Barriers to Forest Regeneration in an Abandoned Pasture in Puerto Rico

Sources of forest regeneration (soil seed bank, seed rain) and barriers to seedling establishment were examined in a recently abandoned pasture in eastern Puerto Rico. Few woody species were found in the soil seed bank or in the seed rain. The number of seeds and species in the seed rain and soil seed bank declined with distance from the adjacent secondary forest. Nine species naturally dispersed and colonized plots during the study, with the wind‐dispersed tree Tabebuia heterophylla being the predominant colonizer (91% of all seedlings). Barriers to seedling establishment were determined using a blocked field experiment with eleven woody species representative of a variety of life histories. Each species was planted under the pasture vegetation (control) or in areas where all vegetation was removed (removal). Germination was enhanced for four species in the control treatment, five species were not affected, and two species did not germinate under either treatment. Survival to 6 months was higher in the removal treatment for two species. Seedling biomass was greater in the removal treatment at 12 months for one species. Seed mass was a good predictor of germination success and final shoot biomass, but not survival. This study demonstrates that seeding recently abandoned pastures with a mix of known pioneer species may accelerate the rate of secondary succession, but some species will have to be planted in later successional stages in order to overcome strong barriers to establishment.     

西双版纳热带森林土壤种子库与地上植被的关系

通过实验研究探讨了西双版纳几类热带森林的土壤种子库与地上植被的关系．结果表明,在森林演替的初期,土壤种子库与地上植被共有的种类和种子储量较多,随着林龄的增大,外来种子的比例逐渐增加,到季节雨林阶段,土壤种子库中的种子大部分为来自群落外的先锋种类．这些种子在郁闭的林冠下很难萌发,一旦森林受到干扰出现林窗或空旷地,这些潜在的种源将迅速萌发,参与植被的恢复或演替     

Postdispersal Seed Fate of Tropical Montane Trees in an Agricultural Landscape, Southern Costa Rica

Variation in postdispersal seed fate is an important factor driving patterns of forest regeneration. Because most previous studies have not tracked final seed fate and have commonly equated seed removal with predation without considering the possibility of secondary dispersal, little is known about individual seed mortality factors in successional and degraded habitats. This study tracked the postdispersal fate of large-seeded tropical montane trees in abandoned pasture, young secondary forests, and small forest fragments during two consecutive years in an agricultural landscape in southern Costa Rica. The incidence of secondary dispersal by animals, scatterhoarding in particular, and the effects of seed burial on germination were measured. Overall, seeds survived through germination more often in secondary forests with high levels of mortality occurring in abandoned pastures and forest fragments. The majority of seed mortality was caused by rodent predation in forest fragments, insects and fungal pathogens in secondary forests, and a combination of desiccation, insects, and fungal pathogens in pastures. Seeds were frequently secondarily dispersed in larger forest fragments, whereas they were only rarely moved in pastures and secondary forests. Burial tended to improve germination in pastures and was important for an often scatterhoarded species, Otoba novogranatensis, in all habitats. The results of this study suggest that: (1) seed mortality factors differ in response to the type of habitat degradation; (2) large-seeded species have high potential for survival when dispersed to young secondary forests; and (3) seed removal is not a reliable proxy for seed predation, particularly in less degraded forest fragments.     

Effects of Forest Management on Diversity and Abundance of Fruit-feeding Nymphalid Butterflies in South-eastern Côte d'Ivoire

The effects of forest management on vegetation structure and capture frequencies of nymphalid butterflies were studied in a logged rain forest in south-east Côte d'Ivoire. An experimental compartment, where liberation thinning was carried out 3 years before, and a 5-year-old mono-dominant tree plantation were compared to a regenerating control compartment; 3642 specimens of 97 species were trapped. Accumulated species richness and diversity indices were lower in the control compared to the liberation thinning compartment, but lowest in the plantation. However, the habitat preference for traps situated in the control compartment was negatively correlated with the size of the species geographical range. Four out of five species with a lower capture frequency in the liberation thinning compartment showed preferences for mature succession stages and were either Upper Guinean endemics or Guinea–Congolian–restricted species. The seven species with higher capture frequencies in the liberation thinning compartment were all geographically widespread. Five of them showed higher frequencies in younger succession stages. Eight species, three canopy specialists and two Guinea savannah species, significantly preferred the plantation, while 17 species avoided this management type. Liberation thinning seems to affect the more specialised species with smaller geographic ranges, thus risking loss of regional diversity.     

普洱市周边地区4种土地利用类型土壤种子库特征

通过对云南普洱市周边地区次生季风常绿阔叶林、针阔混交林、人工更新形成的针叶林及茶园等4种土地利用类型的野外调查及土壤种子库的萌发实验,探讨其土壤种子库的密度大小、物种丰富度和组成及与地上植被的关系。结果表明:干扰强度与频度不同导致土地利用类型之间土壤种子库密度与物种丰富度存在较大差异,土壤种子库密度大小顺序为:针叶林(248.67±116.86)粒·m-2>针阔混交林(186.00±43.27)粒·m-2>次生季风常绿阔叶林(107.33±16.48)粒·m-2>茶园(51.67±10.17)粒·m-2;茶园土壤种子库物种丰富度要显著低于其他类型。4种土地利用类型土壤种子库生活型组成差异极显著,主要以草本植物组成,以菊科与禾本科占优势;针阔混交林的草本植物种子密度最多,非森林的原生物种是草本植物的主要组成;针叶林外来物种的种子密度要显著高于其他类型,紫茎泽兰(Eupatorium adenophorum)是其主要组成。土壤种子库与地上植被的相似性系数较低,其大小顺序为:次生季风常绿阔叶林(0.175)<针阔混交林(0.176)<针叶林(0.215)。     

Seed bank and vegetation composition of forest stands of varying age in central Belgium: consequences for regeneration of ancient forest vegetation

Vegetation composition differs significantly between ancientand recent forest, due to slow colonization capacity of typical forest speciesand the higher abundance of early successional species in recent forest.However, little is known about differences in persistent seed bank compositionbetween ancient and recent forest and about the interaction between seed bankand vegetation in relation with forest age. We surveyed the seed bank and theunderstorey vegetationcomposition in transects from ancient to recent forest. Seed bank and fieldlayer vegetation characteristics and similarity between seed bank andvegetationwere analysed in relation to recent forest age and distance to the ancientforest. A total of 39 species and 14,911 seedlings germinated, whichcorresponds with a seed density of 12,426 seeds/m².Total seed density is significantly higher in the youngest recent forest parcel(55 years). Also the seed bank composition in the youngest forest parceldifferssignificantly from the other parcels. After a longer period of reforestation,the seed bank approaches that of the ancient forest, suggesting seed bankdepletion, although the seed bank is permanently replenished to some extent byseed bank forming species from local disturbances. Seed bank composition doesnot change significantly with distance to the ancient forest. Similaritybetween seed bank and vegetation composition, nomatter the forest age, is very low, but decreases with increasing forest age.The most frequent species in the vegetation are absent in the seed bank andvice versa. The contribution of forest species is highin the vegetation and they almost not occur in the seed bank, while species offorest edges and clearings, and species of disturbed environments are morefrequent in the seed bank. The seed bank is mainly composed of earlysuccessional species of former forest stages or species which temporary occurinsmall-scale disturbances. The seed bank may enhance the negative effects ofearlysuccessional, mainly competitive species to the forest species richness in therecent forest. In this respect, forest management should minimise forestdisturbances, to prevent germination of competitive species form buriedseeds.     

Inflow of seeds through the forest edge: evidence from seed bank and vegetation patterns

To determine the influence of the proximity of a forest edge on seed bank composition and diversity, we performed a seed bank sampling at ancient deciduous forests bordering intensive arable fields. Also vegetation patterns were taken into account. We hypothised that forest edges may facilitate the entrance of diaspores of invasive species into the forest and the successive incorporation of these species in the forest seed bank. We noticed a substantial influence of the proximity of an edge on seed bank composition at as well the forested side of the edge as the field side. The forest edge zone was limited to 3 m into the forest and the field edge zone extended 3m into the field. The seed bank samples of field and forest edge are characterised by a higher species diversity and seed density and a higher similarity between seed bank and vegetation, compared to field or forest samples. The forest edges contains fewer pioneer species in comparison with the forest interior and more competitive species and species of edges and clearings compared with field and forest samples. The seed longevity index increases towards the forest interior. We can conclude from our data that the forest and edge seed bank are composed by both seeds from recent dispersal processes and local seed set and by seeds originating from past vegetation on the site. Near the edge, actual seed input seems of primal importance. Further towards the forest interior seed input decreases and long-living seeds of past vegetation become more important. Ancient forest edges thus act as a barrier for seeds of species of the surrounding arable field.     

Regeneration Potential of Floodplain Forests Under the Influence of Nonnative Tree Species: Soil Seed Bank Analysis in Northern Italy

Soil seed banks are the ecological memory of plant communities and might represent their regeneration potential. This study examines the soil seed bank in hardwood floodplain forests of the biosphere reserve “Valle del Ticino” (Northern Italy) to find out whether the natural forest vegetation can potentially be restored by the soil seed bank. We compared near natural forests of the phytosociological association Polygonato multiflori–Quercetum roboris with stands dominated by the nonnative tree species Robinia pseudoacacia and Prunus serotina in order to investigate whether the composition of the soil seed bank is significantly influenced by the composition of the main canopy tree species and soil properties. Soil seed bank samples were taken from 20 randomly selected plots in stands that were differentiated into four groups related to the dominant forest canopy species. The germinated plants were counted and their species determined. A total of 2,427 plants belonging to 84 species were recorded. The composition of the dominant tree species and soil parameters significantly influence the composition of the seed bank. The similarity with the standing vegetation was very low. Only 13% of the species in the soil seed bank represent the target vegetation. The low percentage of target species and the high percentage of nonnative species imply that the regeneration of near‐natural forest vegetation from the soil seed bank is not feasible. Consequently, disturbances that may activate the soil seed bank should be minimized. Thus, we recommend stopping the mechanical removal of the nonnative tree species in the Ticino Park .     

间伐强度对阿尔泰山天然林下植被的影响

以阿尔泰山天然林为研究对象,探究不同间伐强度（19%、33%、55%、62%）对林下植被的影响,为该区天然林的经营管理提供理论依据。运用样地调查法,对间伐30 a后的林木进行每木检尺,调查林下植被结构、物种多样性,对测定数据进行单因素方差分析和LSD多重比较检验。结果表明：（1）草本层物种丰富度随间伐强度的增大而增大,62%间伐强度显著大于其他处理;灌木层物种丰富度随间伐强度的增大呈先增后减的趋势,55%间伐强度最大（P>0.05）;不同间伐强度处理之间,林下植被的Simpson指数、Shannon-Wiener指数和Pielou均匀度指数均无显著差异（P>0.05）,草本植物优势种各间伐强度之间大致相同;（2）适当增加间伐强度可以明显提高灌木层和草本层密度,以55%间伐强度为优（P<0.05）,林下植被的盖度随着间伐强度的增大先增后减,间伐显著影响灌木植物盖度,而对草本层盖度则影响不显著（P>0.05）。以上研究表明从恢复林地多样性的角度考虑,间伐强度选取55%更合理。     

生境对啮齿动物介导下锐齿槲栎种子命运的影响

干扰作为森林演替的驱动力,在改变森林结构和地表植被的同时也对林内动物介导下的种子扩散过程产生影响.本研究采用塑料标签标记法,选取秦岭林区不同人为干扰后形成的3种类型生境(未抚育林分、抚育3年后林分、裸地),研究啮齿动物扩散行为对不同生境类型锐齿槲栎种子命运的响应.结果表明:抚育3年后林分的种子扩散速率最大,裸地扩散速率最小.裸地的种子原地取食率显著低于未抚育(25.0%)和抚育3年后林分(36.3%),而抚育3年后林分的种子搬运后取食率显著高于未抚育林分(17.3%)和裸地(5.0%),且抚育3年后的林分中被啮齿动物分散埋藏的种子比例最高(4.3%).种子扩散距离以抚育3年后林分最大(26 m),且扩散距离显著大于裸地和未抚育林分.锐齿槲栎种子的初次扩散过程在不同生境中表现不同,其种子后期成苗率也存在差异.不同类型生境在一定程度上影响啮齿动物的觅食策略,最终导致不同的种子扩散模式和森林天然更新格局的形成.     

宝华山主要植被类型土壤种子库存初探

落叶常绿阔叶混交林和落叶阔叶林的土壤种子库分别有49种和33种植物,其中,前者有草本42种,灌木3种,乔木4种,密度为255粒／m2;后者有草本29种,灌木1种,乔木3种,密度为145粒／m2。两植被类型的土壤种子库组成与其现存地上植被的组成相似性极小。落叶常绿阔叶混交林林窗的土壤种子库有33个种,密度为413粒／m2,与非林窗区域的土壤种子库组成仅有53.1%的相似性。     

Forest Succession and Distance from Forest Edge in an Afro‐Tropical Grassland1

Forest succession on degraded tropical lands often is slowed by impoverished seed banks and low rates of seed dispersal. Within degraded landscapes, remnant forests are potential seed sources that could enhance nearby forest succession. The spatial extent that forest can influence succession, however, remains largely unstudied. In abandoned agricultural lands in Kibale National Park, Uganda, recurrent fires have helped perpetuate the dominance of tall (2–3 m) grasses. We examined the effects of distance from forest and grassland vegetation structure on succession in a grassland having several years of fire exclusion. At 10 and 25 m from forest edge, we quantified vegetation patterns, seed predation, and survival of planted tree seedlings. Natural vegetation was similar at both distances, as was seed (eight species) and seedling (six species) survival; however, distance may be important at spatial or temporal scales not examined in this study. Our results offer insight into forest succession on degraded tropical grasslands following fire exclusion. Naturally recruited trees and tree seedlings were scarce, and seed survival was low (20% after 7 mo). While seedling survival was high (95% after 6 to 8 mo), seedling shoot growth was very slow (x?= 0.5 cm/100 d), suggesting that survivorship eventually may decline. Recurrent fires often impede forest succession in degraded tropical grasslands; however, even with fire exclusion, our study suggests that forest succession can be very slow, even in close proximity to forest.     

Recovery of amphibian,reptile, bird and mammal diversity during secondary forest succession in the tropics

In tropical regions, many studies have focused on how vegetation and ecosystem processes recover following the abandonment of anthropogenic activities, but less attention has been given to the recovery patterns of vertebrates. Here we conduct a meta‐analysis (n = 147 studies) of amphibian, reptile, bird and mammal recovery during tropical secondary forest succession (i.e. natural regeneration). For each taxonomic group, we compared changes in species richness and compositional similarity during natural secondary succession to reference forests (mature or old growth forest). In addition, we evaluated the response of forest specialists and the change in bird and mammal functional groups during natural secondary succession in the tropical moist forest biome. Overall, species richness of all groups reached levels of the reference forests during natural secondary succession, but this was not the case for species compositional similarity. The delay in recovery of forest specialists may be the reason for the delay in recovery of species compositional similarity. Overall, vertebrate recovery increased with successional stage, but other potential predictors of diversity recovery, such as, the geographical setting (amphibian and reptile species compositional similarity recovered more rapidly on islands), rainfall (mammal species richness and compositional similarity recovered faster in regions of low rainfall), and the landscape context (amphibian, reptile and mammal species compositional similarity recovered faster in regions with more forest patches) influenced vertebrate recovery. These results demonstrate the important role of secondary forests in providing habitat for many vertebrates, but the slow recovery of species compositional similarity, forest specialists and some functional groups (e.g. insectivorous birds) highlighted the challenge of secondary forest persistence, and strongly argues for the continued protection of old growth/mature forest as habitat for forest specialists and as sources for secondary forest sites.     

Bone-seed (

Bone-seed, Chrysanthemoides monilifera ssp. monilifera (L.), is an environmental weed of coastal vegetation communities scattered throughout New Zealand. To assess the long-term implications for native forest regeneration in sites where bone-seed is present, we selected four study sites around Wellington, New Zealand, where bone-seed was abundant. We compared seed bank composition in bone-seed-invaded sites with nearby native forest patches, and monitored bone-seed and native seedling recruitment with and without control of mature bone-seed plants. We also tested the potential effects of fire on bone-seed recruitment in these communities by heating seeds prior to germination. Bone-seed, gorse (Ulex europaeus), and native species emerged from seed bank samples taken from bone-seed-invaded sites, but only native species and (less) gorse emerged from seed bank samples taken from native forest patches. Gorse germination was strongly promoted by heat but bone-seed germination was less affected by heat. Bone-seed seedling abundance increased dramatically following canopy removal, whereas native seedling abundance decreased dramatically. This suggests that disturbance of any form is likely to favour recruitment of bone-seed (and gorse) over native species, although in the long term, native seedlings can establish beneath the canopy of mature bone-seed plants. It is not yet known if, in the absence of further disturbance, regenerating native vegetation will eventually replace bone-seed in New Zealand.     

Post-cultivation Secondary Succession in a Venezuelan Lower Montane Rain Forest

Since tropical rain forests are widely threatened by conversion to agriculture, even within protected areas, an understanding of recovery processes is important for restoration of forest ecosystems and thus conservation of their biodiversity. Secondary succession following land clearance and crop cultivation was studied in a lower montane rain forest in a protected area of the Venezuelan Cordillera de la Costa Central. Forest recovery was studied using a chronosequence of eight 20 × 20 m plots which represented four forest types ca.10 year-old Secondary Forest, ca. 20 year-old Secondary Forest, ca. 35 year-old (uncultivated) secondary forest and mature forest. Species richness and structural complexity increased during succession, with the oldest secondary forest having a physiognomy comparable to the mature forest. Species diversity was lower in the secondary forests than the mature forest, and their floristic composition was distinct. Four phases are hypothesized to occur in the succession process, each with a distinctive species assemblage: initial colonisation by non-woody vegetation; establishment and canopy closure by short-lived small-seeded woody pioneer species; replacement by longer-lived secondary species; and gradual replacement by mature forest large-seeded climax species. Full recovery of the forests in the protected area is likely to take many years, although it may be assisted through conservation management measures.     

Seed bank dynamics and tree‐fall gaps in a northwestern Mexican Quercus‐Pinus forest

Questions: How does the seed bank respond to different types of tree‐fall gaps and seasonal variations? How does the soil seed bank influence recovery of the standing vegetation in the mature forest and tree‐fall gaps? Location: 1800 — 2020 m a.s.l., Quercus‐Pinus forest, Baja California Sur, Mexico. Methods: Seed size, species composition and germination were estimated under different environmental conditions during dry and rainy seasons: a mature forest plot and gaps created by dead standing trees, snapped‐of f trees and uprooted trees. The soil seed bank was investigated using direct propagule emergence under laboratory conditions, from soil cores obtained during both seasons. Results: 21 species, 20 genera and 14 families constitute the seed bank of this forest community. Fabaceae, Asteraceae, Euphorbiaceae and Lamiaceae were the most frequently represented families in the seed bank. Floristic composition and species richness varied according to the different modes of tree death. Species composition of seed banks and standing vegetation had very low similarity coefficients and were statistically different. Seed bank sizes varied between 164 and 362 ind.m^‐2 in the mature forest plot for the dry and rainy seasons, respectively, while soil seed bank sizes for gaps ranged between 23–208 ind.m^‐2 forthe dry season and between 81–282 ind.m^‐2 for the rainy season. Conclusions: Seed bank sizes and germination response were always higher in the rainy season under all the environmental conditions analysed. Results suggest that timing responses to gap formation of the soil seed bank could be more delayed in this temperate forest than expected.     

Soil seed bank and floristic diversity in a forest‐grassland mosaic in southern Spain

Abstract. Soil seed bank and floristic diversity were studied in a forest of Quercus suber, a forest of Quercus canariensis and a grassland, forming a vegetation mosaic in Los Alcornocales Natural Park, southern Spain. The soil seed bank was estimated by the germination technique. In each community patch, diversity, woody species cover and herbaceous species frequency was measured. Three biodiversity components – species richness, endemism and taxonomic singularity – were considered in the vegetation and the seed bank. Forest patches had a soil seed bank of ca. 11 200–14 100 seed.m^?2 and their composition had low resemblance to (epigeal) vegetation. The grassland patch had a more dense seed bank (ca. 31 800 seed.m^?2) and a higher index of similarity with vegetation, compared with the forests nearby. The complete forest diversity was 71–78 species on 0.1 ha, including 12–15 species found only in the seed bank; the grassland species richness was higher (113 species on 0.1 ha). We discuss the role of soil seed banks in the vegetation dynamics and in the complete plant biodiversity of the mosaic landscape studied.     

Establishment of Epiphytic Bromeliads in Successional Tropical Premontane Forests in Costa Rica

Plant community composition is the combined result of species-specific competitive abilities and the availability of propagules. For epiphytic plants, current hypotheses consider that dispersal-related factors are most important. By controlling seed dispersal constraints, we experimentally examined whether the community composition of epiphytic bromeliads in a tropical premontane area is determined during early phases of seedling recruitment. Also, we tested whether establishment success was related to eco-physiological traits of the species. A total of 7200 seeds were artificially affixed on several host trees in two secondary forest patches and in a mature forest stand. Four bromeliad species with differing physiological characteristics (CAM, C₃-CAM, and C₃) and habitat preference (secondary vs. primary forest) were selected. We found that differences in seed germination probability among habitats and species were not likely to influence community assembly. After 2 yr, seedling survival and plant development were relatively higher in the early-successional forest. Seedling establishment success was not associated with specific physiological and morphological adaptations or habitat preference of the studied species. Our results were not consistent with the described community composition and rates of population recruitment of the studied species in the same successional habitats. The results support the hypothesis that chance and historic events related to seed dispersal have an important influence on community assembly of epiphytic plants. In addition, differences in growth rates and reproductive turnover among species are expected to influence the relative abundance and recruitment rates in a particular habitat.     

Revealing secondary seed removers: results from camera trapping

This paper reports the results of the first study on secondary seed removal of seeds dispersed by Sykes’ monkeys (Cercopithecus albogularis) using camera traps in Africa. Patterns of primary seed dispersal are often superimposed by secondary conveyance, emphasising the need to study these secondary processes carefully. As the agents and mechanisms of seed dispersal are often concealed, being carried out by cryptic or nocturnal animals in dense vegetation, camera trapping was deemed a viable means to investigate secondary removal of seeds disseminated by C. albogularis in the Western Soutpansberg, South Africa. Camera traps were established at preferred feeding trees of the focal Sykes’ monkey group to identify animal species that remove seeds and fruits spat and dropped to the forest floor and seed removal observations were carried out. This method proved to be effective in identifying seed removers and also allowed to get indications about the quantities of seeds removed. Ten animal species were recorded visiting the trees, of which eight were observed removing seeds and fruits. Overall seed and fruit removal rates were high, indicating that the foraging behaviour of C. albogularis attracts many terrestrial frugivores.     

南亚热带森林不同演替阶段土壤种子库的初步研究

本文对鼎湖山不同演替阶段的森林（马尾松林、针阔叶混交林、季风常绿阔叶林）的土壤种子库进行了初步研究．分别从各个阶段的样地内抽取４或８个１×１ｍ的小样地,分三层（共１０ｃｍ厚）采集土样,带回实验室,通过萌发法观察记录其土壤种子库状况．通过统计分析,其结果如下：１．种子数量和物种多样性一般随演替发展而减少,种子数目以３—５ｃｍ厚土层为最多．２．各演替阶段土壤种子库的种类组成均以草本植物为主．３．在光照和湿度基本一致的情况下,萌发种子数与温度成正相关．４．雨季取的种子数目较旱季多,且种类组成有所不同．这是因为南亚热带森林内植物种子的休眠期短,在旱季采集土样时许多种的种子尚未下落．５．种子库组成与地上植物相关性不明显,但演替早期阶段的相关性比演替后期更密切。