Ilex Aquifolium Research Articles

The enigma of Calonectria species occurring on leaves of Ilex aquifolium in Europe

Species of Calonectria are common saprobes and plant pathogens on a wide range of hosts occurring in subtropical to tropical regions of the world. The aim of the present study was to resolve the status of new Calonectria collections obtained on Ilex leaves from France. Based on DNA sequence data of their β-tubulin and histone gene regions, as well as morphology, the new collections matched the ex-type strain of Cylindrocladium ilicicola. On the host and in culture, yellow to brownish-yellow perithecia were observed that did not strain red in 3 % KOH. Based on these results, C. ilicicola and its purported teleomorph, Ca. pyrochroa, were shown to represent two distinct species, as the latter has bright red perithecia that strain purple in KOH. A new combination, Ca. lauri, based on Tetracytumlauri, is subsequently proposed for C. ilicicola. Calonectria lauri is distinct from Ca. ilicicola, a pathogen commonly associated with Cylindrocladium black rot of peanut. Finally, Ca. canadiana is proposed as new name for Cy. canadiense, which is a nursery pathogen involved with root rot of several tree genera in Quebec, Canada.

Resistance to wildfire and early regeneration in natural broadleaved forest and pine plantation

The response of an ecosystem to disturbance reflects its stability, which is determined by two components: resistance and resilience. We addressed both components in a study of early post-fire response of natural broadleaved forest ( Quercus robur, Ilex aquifolium) and pine plantation ( Pinus pinaster, Pinus sylvestris) to a wildfire that burned over 6000 ha in NW Portugal. Fire resistance was assessed from fire severity, tree mortality and sapling persistence. Understory fire resistance was similar between forests: fire severity at the surface level was moderate to low, and sapling persistence was low. At the canopy level, fire severity was generally low in broadleaved forest but heterogeneous in pine forest, and mean tree mortality was significantly higher in pine forest. Forest resilience was assessed by the comparison of the understory composition, species diversity and seedling abundance in unburned and burned plots in each forest type. Unburned broadleaved communities were dominated by perennial herbs (e.g., Arrhenatherum elatius) and woody species (e.g., Hedera hibernica, Erica arborea), all able to regenerate vegetatively. Unburned pine communities presented a higher abundance of shrubs, and most dominant species relied on post-fire seeding, with some species also being able to regenerate vegetatively (e.g., Ulex minor, Daboecia cantabrica). There were no differences in diversity measures in broadleaved forest, but burned communities in pine forest shared less species and were less rich and diverse than unburned communities. Seedling abundance was similar in burned and unburned plots in both forests. The slower reestablishment of understory pine communities is probably explained by the slower recovery rate of dominant species. These findings are ecologically relevant: the higher resistance and resilience of native broadleaved forest implies a higher stability in the maintenance of forest processes and the delivery of ecosystem services.

Effects of Forest Fragmentation on Seed Dispersal and Seedling Establishment in Ornithochorous Trees

Habitat fragmentation increases seed dispersal limitation across the landscape and may also affect subsequent demographic stages such as seedling establishment. Thus, the development of adequate plans for forest restoration requires an understanding of mechanisms by which fragmentation hampers seed delivery to deforested areas and knowledge of how fragmentation affects the relationship between seed-deposition patterns and seedling establishment. We evaluated the dispersal and recruitment of two bird-dispersed, fleshy-fruited tree species (Crataegus monogyna and Ilex aquifolium) in fragmented secondary forests of northern Spain. Forest fragmentation reduced the probability of seed deposition for both trees because of decreased availability of woody perches and fruit-rich neighborhoods for seed dispersers, rather than because of reductions in tree cover by itself. The effects of fragmentation went beyond effects on the dispersal stage in Crataegus because seedling establishment was proportional to the quantities of bird-dispersed seeds arriving at microsites. In contrast, postdispersal mortality in Ilex was so high that it obscured the seed-to-seedling transition. These results suggest that the effects of fragmentation are not necessarily consistent across stages of recruitment across species. Habitat management seeking to overcome barriers to forest recovery must include the preservation, and even the planting, of fleshy-fruited trees in the unforested matrix as a measure to encourage frugivorous birds to enter into open and degraded areas. An integrative management strategy should also explicitly consider seed-survival expectancies at microhabitats to preserve plant-population dynamics and community structure in fragmented landscapes.

Cattle exclosure and vegetation dynamics in an ancient, Irish wet oakwood

The effects of a 10-year period of cattle exclosure on the ground flora and tree regeneration of an Irish ancient lowland wet oakwood (Corylo-Fraxinetum deschampsietosum) community are assessed. The approach was to record changes in a quadrat sample located in a National Nature Reserve and to model change by multivariate statistical analysis using Principal Components Analysis and Redundancy Analysis ordination. Exclosure gave significant increases in the abundance of regenerating ash Fraxinus excelsior and holly Ilex aquifolium. Non-native tree species establishment also occurred. In the ground flora, there were significant decreases in the abundance of ruderal species and grasses and significant increases in graze-resistant and graze-sensitive species usually restricted to broadleaf woodland habitats. Changes were similar under low and high canopy sites, but greater under a high canopy. In Irish lowland wet oakwood, managed historically to favour oak and currently grazed by cattle, exclosure promotes the succession of ash to the canopy and holly to the understorey. With time, this is likely to lead to a shift from oak dominance. In a conservation context this, together with an increased risk of non-native tree species establishment, can be ecologically damaging, particularly in small sites without natural dynamics, as is the case in the hedged agricultural landscapes of western Europe. We conclude that site-based conservation objectives should be prioritised when implementing generic, landscape-scale management strategies, such as rotational exclosure.

Spatial associations among tree species in a temperate forest community in North-western Spain

Analyzing spatial patterns in plant communities may provide insights in the importance of different processes for community assembly and dynamics. We applied techniques of spatial point pattern analysis to data from a fully mapped plot of a temperate forest community ( Corylus avellana, Crataegus monogyna, Fagus sylvatica, Ilex aquifolium and Taxus baccata) in North-western Spain to conduct a community wide assessment of the type and frequency of intra and interspecific spatial association patterns. We first explored the overall intra and interspecific patterning, and then classified the types of association patterns at various neighbourhoods. By conditioning on the larger scale pattern we then explored small-scale (0–15 m) intraspecific and interspecific patterns. Association patterns varied from strong positive association at small scales to, as a by-product, repulsion at intermediate scales. Surprisingly, there were no negative associations at small scales, but trees were arranged in multi-species clumps, up to 2.5 m in diameter and comprising a few individuals. Ilex and Corylus, the understorey species, were frequently involved in the clumps, showing positive small-scale association with the other tree species. Our analyses highlighted that animal mediated seed dispersal, interspecific facilitation and perturbation processes may operate successively to shape tree distributional patterns, although their relative importance vary among species. Given the complexity of the patterns described and the current threats to some of the species studied, directed experiments in the field are needed to further elucidate some of the hypotheses derived.

Invasive Hollies (Ilex, Aquifoliaceae) and Their Dispersers in the Pacific Northwest

Abstract Naturalized Ilex aquifolium L. (English holly) was first collected in the Pacific Northwest in 1953, based on herbarium records. Field surveys showed it is now commonly naturalized from northwestern California to coastal British Columbia. Ilex crenata Thunb. and I. opaca Aiton were also found growing outside of cultivation, but rarely. A key and seed illustrations are provided to distinguish these three Ilex species. Between 2003 and 2006 twice-weekly visits to naturalized and cultivated hollies in Seattle revealed seven species of birds disseminating seeds by eating the fruits. American robins, Turdus migratorius, accounted for 96% of 2796 frugivory observations on I. aquifolium, followed by European starlings, Sturnus vulgaris (3.2%). Ilex aquifolium fruits ripened in October and persisted for six months, yet 99% of all fruit was consumed between November and February. A study of I. aquifolium seed fate found pre-dispersal diurnal seed predation was rarely observed. Bird-regurgitated seed was m...

Modelling the spatial distribution of tree species with fragmented populations from abundance data

Spatial distribution modelling can be a useful tool for elaborating conservation strategies for tree species characterized by fragmented and sparse populations. We tested five statistical models—Support Vector Regression (SVR), Multivariate Adaptive Regression Splines (MARS), Gaussian processes with radial basis kernel functions (GP), Regression Tree Analysis (RTA) and Random Forests (RF)—for their predictive performances. To perform the evaluation, we applied these techniques to three tree species for which conservation measures should be elaborated and implemented: one Mediterranean species (Quercus suber) and two temperate species (Ilex aquifolium and Taxus baccata). Model evaluation was measured by MSE, Goodman-Kruskal and sensitivity statistics and map outputs based on the minimal predicted area criterion. All the models performed well, confirming the validity of this approach when dealing with species characterized by narrow and specialized niches and when adequate data (more than 40–50 samples) and environmental and climatic variables, recognized as important determinants of plant distribution patterns, are available. Based on the evaluation processes, RF resulted the most accurate algorithm thanks to bootstrap-resampling, trees averaging, randomization of predictors and smoother response surface.

Diet and habitat selection in Cantabrian Capercaillie (Tetrao urogallus cantabricus): ecological differentiation of a rear-edge population

Cantabrian capercaillie Tetrao urogallus cantabricus is a peripheral population with distinctive phenotypic, biogeographic, and genetic characteristics. Hence, the population may also show substantial ecological differentiation associated with its habitat in purely deciduous forests. We assessed seasonal diet selection, small-scale habitat selection, and patterns of trophic niche width in Cantabrian capercaillie over two years. Diet was found to be a driver of small-scale habitat selection, a result consistent with previous studies of stand-scale habitat selection. Diet and habitat selection showed the importance of beech Fagus sylvatica, holly Ilex aquifolium, bilberry Vaccinium myrtillus, and ferns in Cantabrian capercaillie’s resource selection. Conversely, the abundant oaks Quercus petraea, birches Betula pubescens, and heaths Erica sp. were used below their availability. The reliance on bilberry appears as a unifying characteristic between central and peripheral capercaillie populations. Cantabrian capercaillie showed stronger reliance on understory resources than range-central populations. It also showed wider trophic niche and higher specialization of feeding events. Trophic niche patterns and reliance on ground resources indicated a marked ecological differentiation, which stresses the need for local data and specific conservation actions.

Current and Potential Distributions of Three Non-Native Invasive Plants in the Contiguous USA

ABSTRACT: Biological invasions pose a serious threat to biodiversity, but monitoring for invasive species is time consuming and costly. Understanding where species have the potential to invade enables land managers to focus monitoring efforts. In this paper, we compared two simple types of models to predict the potential distributions of three non-native invasive plants (Geranium robertianum, Hedera spp., and Ilex aquifolium) in the contiguous USA. We developed models based on the climatic requirements of the species as reported in the literature (literature-based) and simple climate envelope models based on the climate where the species already occur (observation-based). We then compared the results of these models with the current species distributions. Most models accurately predicted occurrences, but overall accuracy was often low because these species have not yet spread throughout their potential ranges. However, literature-based models for Geranium and observation-based models for Ilex illustrated ...

In VitroAnticancer Potential of Tree Extracts from the Walloon Region Forest

Forty-eight extracts from 16 common Belgian trees from the Walloon Region forest were evaluated for in vitro growth inhibitory activity against the human LoVo colon cancer, PC3 prostate cancer, and U373 glioblastoma cell lines. Our study was performed with the aim of selecting plant candidates in order to later isolate new anticancer compounds from an easily affordable tree material. Extracts from Alnus glutinosa (stem bark), Carpinus betulus (leaves and stem bark), Castanea sativa (stem bark), Fagus sylvatica (leaves), Ilex aquifolium (leaves), Larix decidua (leaves), Quercus petraea (stem bark), and Quercus robur (leaves) showed for the first time potent in vitro growth inhibitory activity and could become easily affordable sources of potential new anticancer agents. Root extracts from Robinia pseudoacacia, already known for containing cytotoxic lectins, also showed interesting activity.

Altitudinal differentiation in growth and phenology among populations of temperate-zone tree species growing in a common garden

The aim of the study was to determine whether there are genetic variations in growth and leaf phenology (flushing and senescence) among populations of six woody species ( Abies alba Mill., Acer pseudoplatanus L., Fagus sylvatica L., Fraxinus excelsior L., Ilex aquifolium L., and Quercus petraea (Matt.) Liebl.) along altitudinal gradients, using a common-garden experiment. We found (i) significant differences in phenology and growth among provenances for most species and (ii) evidence that these among-population differences in phenology were related to the annual temperature at the provenance sites for ash, beech, and oak. It is noteworthy that along the same climatic gradient, species can exhibit opposing genetic clines: beech populations from high elevations flushed earlier than those from low elevations, whereas we observed the opposite trend for ash and oak. For most species, significant altitudinal clines for growth were also revealed. Finally, we highlighted the fact that both phenology timing and growth rate were highly consistent from year to year. The results demonstrated that despite the proximity of the populations in their natural area, differences in altitude led to genetic differentiation in their phenology and growth. These adaptive capacities acting along a natural climatic gradient could allow populations to cope with current climate change.

Antibacterial, antifungal, and antimycobacterial activity of Ilex aquifolium leaves

The objective of this study was to examine antibacterial, antifungal, and antimycobacterial properties of Ilex aquifolium L. (Aquifoliaceae) growing in Turkey. The ethanol, ethyl acetate, chloroform, and n-hexane extracts prepared from the leaves of I. aquifolium were tested against Escherichia coli, Staphylococcus aureus, Pseudomonas aeruginosa, Enterobacter aerogenes, Proteus vulgaris, Salmonella typhimurium, and Candida albicans for antibacterial and antifungal evaluation using the microdilution broth susceptibility assay. In addition, antimycobacterial activity of the crude extracts of I. aquifolium was evaluated by microplate Alamar blue assay. The results showed that the extracts tested, except n-hexane, possessed moderate antibacterial and antifungal activity varying from 62.5 to 250 μg/mL. On the other hand, the ethanol extract of the leaves exhibited a minimum inhibitory concentration value (MIC) against Mycobacterium tuberculosis H37Ra strain of 200 μg/mL.

A new root‐knot nematode, Meloidogyne silvestris n. sp. (Nematoda: Meloidogynidae), parasitizing European holly in northern Spain

High infection rates of European holly (Ilex aquifolium) feeder roots by an unknown root‐knot nematode were found in a holly forest at Arévalo de la Sierra (Soria province) in northern Spain. Holly trees infected by the root‐knot nematode showed some decline and low growth. Infected feeder roots were distorted and showed numerous root galls of large (8–10 mm) to moderate (2–3 mm) size. Morphometry, esterase and malate dehydrogenase electrophoretic phenotypes and phylogenetic trees of sequences within the ribosomal DNA (rDNA) demonstrated that this nematode species differs clearly from other previously described root‐knot nematodes. Studies of host‐parasite relationships showed a typical susceptible reaction in naturally infected European holly plants, but did not reproduce on a number of cultivated plants, including tomato, grapevine, princess‐tree and olive. The species is described here, illustrated and named as Meloidogyne silvestris n. sp. The new root‐knot nematode can be morphologically distinguished from other Meloidogyne spp. by: (i) roundish perineal pattern, dorsal arch low, with fine, sinuous cuticle striae, lateral fields faintly visible; (ii) female excretory pore level with stylet knobs, or just anterior to them, EP/ST ratio about 0·8; (iii) second‐stage juveniles with hemizonid located 1 to 2 annuli anterior to excretory pore and short, sub‐digitate tail; and (iv) males with lateral fields composed of four incisures, with areolated outer bands. Phylogenetic trees derived from maximum parsimony analysis based on 18S, ITS1‐5·8S‐ITS2 and D2–D3 of 28S rDNA showed that M. silvestris n. sp. can be differentiated from all described root‐knot nematode species, and it is clearly separated from other species with resemblance in morphology, such as M. ardenensis, M. dunensis and M. lusitanica.

Ilex geissertii sp. n. (Aquifoliaceae), a fossil ancestor of Ilex sect. Ilex in the upper Miocene and Pliocene of Europe

Leaf remains of Ilex L. sect. Ilex known from the Pliocene of Auenheim (Alsace, France) and Frankfurt a. M. - Niederrad (Hesse, Germany) as Ilex aff. cornuta Lindley fossilis Geissert and Ilex aquifolium L. fossilis Engelhardt, respectively, are re-interpreted as representing a new species Ilex geissertii Kvaček, Teodoridis et Wang Qing, sp. nov. Similar leaf fossils occur also in the upper Miocene of the Netherlands (Brunssum), France (Murat), and the Pliocene of Italy (Valdarno Superiore). Although these fossil records were considered either direct ancestors of I. aquifolium L. (Near-East, Europe, N Africa) or allied to I. cornuta Lindley and Paxton (China and Korea), both the mentioned extant species differ in the leaf morphology. Close affinities of I. geissertii to several extant representatives of Ilex subsect. Ilex (= Oxyodontae Loesener nom. illegit.) are indisputable and stress a common origin of this group of the hollies known already from the Oligocene in Europe ( Ilex castellii Kvaček et Walther). Several extinct elements of the European late Miocene and Pliocene flora accompanying this new holly (e.g., Ginkgo, Taxodium, Fagus kraeuselii, Parrotia, Craigia, and Trichosanthes) share similar former phyllogeography in having their modern nearest relatives outside Europe.

A multiple approach for the evaluation of the spatial distribution and dynamics of a forest habitat: the case of Apennine beech forests with Taxus baccata and Ilex aquifolium

An approach integrating phytosociological and stand structure surveys with the predictive modelling of species distribution was applied to analyse the spatial distribution and dynamics of the Apennine beech forests with Taxus and Ilex, a high conservation priority forest habitat in Europe. The homogeneity of the habitat was tested trough a Mann-Whitney test between beech woods with Taxus and those with Ilex with respect to climatic, topographic, structural and environmental parameters: the former have proven to be more microthermic, mesophilous and characterised by a closer canopy. Five statistical models were compared to analyse the relationship between bioclimatic parameters and Taxus and Ilex spatial distribution: Regression Tree Analysis, the most efficient model, has shown that the distribution of Taxus is influenced by precipitation variables, while Ilex is mainly influenced by temperature variables. This model highlighted that Ilex has a potential area that surrounds, at lower altitudes, that of Taxus. A stepwise multiple regression analysis has been applied to identify the factors influencing the regeneration of the two species: beside climatic parameters, Taxus regeneration is negatively influenced by soil nitrate concentration (an indicator of livestock disturbance) while Ilex is negatively influenced by beech forest cover. Traditional management practices seem to have an effect on the regeneration of the two species: frequent cuts favour the regeneration of Ilex, reducing the forest cover and allowing more light penetration, while Taxus, less resistant to grazing livestock, is confined to more inaccessible places. The multiple approach has proven to be useful for the elaboration of two differentiated conservation strategies for the two beech forest types.

Modifiche compositive e strutturali in soprassuoli in evoluzione naturale della riserva M.a.B. di Montedimezzo (Isernia)

Tree species composition and structural dynamics in forest stands under natural evolution in the Montedimezzo M.a.B. reserve. This study is included in an experimental protocol established in 1954 aimed at examining the natural dynamics in mixed broadleaves high forests dominated by Turkey oak (Quercus cerris L.) and beech (Fagus sylvatica L.). Stand development patterns in the last fifteen years are reported. Repeated quantitative and qualitative inventories were carried out in two permanent plots (3 ha each) and in two structural transects (0.2 ha each) located in Collemeluccio-Montedimezzo M.a.B. reserve (Is). The study highlighted, as already observed in 1954, the differences between the two multilayered stands. In the higher elevation beech is the predominant species recording 53% of importance value index (I), followed by Turkey oak (I=25%). In the lower elevation the dominant species is Turkey oak (I=46%), followed by hornbeam (Carpinus betulus L.) (I=28%) and beech (I=12%). The decreased presence of Turkey oak and the increase of beech density are confirmed in each plot over the last fifteen years. Among the other tree species, importance values rise only for silver fir (Abies alba Mill.) (introduced) and holly tree (Ilex aquifolium L.), while hornbeam and hedge maple have been reducing their own presence.

Um método eficiente para a detecção de Phytophthora cinnamomi associada com a Doença da Tinta do Castanheiro na rizosfera de Castanheiro (Castanea sativa Mill.)

The life cycle of Phytophthora cinnamomi Rands and P. cambivora (Petri)Buisman, both species associated with Ink Disease of Chestnut, occurs entirely in soil. The presence of any of these parasitic species is a limiting constraint for new plantations of sweet chestnut and represents a serious problem for established groves. Sensitive and fast detection methods are necessary to guarantee the absence of inoculum in the propagation material and to evaluate the sanitary condition of soils. In this study a method that involved a baiting assay conjugated with selective agar medium was developed. As biological baits, leaf discs of sweet chestnut (Castanea sativa), holly trees (Ilex aquifolium) and camellia (Camellia japonica) were used, at 25 oC and normal laboratory conditions with or without addition of biocides (pimaricin and penicillin) in the water of soil dilution. The leaf discs of sweet chestnut were more efficient for detection of Phytophthora and greater percentage of positive isolations were always obtained in the selective medium P10VPH. Results of this study allow us to establish an efficient protocol for detection of P. cinnamomi associated with Ink Disease of Chestnut in the rhizosphere of sweet chestnut.

Hazel dormouse Muscardinus avellanarius (L.) nest site selection in hedgerows

The natural nest sites in hedgerows of the hazel dormouse Muscardinus avellanarius were investigated in Devon, in south-west England. A total of 73 nests were found in the autumn by careful searching of hedgerows, all of which were species-rich. In hedgerows that had not been cut for 6 years or more, most nests were in bramble Rubus fruticosus agg. and field rose Rosa arvensis margins, with a few in holly Ilex aquifolium bushes. In contrast, in hedgerows that had been cut within 6 years, the majority of nests were in shrubs, the dense protective growth at the cut line being strongly favoured. Blackthorn Prunus spinosa was preferentially selected, hazel Corylus avellana was avoided. It is suggested that the availability of safe nest sites may limit population density and that where nesting conditions are good the hazel dormouse may prefer to build nests in unenclosed situations rather than in hollows, whether natural hollows, such as tree holes, or artificial ones, such as nest boxes. To encourage hazel dormice in hedgerows, bramble and rose margins should be allowed to develop, and some lengths of hedgerow should be cut occasionally, particularly where blackthorn is present.

Leaf phenology sensitivity to temperature in European trees: Do within-species populations exhibit similar responses?

Consequences of climate warming on tree phenology are readily observable, but little is known about the differences in phenological sensitivity to temperature between species and between populations within a species. The aim of the present study is to compare phenological sensitivities to temperature of seven woody species between each other and within-species between two geographical areas using both altitudinal and temporal gradients ( Abies alba, Acer pseudoplatanus, Carpinus betulus, Fagus sylvatica, Fraxinus excelsior, Ilex aquifolium and Quercus petraea). The timing of leaf unfolding was monitored (i) over 2 years along two altitudinal gradients in the Pyrénées mountains (six species), and (ii) over 22 years in Fontainebleau forest (four species). Three species were present in both areas which allowed us to compare their phenological sensitivity to temperature over altitudinal and temporal gradients. Along altitudinal gradients, we observed for all species an advance in leaf unfolding with decreasing elevation, ranging from 11 to 34 days 1000 m −1 for beech and oak, respectively. Across the temporal gradient, we found significant advances in leaf unfolding for oak (−0.42 days year −1) and ash (−0.78 days year −1) since 1976, whereas no significant advance was observed for beech and hornbeam. For both gradients and for all species, significant correlations were found between leaf unfolding dates and temperature, except for beech in the temporal study. Moreover, we highlighted that phenological sensitivity to temperature was very similar between the two geographically separated populations (Pyrénées and Fontainebleau forests). Thus, oak had the strongest sensitivity (−7.48 and −7.26 days °C −1 in altitudinal and temporal gradient, respectively) and beech had the lowest (−2.09 and −2.03 days °C −1). Our results suggest that population sensitivity to global warming might be stable for a given species, in spite of its possible local adaptation.

The role of remnant trees in seed dispersal through the matrix: Being alone is not always so sad

Remnant trees within the non-forest matrix are common structures of forest landscapes which have high conservation value due to their supposed roles as biological legacies and stepping stones in fragmented forest scenarios. Fleshy-fruited remnant trees are dispersal foci for many forest plants, as seeds accumulate under their canopies after visitation by forest frugivores. Despite this recognized effect, little is known about the relative role of remnant trees in maintaining their seed dispersal function after forest fragmentation. In this work, conducted in the Cantabrian Range (Northern Spain), seed deposition by frugivorous birds and post-dispersal seed predation by rodents were compared beneath the canopies of hawthorn ( Crataegus monogyna) trees scattered in the non-forested matrix, to those embedded in forest patches. We studied two years which had strong differences in community-wide fruit abundance (high in 2004, and low in 2005). Hawthorn and holly ( Ilex aquifolium) seeds dominated the seed rain. The density of dispersed seeds differed between years and tree type, with higher values under patch trees relative to remnant ones. However, the effect of tree type depended on both the year and the seed species, as larger differences were found in the year of high fruit availability, and with holly seeds. Higher levels of post-dispersal seed predation on holly seeds also contributed to offset the differences between tree types. Our results suggest that remnant trees, by functioning as dispersal foci, facilitate the dispersal of the two most abundant plant species for forest succession through the matrix. More importantly, we demonstrated that in years of low fruit availability in which forest frugivores are forced to exploit scattered fruit resources, the role of remnant trees may even be equivalent to that played by forest trees.