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.     

Incorporating patterns of disperser behaviour into models of seed dispersal and its effects on estimated dispersal curves

The processes determining where seeds fall relative to their parent plant influence the spatial structure and dynamics of plant populations and communities. For animal dispersed species the factors influencing seed shadows are poorly understood. In this paper we test the hypothesis that the daily temporal distribution of disperser behaviours, for example, foraging and movement, influences dispersal outcomes, in particular the shape and scale of dispersal curves. To do this, we describe frugivory and the dispersal curves produced by the southern cassowary, Casuarius casuarius, the only large-bodied disperser in Australia’s rainforests. We found C. casuarius consumed fruits of 238 species and of all fleshy-fruit types. In feeding trials, seeds of 11 species were retained on average for 309 min (±256 SD). Sampling radio-telemetry data randomly, that is, assuming foraging occurs at random times during the day, gives an estimated average dispersal distance of 239 m (±207 SD) for seeds consumed by C. casuarius. Approximately 4% of seeds were dispersed further than 1,000 m. However, observation of wild birds indicated that foraging and movement occur more frequently early and late in the day. Seeds consumed early in the day were estimated to receive dispersal distances 1.4 times the ‘random’ average estimate, while afternoon consumed seeds received estimated mean dispersal distances of 0.46 times the ‘random’ estimate. Sampling movement data according to the daily distribution of C. casuarius foraging gives an estimated mean dispersal distance of 337 m (±194 SD). Most animals’ behaviour has a non-random temporal distribution. Consequently such effects should be common and need to be incorporated into seed shadow estimation. Our results point to dispersal curves being an emergent property of the plant–disperser interaction rather than being a property of a plant or species.     

种子产量对鼠类扩散栓皮栎坚果的影响

为了解林木种子产量对鼠类种子扩散行为的影响,于2008 年和2009 年,在国有济源市愚公林场调查了栓皮栎的种子雨;在每年的种子雨结束后,选择次生林生境,研究了鼠类对人工释放栓皮栎坚果的扩散,旨在探讨种子产量与种子扩散间的关系。结果表明:1)取食栓皮栎坚果的鼠类主要有大林姬鼠、社鼠和岩松鼠,2008 年的鼠类捕获率(2% )低于2009 年(10%),其差异并未达到显著性水平;2)栓皮栎的种子雨构成和产量存在年际差异,2008 年以完好种子为主且产量高于2009 年,而2009 年以败育种子为主;3)2008 年人工释放栓皮栎坚果的中位存留时间显著高于2009 年;4)2008 年鼠类对栓皮栎坚果的平均扩散距离显著低于2009 年,且两年的搬运距离几乎都集中在9 m 以内;5)2008 年鼠类对栓皮栎坚果的埋藏比例显著高于2009 年。结果提示,在种子高产年份,可能有较多的种子逃脱动物的取食,从而增加种子萌发和幼苗建成的机会,最终促进植物的更新。     

Synthetic seed production from encapsulated somatic embryos of cork oak (Quercus suber L.) and automated growth monitoring

Cork oak (Quercus suber) somatic embryos were coated with alginate for the production of synthetic seeds and their storability for commercialization was investigated. Also, the automatic monitoring of somatic embryo growth with a digital system of image capture was tested. A power regression model was fitted between size and fresh weight (Adjusted R-squared = 0.96). This method permitted growth assessment without contamination risk and opens the possibility of an automated control of culture growth for the future up scaling of plant production. Conversion rate of synthetic seeds was higher on medium supplemented with mineral nutrients than on medium without nutrients. Also, when the somatic embryos were coated without mineral nutrients added to the capsule, conversion rate was significantly lower. The addition of sucrose to the capsule had no significant effect on the conversion rate. No differences were recorded between 50 and 100 mM CaCl₂ for capsule complexation. Synthetic seeds were cold stored at 4°C for two months without significant loss of conversion capacity. The present study reports the first attempts to determine optimal storage time and conditions for conversion of encapsulated somatic embryos of cork oak.     

Modelling the drought impact on monoterpene fluxes from an evergreen Mediterranean forest canopy

In many ecosystems drought cycles are common during the growing season but their impact on volatile monoterpene emissions is unclear. Therefore, we aimed to develop and evaluate a process-based modelling approach to explore the explanatory power of likely mechanisms. The biochemically based isoprene and monoterpene emission model SIM-BIM2 has been modified and linked to a canopy model and a soil water balance model. Simulations are carried out for Quercus ilex forest sites and results are compared to measured soil water, photosynthesis, terpene-synthase activity, and monoterpene emission rates. Finally, the coupled model system is used to estimate the annual drought impact on photosynthesis and emission. The combined and adjusted vegetation model was able to simulate photosynthesis and monoterpene emission under dry and irrigated conditions with an R ² of 0.74 and 0.52, respectively. We estimated an annual reduction of monoterpene emission of 67% for the extended and severe drought period in 2006 in the investigated Mediterranean ecosystem. It is concluded that process-based ecosystem models can provide a useful tool to investigate the involved mechanisms and to quantify the importance of specific environmental constraints.     

Effect of water stress on monoterpene emissions from young potted holm oak (Quercus ilex L.) trees

We investigated the effects of a short period of water stress on monoterpene emissions from Quercus ilex, a common oak species of the Mediterranean vegetation and a strong emitter of monoterpenes. The experiment was carried out on two young saplings with a branch enclosure system under semi-controlled conditions. Under unstressed conditions, small qualitative (cis--ocimene, trans--ocimene, -caryophyllene and 1,8-cineol) and large quantitative (as much as 40% for the main compounds emitted) differences were observed between the two apparently similar trees. Nevertheless these differences did not affect the short-and long-term responses to temperature and water stress. Daily courses of emissions and gas exchanges were similar before and after the stress. During the most severe stress, emissions were reduced by a factor of two orders of magnitude and the log-linear relationship between emissions and temperature no longer existed. Photosynthesis and transpiration rates decreased as soon as the soil started to dry, whereas monoterpene emissions slightly increased for few days and then dropped when the daily CO₂ balance approached zero. We concluded that under water stress monoterpene emissions were highly limited by monoterpene synthesis resulting from a lack of carbon substrate and/or ATP. After rewatering, both emissions and gas exchanges recovered immediately, but to a level lower than the pre-stress level. These results have many implications for monoterpene emission modelling in the Mediterranean area, since the dry period generally extends from May to August. If our results are confirmed by field experiments, water stress could lead to a large overestimation of the emissions under summer conditions, when the algorithms based on light and temperature would give high emission rates.     

Spatial patterns of dispersal, seed predation and germination during colonization of abandoned grassland by Quercus petraea and Corylus avellana

Key processes during recruitment of Quercus petraea and Corylus avellana were investigated in abandoned calcareous grasslands and adjacent scrub using the following methods: (1) observation of hoarding animals during the main period of ripening of acorns and hazel nuts, (2) exposition of seeds on the soil surface and in 5–6 cm depth to test differences in predation and germination, and (3) mapping of seedlings in the grasslands. European jays (Garrulus glandarius) and mice were the main disperser of seeds. Jays preferred acorns, whereas the rodents were less selective, but probably more important for dispersal of nuts. The maximum dispersal distance was about 10–20 m for mice and was estimated several hundred metres for jays. Mice collected hoards of several seeds in about 2 cm depth in the soil, whereas jays stored single seeds. Seed predation and probably hoarding by mice were highest under scrub and in unmown grassland, while jays preferred mown sites for hoarding. However, hiding of seeds in the soil reduced losses in all sites. Predation of nuts was slightly less intensive than that of acorns. Seeds of Corylus were more sensitive to desiccation than Quercus, but in both species germination was lower for seeds exposed on the soil surface and in drier sites. Quercus and Corylus were the most abundant woody species in the fallow grasslands, probably due to the effective multi-staged dispersal by jays and mice, whereas wind-dispersed and fleshy-fruited species were less common; the latter restricted to margins of adjacent scrubland. The study provides several examples for discordance in suitability of patches for seeds and seedlings due to different habitat requirements of successive developmental stages. This result emphasizes the need for studies in the multiple stages during recruitment of vertebrate-dispersed plants.     

生境类型对啮齿动物扩散和贮藏栓皮栎坚果的影响

在河南省济源市王屋山地区,研究了次生林、灌丛、退耕地三个生境中啮齿动物对栓皮栎坚果的取食、
扩散与贮藏情况。结果表明:1)在研究地区,取食栓皮栎坚果的啮齿动物主要有大林姬鼠、社鼠和黑线姬鼠;
2)在次生林中,啮齿动物对坚果的搬运速率慢于在灌丛和退耕地生境中的搬运速率;3)次生林生境中,贮藏
坚果占被扩散坚果的比例高于灌丛和退耕地,而灌丛生境中被取食坚果占被扩散种子的比例高于退耕地和次生
林;4)栓皮栎坚果的中位存留时间在三个生境间有显著性差异。研究结果提示,在人工造林时,可通过对栓皮
栎坚果进行埋藏处理,以提高种子萌发率。     

Cytological analysis of early microspore divisions leading to gametic embryo formation in <Emphasis Type="Italic">Quercus suber</Emphasis> L. anther cultures

The correlation between the phenologic stage of the inflorescence and the microspore development stage was studied. Cytological examinations of the development of microspores during in vitro anther culture of cork oak (Quercus suber L.), were carried out during the first four weeks of culture. To observe the division occurring in the microspores, anthers were taken randomly from the cultures after heat shock treatment and were stained with DAPI. Most of the anthers responding to a heat stress treatment contained 91 % vacuolated microspores, indicating that this developmental stage is responsive to embryogenesis induction in cork-oak microspores. After the heat shock treatment some cork-oak microspores were induced and initiated the embryogenic pathway with the occurrence of numerous symmetric mitosis, producing structures with two to ten or more nuclei. These lead to the formation of high numbers of multicellular cork-oak microspores (pro-embryos). Twenty-forty days after induction, small white globular and cotyledonal embryos were observed, which further developed root and shoot, regenerating plantlets.     

Micropropagation of juvenile and adult Quercus suber L.

This paper describes research on the application of tissue culture techniques to the micropropagation of cork oak (Quercus suber L.), a forest species of ecological and industrial importance in the Mediterranean area. Apical buds and nodal stem segments were employed as initial explants. Their origins were young seedlings, stump sprouts and sprouts formed on cuttings collected from old trees.The action of the mineral medium and growth regulators was studied in the multiplication stage. Media with low concentrations of ions, such as Sommer's or Heller's, are more suitable for growth and proliferation of explants than other media richer in salts. It was also observed that cytokinin (BA) must be present for the culture development. Adding low concentrations of auxin (NAA) to the medium improves the multiplication rate, especially in vegetative material of adult origin.The auxin type is the most important factor in the promotion of rhizogenesis. The method of application determines the quality of the root system. Treatment with low concentrations of IBA added to the rooting medium gives the best results.High sucrose concentration also improves rooting. Diluting the mineral rooting medium is slightly favourable, although there is no significant difference between it and the standard mineral concentration.Abbreviations D Durzan's - GD Gresshoff & Doy's - H Heller's - L Lepoivre's - MS Murashige & Skoog's - SH Schenk & Hildebrandt's - S Sommer's medium     

Target regions for silvoarable agroforestry in Europe

Silvoarable agroforestry (SAF) has recently been proposed as an alternative land-use system for Europe. Data on soil, climate, topography, and land cover were integrated in a geographic information system (GIS) to identify agroforestry target regions where (i) productive growth of trees (Juglans spp., Prunus avium, Populus spp., Pinus pinea, and Quercus ilex) in SAF systems could be expected and where (ii) SAF systems could potentially reduce the risk of soil erosion, nitrate leaching and increase landscape diversity.The analysis shows that the investigated tree species could grow productively in SAF systems on 56% of the arable land throughout Europe (potential productive tree growth area). 80% of the European arable land was classified as potential risk area for soil erosion, nitrate leaching, and/or landscape diversity. Overlaying potential productive tree growth areas with the arable land that was considered as environmental risk area yielded target regions. They were found to make up about 40% of the European arable land and thus SAF could contribute to soil protection on 4%, to mitigate nitrate leaching on 18% and to increase landscape diversity on 32% of European arable land.Although limited by constrained data availability, the study shows that SAF could be implemented in a productive way throughout Europe and that it could help resolve some of the major land-use problems. The environmental benefits could justify the support of SAF by subsidies.     

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.     

Bark stripping in cork oak (Quercus suber L.): effect of an antitranspirant application on gas exchange and water relations of the stripped surface

Quercus suber L. is the primary source of industrial cork, which can be legally collected every 9 years. The main objective of this work was to test the efficiency of an application of an antitranspirant, at three different concentrations, after the bark stripping. For this purpose, several measurements of the gas exchange, water potential, total chlorophylls and the carotenoids contents were determined in cork oak trees, at two times in a day, morning and afternoon. The antitranspirant film was applied immediately after stripping. Transpiration rate showed a significant increase in the afternoon. The parameters, water potential, photosynthetic rates, stomatal conductance and the intrinsic water use efficiency, showed a significant decrease from morning to afternoon. The difference between pigments concentration was not significant throughout the day. Water potential and transpiration rate were high in the treatments with lower antitranspirant concentration. However, the treatment with a higher paraffin concentration showed larger photosynthesis rate. This result suggests that the loss of water observed for the stripping surface can be minimized by a larger concentration of the antitranspirant.     

Predating strategy of rodents on acorns of Quercusaliena var. acuteserrata under different predating risks and fate of acorns

The effects of rodents on forest regeneration have been highlighted in many ecological studies. In 2002 and 2003, the acorns of Quercus aliena var. acuteserrata were subjected to 12 different treatments. The daily dynamics and the amount of acorns that were finally left intact, predated in situ, or removed off were examined and documented. The ratios of acorns that were infested by bugs before and after predation by rodents were carefully documented. It was found that: (1) the ratios of acorns infested by bugs before and after predation by rodents showed significant difference (P > 0.05), suggesting that rodents would not prey on acorns during the predating process if acorns had been already infested by bugs. (2) When compared with the controls, the fate of acorns could be roughly classified into four types: acorns that were simply buried or placed on black paper showed no significant differences with the controls in their response to rodents, suggesting that rodents have no sensitivity to the little change of odor resulting from burying and may be more adapted to black background. Acorns attached with strings, dyed with black ink, cut into halves, or placed on white/green/red paper were predated in situ with much lower predation rates and lasted the same duration. Compared with the controls, the acorns were removed away by rodents with the same first day rate and the removal lasted longer. The final removal rate of the acorns by rodents was increased, whereas that of predation in situ was decreased, and none were left in the spot. This may suggest that rodents in this case increased their predation risk expectation and adopted a strategy of “less predation proportion in situ”. Acorns that were burnt were not predated in situ as well as removed in the first day, but the rates were raised to the maximum in the forth and fifth day and then declined to zero. Consequently, the final ratios of predation in situ and removal considerably decreased with many acorns, nearly 50%, being left intact. It can be hypothesized that the rodents responded to the change in odor of acorns by using the strategies of “less predation proportion in situ” and “less predating activities”, thereby resulting in larger number of acorns remained intact. Acorns that were enwrapped were barely predated in situ, removed in the first day with a normal predation rate, which declined to nearly zero in the second day and then increased till it declined again from the fourth day to the final day. The final rate of predation in situ was lower than 5% without acorns remained intact, and the rate of removal of acorns showed a considerable relative increase. This suggests that when there was a change in both odor and status of acorns, rodents also adopted the “spying” and “less predation proportion in situ” strategy, Because the predation risk is critical, the sense of sight would play an important role in rodent's living. (3) Definite amounts of acorns were predated in situ, discarded, removed away, predated ex situ, lost and hoarded separately.     

Quercus ilex browse utilization by Caprini in Sierra de Cazorla and Segura (Spain)

The impact of domestic and wild Caprini browsing on Quercus ilex has been examined in an area of the Sierra de Cazorla. Vegetation as a herbivore food supply, herbivore feeding regime and density in the study area during six sampling periods throughout two years, has been quantified. Wild Caprini show diets similar to the available vegetation, whereas domestic Caprini tend more towards the trophic specialities (browsing or grazing) of their genus. Nevertheless, this tendency was more pronounced in domestic goats than in sheep. A hypothetical estimate of Q. ilex intake by each species under the study conditions was carried out. It was found that domestic Caprini have a greater impact on the holm oak than wild Caprini, density and feeding-niche deviations being the main factors responsible for this situation.Abbreviations DM= dry matter     

Restricted dispersal in a flying beetle assessed by telemetry

Many insects living in ancient trees are assumed to be threatened as a result of habitat loss and fragmentation. It is generally expected that species in habitats with low temporal variability in carrying capacity have lower degree of dispersal in comparison to those in more ephemeral habitats. As hollow trees are long-lived, species in that habitat are expected to be sensitive to habitat fragmentation, due to a low capacity to establish new populations far from present ones. Using radio telemetry, we studied the dispersal for a beetle, Osmoderma eremita, living in hollow trees. O. eremita exhibited philopatry and only dispersed over short ranges. About 82–88% of the adults remained in the tree where they were caught. All observed dispersal movements ended up in nearby hollow trees and 62% in the neighbouring hollow tree. These results corroborate the suggestion that habitat fragmentation may be detrimental to insects living in temporally stable but spatially variable habitats. In order to preserve such species, we propose that conservation efforts should be focused on maintaining or increasing the number of suitable trees in and near presently occupied stands.     

Aggregated seed dispersal by wreathed hornbills at a roost site in a moist evergreen forest of Thailand

Hornbills (Bucerotidae) are widely regarded as important seed dispersers in tropical forests in Africa and Asia. We investigated how the roosting behavior of wreathed hornbills (Aceros undulatus) influences seed deposition and seedling survival at a roost site in a moist evergreen forest of Khao Yai National Park, Thailand. Fallen fruits and seeds were collected in traps that were placed around a roosting site for 14 months, and seedlings were monitored in adjacent quadrats for 3 years. Seedfall and seedlings of species represented in the hornbill diet occurred at significantly higher densities in the traps and quadrats located beneath the crown of the roosting tree than in those located beyond the crown. With the exception of Cinnamomum subavenium, the seeds and seedlings of most diet species rarely survived beyond the first year. The quality of hornbill dispersal to this roosting site may be poor due to the highly concentrated seedfall, which results in high seed and seedling mortality. However, the number of seeds deposited by each hornbill each day at roosting sites is relatively low. Wreathed hornbills are primarily scatter dispersers during the day and probably serve as agents of seed dispersal in the moist evergreen forest of Khao Yai.     

Relevance of genetics for conservation policies: the case of Minorcan cork oaks

Background and Aims

Marginal populations of widely distributed species can be of high conservation interest when they hold a significant or unique portion of the genetic diversity of the species. However, such genetic information is frequently lacking. Here the relevance of genetic surveys to develop efficient conservation strategies for such populations is illustrated using cork oak (Quercus suber) from Minorca (Balearic Islands, Spain) as a case study. Cork oak is highly endangered on the island, where no more than 67 individuals live in small, isolated stands in siliceous sites. As a consequence, it was recently granted protected status.

Methods

Two Bayesian clustering approaches were used to analyse the genetic structure of the Minorcan population, on the basis of nuclear microsatellite data. The different groups within the island were also compared with additional island and continental populations surrounding Minorca.

Key Results

Very high genetic diversity was found, with values comparable with those observed in continental parts of the species'' range. Furthermore, the Minorcan oak stands were highly differentiated from one another and were genetically related to different continental populations of France and Spain.

Conclusions

The high levels of genetic diversity and inter-stands differentiation make Minorcan cork oak eligible for specific conservation efforts. The relationship of Minorcan stands to different continental populations in France and Spain probably reflects multiple colonization events. However, discrepancy between chloroplast DNA- and nuclear DNA-based groups does not support a simple scenario of recent introduction. Gene exchanges between neighbouring cork oak stands and with holm oak have created specific and exceptional genetic combinations. They also constitute a wide range of potential genetic resources for research on adaptation to new environmental conditions. Conservation guidelines that take into account these findings are provided.     

Localized migration and dispersal by the sweet potato whitefly,Bemisia tabaci

Laboratory populations of the sweet potato whitefly, Bemisia tabaci, have been shown to consist of both migratory and trivial flying morphs. The behavior of these forms as part of the process of short-range migration was examined under field conditions. Insects were marked in a field of melons using fluorescent dust during two consecutive growing seasons. During the first growing season, passive traps used to collect living whiteflies, were placed along 16 equally spaced transects radiating from the field to a distance of up to 1.0 km. Wind out of the north-east consistently carried migrating whiteflies to traps placed along transects in the south-western quadrant because cold air drainages dictate wind direction during early morning hours in the desert South-west. For this reason, during the second season traps were laid out over fallow ground in a rectangular grid extending 2.7 km to the south-west of the marked field. If dispersal was entirely passive, patterns could be described using a diffusion model. Statistical examination of the data, howèver, demonstrated that the distribution on all days was patchy. Geostatistical techniques were used to describe the observed patchiness. Traps in the immediate vicinity of the marked field caught more whiteflies than the daily median. Large numbers were also collected from near the periphery of the grid. White-flies were far less prevalent in the grid's center. As a result, the distribution of captured whiteflies can be described as bimodal. These patterns confirm behavior observed in the laboratory, i.e., a portion of the population are trivial fliers that do not engage in migration and are consequently captured in traps near the field, and a portion initially respond to cues associated with skylight, ignoring cues provided by the ground, and fly for a period of time before landing in distant traps. During both years movement out of the field had an exaggerated directional component on 13 of 14 days.     

Biological control of an alien tree, Acacia cyclops, in South Africa: impact and dispersal of a seed-feeding weevil, Melanterius servulus

Several Australian Acacia species have become highly invasive in a number of ecologically sensitive areas of South Africa. Many have useful attributes that, to an extent, have hampered biocontrol efforts by restricting potential agents to those reducing seed production whilst not affecting vegetative growth. The outcome of the introduction of a seed-feeding weevil, Melanterius servulus, against Acacia cyclops in 1994, was assessed by investigating the plant’s reproductive phenology, as well as levels of damage caused by the beetles. The study provided essential information on host/agent interactions and, on the build up and spread of populations of the agent. Seed damage is commonly as high as 90% (exceptionally it reaches 100%), and dispersal rates of the beetles on average is almost 2 km per year. The consistently high levels of seed damage achieved hold promise for the future of the program, however, its ultimate success will be further enhanced through the integration of biological control with other manual clearing operations that are currently employed on a large scale in South Africa.