size selection was detected for the latter two species. Thus, we conclude that rodents show preference for some oak acorns and that landscape context contributes significantly to rodent activities and decisions.     

Comparative structural study ofQuercus serrata andQ. acutissima formed byPisolithus tinctorius andHebeloma cylindrosporum

Ectomycorrhizas were synthesized in pots and growth pouches betweenQuercus serrata, Q. acutissima, and two ectomycorrhizal fungi,Pisolithus tinctorius andHebeloma cylindrosporum. Root morphology and the structure of the mantle and Hartig net were compared using light, fluorescence, scanning and transmission electron microscopy.P. tinctorius initially colonized root cap cells, and eventually produced a highly branched lateral root system with a complete mantle, whereasH. cylindrosporum promoted root elongation with few hyphae on the root apex surface indicating that interaction between roots differs with fungal species. Hartig net structure and hyphal inclusions varied between all the combinations tested. There were structural differences between mycorrhizas ofH. cylindrosporum/Q. acutissima grown in soil and growth pouches, which indicate that the growth pouch environment can induce artefacts in roots. Fruit bodies ofH. cylindrosporum developed in pots withQ. acutissima. AlthoughP. tinctorius has been used to inoculate oak seedlings in the nursery, results of this study indicate thatH. cylindrosporum may also be an effective ectomycorrhizal fungus forQ. serrata andQ. acutissima.     

No heat-stimulated germination found in herbaceous species from burned subtropical grassland

The inverse relationship between numbers of stomata (stomatal frequency) on tree leaves and ambient CO₂ concentration is increasingly applied for reconstructing past atmospheric CO₂ levels. The abundance of leaf remains of Quercus robur in Holocene peat and lake deposits in Europe makes this species potentially suitable for high-resolution stomatal frequency analysis. In order to quantify the CO₂ responsiveness of the species, the behavior of the stomatal index for Q. robur during the current anthropogenic CO₂ increase is determined on the basis of buried, herbarium and modern leaf material from the Netherlands. The stomatal index (SI), expressing the ratio of the number of stomata in a given area divided by the total number of stomata and other epidermal cells in that same area, is used in order to minimize influences on stomatal frequency of environmental conditions other than CO₂. The sigmoid SI response pattern recorded for Q. robur resembles that of the closely related species Q. petraea, although there is a difference in the timing of the response limitation of the two species to increasing atmospheric CO₂. For calibration purposes only the linear phase of the sigmoidal response curve is taken into consideration in the presented CO₂ response model, which allows confident combination of Q. robur and Q. petraea over the interval from 290 to 325 ppmv CO₂. The model is conservative in reconstructing past CO₂ mixing ratios outside the range of monitored response. As a result of the observed SI response limit, the model predicts CO₂ levels below 325 ppmv with a mean error of 10.2 ppmv, whereas higher CO₂ levels are underestimated.     

Regeneration by sprouting of holm-oak (Quercus ilex) stands exploited by selection thinning

Holm-oak forests of the Montseny Massif have traditionally been exploited by selection thinning, which consists of a partial removal of standing trees and biomass. Cutting percentages within each size class indicate that this management technique represents a compromise between forest improvement and harvesting.Sprouting is the mean regeneration process of these holm-oak stands. The mean number of living sprouts per stool is considerably higher than in other coppiced hardwood species, and decreases along the cutting-cycle according to a negative exponential function. Mean number of surviving sprouts per stool is significantly correlated with regenerative age and stool size. Death of sprouts throughout the cutting cycle is compensated by growth of surviving sprouts. Thus, the weight of living sprouts per stool increases linearly with time, and also with the basal area coppiced.Sprouts developing within the first fifteen years of regeneration belong to a single cohort that appears during the first year after cutting. After that period of time, new sprouts appear from dormant buds, so that after 30 years of regeneration a wide range of cohorts of different ages are present.     

Patterns and ecological consequences of abiotic heterogeneity in managed cork oak forests of Southern Spain

Spatial heterogeneity of abiotic factors influences the structure and function of forests and must be taken into account for their conservation and sustainable management. In this study, we evaluate the heterogeneity of abiotic environmental variables in managed cork oak (Quercus suber L.) forests in southern Spain at patch, site and regional scales. The extent of spatial heterogeneity depended on the environmental variable examined and the scale considered. For example, soil Mn and P and light availability in the understorey were very heterogeneous at the regional scale, while soil N had low regional heterogeneity, but high spatial variability, at patch scale, attributed to open overstorey and grazing disturbance. There was a general trend of increasing heterogeneity with spatial scale. We also study the effects of a silvicultural practice—shrub clearing on the forest environment and its consequence for spatial heterogeneity. Shrub clearing increased understorey light and decreased its spatial heterogeneity with idiosyncratic effects on soil properties and their spatial heterogeneity at each site. Finally, we compare the heterogeneity (estimated by the coefficient of variation) obtained in these cork oak forests with a database compiled from published studies on other forest environments. The comparison revealed a remarkable extent of abiotic heterogeneity in the cork oak forests studied, suggesting that a sustainable management of these forests should combine intrinsic and human induced abiotic heterogeneity to preserve crucial ecological processes and to maintain high levels of biodiversity.     

The combined role of topography and overstorey tree composition in promoting edaphic and floristic variation in a Mediterranean forest

This study aimed to address which factors, other than topography, contribute to the floristic variation of forested slopes. The natural forest studied is located in the Sudoeste Alentejano e Costa Vicentina Nature Park (southwestern Portugal). We sampled topographic, edaphic, floristic and community structure variables along three bottom–top hillside transects. Multivariate analyses of soil variables (by PCA), and of woody species composition and floristic–environmental relationships (by CCA and pCCA) were performed. Environmental–floristic trends strongly associated with the elevation gradient were identified. At lower altitudes, the lowest species richness, the highest soil fertility, and the tallest and most dense (least available light) canopy occurred. The spatial variation in woody species composition and abundance was closely associated with Zn availability in the soil and litter groundcover, but these varibles had significant spatial structure in the studied forest. The non-spatially structured species variance was better predicted by soil NO₃^– and NH₄⁺. The spatial variation of species data not shared with environmental variables was also calculated. We suggest that the influence of topographic gradient on the variation of edaphic variables and on the distribution and abundance of woody species was mediated by overstorey tree composition. Locally dominant tree species, in particular Quercus faginea and Quercus suber, may function as ecosystem engineers promoting environmental changes (i.e., Zn availability in soil, litter accumulation and light availability) that influence overall floristic variation.     

Seed dispersal by pygmy chimpanzees (Pan paniscus): A preliminary report

The role in seed dispersal played by the pygmy chimpanzees (Pan paniscus) inhabiting Wamba, Republic of Zaïre, was studied. Germination was tested for seeds of 17 plant species recovered from the feces of pygmy chimpanzees at Wamba. The fecal seeds of 13 species germinated, and in six of the species the germination rate for the fecal seeds was higher than that of control seeds. Although five other species showed a higher germination rate in the control seeds than in the fecal seeds, the remaining two species revealed no difference in germination rate between the fecal and control seeds. There was no great difference in germination velocity between the fecal and control seeds of the same species. For comparison, seeds of four plant species collected from the feces of common chimpanzees (Pan troglodytes) and gibbons (Hylobates lar) in captivity in Okinawa were tested for their germinability. In this test, although the seeds had passed through the digestive tract, their germinability demonstrated little change. Based on the behavioral characteristics of the pygmy chimpanzee at Wamba and observations of the captive primates on Okinawa, it seems that pygmy chimpanzees may play an important role in the seed dispersal of fruit plant species at Wamba.