Seed dispersal by wind,birds, and bats between Philippine montane rainforest and successional vegetation

In the moist Neotropics, vertebrate frugivores have a much greater role in the dispersal of forest and successional woody plants than wind, and bats rather than birds play the dominant role in dispersing early successional species. I investigated whether these patterns also occurred in a Philippine montane rainforest and adjacent successional vegetation. I also asked whether seed mass was related to probability of dispersal between habitats. A greater number of woody species and stems in the forest produced vertebrate-dispersed seeds than wind-dispersed seeds. Although input of forest seeds into the successional area was dominated by vertebrate-dispersed seeds in terms of species richness, wind-dispersed seeds landed in densities 15 times higher. Frugivorous birds dispersed more forest seeds and species into the successional area than bats, and more successional seeds and species into the forest. As expected, seed input declined with distance from source habitat. Low input of forest seeds into the successional area at the farthest distance sampled, 40 m from forest edge, particularly for vertebrate-dispersed seeds, suggests very limited dispersal out of forest even into a habitat in which woody successional vegetation provides perches and fruit resources. For species of vertebrate-dispersed successional seeds, probability of dispersal into forest declined significantly with seed mass.     

Fleshy fruits of indigenous and adventive plants in the diet of birds in forest remnants, Nelson, New Zealand

The relationship between fleshy-fruited indigenous species and adventive weeds in the diet of 500 mist-netted birds was studied in forest remnants of differing size and degree of modification. Fruit abundance Peaked in March and April, and most fruit was either red/orange or purple/black. The physical parameters of adventive and indigenous fruits were not significantly different. Six of the 15 passerine species netted are frugivores, and of those netted 77% had eaten fruit. They were divisible into three groups: endemic (bellbirds, Anthornis melanura; tuis, Prosthemadera novaeseelandiae), non-endemic but indigenous (silvereyes, Zosterops lateralis), and adventive (blackbirds, Turdus merula; song thrushes, T. philomelos; starlings, Sturnus vulgaris). Bird diets varied between the groups and according to fruit availability as determined by sires and seasons. Endemic birds ate the least adventive fruit; bellbirds ate mainly Podocarpus hallii and Coprosma robusta fruits at all sites. Tuis had a varied diet, including some adventive fruits. Silvereyes ate the widest range of indigenous and adventive fruits. Blackbirds and, to a smaller extent, song thrushes ate many of the same indigenous fruits as the other bird groups, but their diet included more adventive fruits, e.g., Berberis glaucocarpa. Starlings were caught only when they fed on Sambucus nigra, but they also ate a few indigenous fruits. There was little seasonal variation in bird numbers caught. Adventive species extended the seasonal availability of fruits into winter, particularly in the forest remnant closest to a town, which had the highest proportion of adventive fruits. Several weed species distributed mainly by non-endemic and adventive birds are forming new secondary vegetation. Some have large fruit crops which generally offer little food for endemic birds. Where fruiting weeds pre-empt sites that may have been occupied by native species, they create an inferior habitat for endemic birds. However, the non-endemic and adventive birds also disperse indigenous fruits into early successional vegetation, and the importance of their seed rain for conservation of biodiversity will therefore depend on the site.     

Remnant Trees Affect Species Composition but Not Structure of Tropical Second-Growth Forest

Remnant trees, spared from cutting when tropical forests are cleared for agriculture or grazing, act as nuclei of forest regeneration following field abandonment. Previous studies on remnant trees were primarily conducted in active pasture or old fields abandoned in the previous 2–3 years, and focused on structure and species richness of regenerating forest, but not species composition. Our study is among the first to investigate the effects of remnant trees on neighborhood forest structure, biodiversity, and species composition 20 years post-abandonment. We compared the woody vegetation around individual remnant trees to nearby plots without remnant trees in the same second-growth forests (“control plots”). Forest structure beneath remnant trees did not differ significantly from control plots. Species richness and species diversity were significantly higher around remnant trees. The species composition around remnant trees differed significantly from control plots and more closely resembled the species composition of nearby old-growth forest. The proportion of old-growth specialists and generalists around remnant trees was significantly greater than in control plots. Although previous studies show that remnant trees may initially accelerate secondary forest growth, we found no evidence that they locally affect stem density, basal area, and seedling density at later stages of regrowth. Remnant trees do, however, have a clear effect on the species diversity, composition, and ecological groups of the surrounding woody vegetation, even after 20 years of forest regeneration. To accelerate the return of diversity and old-growth forest species into regrowing forest on abandoned land, landowners should be encouraged to retain remnant trees in agricultural or pastoral fields.     

Patterns of Seed Dispersal and Dispersal Failure in a Hawaiian Dry Forest Having Only Introduced Birds

Dry forests are among the most endangered natural communities in the Hawaiian Islands. Most have been reduced to isolated trees and small forest fragments in which native tree species reproduce poorly. The replacement of native birds by introduced generalists may be contributing to dry forest decline through modification of seed dispersal patterns. To document seed dispersal by introduced birds, we conducted foraging observations on fleshy-fruited trees and measured seed rain under trees and in adjacent open areas for 1 year in a dry forest dominated by native trees. Although trees covered only 15.2 percent of the study area, 96.9 percent of the bird-dispersed seeds were deposited beneath them. The Japanese white-eye (Zosterops japonicus) was the principal dispersal agent. Among bird-dispersed seeds, those of the invasive tree Bocconia frutescens accounted for 75 percent of all seeds collected beneath trees (14.8 seeds/m²/yr) and the invasive shrub Lantana camara accounted for 17 percent. Although nearly 60 percent of the reserve's native woody species possess fleshy fruits, introduced birds rarely disperse their seeds. Native trees accounted for <8 percent of all bird-dispersed seeds and are consequently experiencing dispersal failure by falling directly under parent trees. Smaller-seeded non-native plants, in contrast, may be benefiting from dispersal by introduced birds. Current dispersal patterns suggest that these readily disseminated non-native plants may eventually replace the remaining native flora.     

Recovery of rain forest soil seed banks under different reforestation pathways in eastern Australia

Summary Seed availability is a major factor limiting the recruitment of rain forest to cleared land, but little is known about the composition of the soil seed bank under different reforestation pathways. We quantified changes in the viable soil seed bank following rain forest clearing and pasture establishment and subsequent reforestation in subtropical eastern Australia. Major reforestation pathways in the region include planting of a diverse suite of native trees for ecological restoration purposes, autogenic regrowth dominated by the non‐native tree Camphor Laurel (Cinnamomum camphora) and management of this regrowth to accelerate the development of a native tree community. These pathways differ considerably in cost: restoration plantings are expensive, autogenic regrowth is free, whilst managing regrowth generally costs much less than restoration plantings. We surveyed five sites within each of three reforestation pathways as well as reference sites in remnant rain forest and pasture. The composition of the seed bank was determined by germinating plants from soil samples collected from each site. Germinants were classified into several functional groups according to life form, origin, dispersal mode and successional stage. The majority of functional groups varied significantly in abundance or richness between rain forest and pasture sites. Most of the functional groups that varied between rain forest and pasture were restored to values similar to rain forest by at least one of the three reforestation pathways examined. The species richness of native woody plants in the soil seed bank was slightly higher in restoration plantings than in autogenic or managed regrowth; nevertheless, the species richness and abundance of native woody plants and vines were higher in the seed bank of autogenic regrowth than pasture, and both attributes were enhanced by the management of regrowth sites. The results of this study show that autogenic regrowth can make an important contribution to rain forest restoration at a landscape scale. The optimal reforestation approach or mix of approaches will depend on the desired rate of recovery and the resources available for restoration.     

Post‐Establishment Seedling Success in the Brazilian Cerrado: A Comparison of Savanna and Forest Species1

Due to frequent fire, low nutrient availability, and prolonged drought, tropical savanna is a stressful environment for the survival and growth of woody plant seedlings. To understand why forest species do not succeed in this environment while savanna species are able to persist, the effects of fire and woody cover on seedlings of these two functional groups were investigated in the Brazilian Cerrado. Seedlings were established in experimental plots under three densities of woody cover, in sites protected from fire and sites to be subjected to fire. There was a clear difference in the ability of savanna and forest species to survive fire. None of the three forest species were able to survive fire during the first two years of life, whereas eight of the nine savanna species were able to resprout following fire. The small seed size of the ninth savanna species, Miconia albicans, predisposed its seedlings to be sensitive to fire, because there was a strong positive correlation between seed size and survivorship. Savanna species were less dependent on woody cover than were forest species, which exhibited higher growth and survival under tree canopies than in open grassland. The low rates of establishment and survival of forest trees in savanna, combined with high sensitivity to fire, appear sufficient to prevent the expansion of forest into savanna under current fire regimes in the Cerrado.     

Role of nurse plants in Araucaria Forest expansion over grassland in south Brazil

Abstract Araucaria Forest expansion over grassland takes place under wet climate conditions and low disturbance and it is hypothesized that isolated trees established on grassland facilitate the establishment of forest woody species beneath their canopies. Forest with Araucaria angustifolia is a particular type of Brazilian Atlantic Forest and the main forest type on the highland plateau in south Brazil, often forming mosaics with natural Campos grassland. The objectives of this paper were to evaluate the role of isolated shrubs and trees as colonization sites for seedlings of Araucaria Forest woody species on grassland, to determine which species function as preferential nurse plants in the process and the importance of vertebrate diaspore dispersal on the structure of seedling communities beneath nurse plants. The study was conducted in São Francisco de Paula, Rio Grande do Sul State, where we sampled isolated shrubs and trees in natural grassland near Araucaria Forest edges. Seedlings were counted and identified, and seedling diaspore dispersal syndromes, size and colour were registered. We detected 11 woody species with a potential role in nucleating grassland colonization by forest species. Beneath the canopies of nurse plants more forest species seedlings were found compared with open field grassland and the seedlings had diaspores mostly dispersed by vertebrates. Also, more seedlings were found under the canopy of A. angustifolia than beneath other nurse plant species. We conclude that A. angustifolia trees established on grassland act as nurse plants, by attracting disperser birds that promote colonization of the site by other forest species seedlings, and that under low level of grassland disturbance, conservation of frugivorous vertebrate assemblages may increase forest expansion over natural grassland and also facilitate the regeneration of degraded forest areas.     

Life History Traits of Lianas During Tropical Forest Succession

Tropical secondary forests form an important part of the landscape. Understanding functional traits of species that colonize at different points in succession can provide insight into community assembly. Although studies on functional traits during forest succession have focused on trees, lianas (woody vines) also contribute strongly to forest biomass, species richness, and dynamics. We examined life history traits of lianas in a forest chronosequence in Costa Rica to determine which traits vary consistently over succession. We conducted 0.1 ha vegetation inventories in 30 sites. To examine the establishment of young individuals, we only included small lianas (0.5–1.5 cm diameter at 1.3 m height). For each species, we identified seed size, dispersal mode, climbing mode, and whether or not the seedling is self‐supporting. We found a strong axis of variation determined by seed size and seedling growth habit, with early successional communities dominated by small‐seeded species with abiotic dispersal and climbing seedlings, while large‐seeded, animal‐dispersed species with free‐standing seedlings increased in abundance with stand age. Contrary to previous research and theory, we found a decrease in the abundance of stem twiners and no decrease in the abundance of tendril‐climbers during succession. Seed size appears to be a better indicator of liana successional stage than climbing mode. Liana life history traits change predictably over succession, particularly traits related to seedling establishment. Identifying whether these trait differences persist into the growth strategies of mature lianas is an important research goal, with potential ramifications for understanding the impact of lianas during tropical forest succession.     

Influence of boreal forest succession and dead wood qualities on saproxylic beetles

1 Saproxylic insects, a functional group dominated by beetles, are dependent on dead or moribund trees as habitat elements. 2 Although there are few studies of saproxylic insects from the North American boreal zone, European studies demonstrate that forest harvest can lead to a biologically significant decrease in saproxylic beetle diversity. 3 We studied saproxylic beetles in the North American boreal mixedwood forest using flight intercept traps established on naturally dead and girdled trembling aspen and spruce trees along a successional gradient of undisturbed stands from deciduous to coniferous overstory trees. 4 Composition and diversity of beetle assemblages differed among forest successional types. 5 Snag age class was an important determinant of composition for saproxylic beetle assemblages. 6 Multivariate regression analysis of these data indicated that saproxylic beetles are responding to changes in coarse woody debris, and not to the relative densities of canopy tree species, although these variables are strongly correlated. 7 Coarse woody debris management should be a primary concern in forest management plans seeking to conserve saproxylic organisms and the critical ecosystem functions (i.e. nutrient cycling) in which they participate.     

Woody plant assemblages in isolated forest patches in a semiarid agricultural matrix

Spatial and temporal isolation and environmental variability are important factors explaining variation in plant species composition. The effect of fragmentation and disturbance on woody plant species composition was studied using data from 32 remnant church forest patches in northern Ethiopia. The church forests are remnants of dry Afromontane forest, embedded in a matrix of intensively used crop and grazing lands. We used canonical correspondence analysis and partial canonical correspondence analysis to analyze the effects of fragmented and isolated forest-patch identity, environmental and spatial variables on woody plant species composition in different growth stages. The dominance of late successional species was higher at the adult growth stage than seedlings and saplings growth stages. In the adult stages, late successional species like Olea europaea subsp. cuspidate had high frequency of occurrence. Forest patch identity was more important in explaining woody plant assemblages than environmental and spatial variables. For all growth stages combined, environmental variables explained more of the explained total fraction of variation in species composition than spatial variables. Topographic variables best explained variations in species composition for saplings, adults and all growth stages combined, whereas the management regime was most important for seedlings species composition. Our results show that in a matrix of cultivated and grazing land, fragmented and isolated forest patches differ in woody plant species assemblages. Some species are widely distributed and occurred in many patches while other occurred only in one or a few forest patches. Thus, our results indicate that remnant forest patches are important for preserving rare plant species and therefore management practices should focus on minimizing disturbance to the church forests and if possible increase church forest patch size.     

Spontaneous Establishment of Woody Plants in Central European Derelict Sites and their Potential for Reclamation

The performance of woody plants was analyzed in 15 successional seres starting at bare ground in central European manmade habitats. The total cover of woody species after 10 years of succession was significantly related neither to initial soil moisture nor to nitrogen (expressed using Ellenberg indicator values). But the comparison of seres indicates that establishment of woody plants was easier under moderate environmental conditions and retarded in extreme habitats (dry, nutrient-poor, or acid). The arrival of the first woody plants was delayed in dry sites. No significant differences were found between primary and secondary seres, either with respect to the total cover of woody plants reached after 10 years of succession or considering the time of their arrival. In total, 24 woody species (10 shrubs and 14 trees) appeared in the series investigated. Their successional performance (in terms of the number of seres in which the species occurred and maximum cover reached in any sere) was not related to species traits (life strategy, type of mycorrhizae, mode of dispersal, diaspore weight), except for the regeneration strategy, species with seasonal regeneration by seeds were capable of creating higher cover. Betula pendula (European birch) was the most successful species in spontaneous succession, especially on moist sites. Practical suggestions for the management of particular habitats (sites disturbed by mining, sites reclaimed after acid rain deforestation, urban sites, abandoned fields) are provided regarding the establishment of woody plants.     

The role of farm structure on bird assemblages around a Kenyan tropical rainforest

Although it is clear that the farmlands neighbouring fragmented forests are utilized by some forest birds, it is not clear how birds in general respond to farmland habitat mosaic. An effort was made to determine how bird density and foraging assemblages were influenced by farm structural characteristics and distance from forest edge. Thirty farms up to a distance of 12 km around Kakamega forest in western Kenya were studied. Farm structure entailed size, hedge volume, habitat heterogeneity, woody plant density, plant diversity and crop cover. Birds were surveyed using line transects and DISTANCE analyses and classified into six feeding guilds and three habitat associations. Size of farms increased away from the forest, as woody plant density, plant diversity, indigenous trees and subsistence crop cover declined. The most important farm structure variable was hedge volume, which enhanced bird species richness, richness of shrub‐land bird species and insectivorous bird density (R = 0.58, P < 0.01). Bird density increased with tree density while indigenous trees were suitable for insectivores and nectarivores. There were very few forest bird encounters. Agricultural practices incorporating maintenance of hedges and sound selection of agroforestry trees can enhance conservation of birds on farmland, though, not significantly for forest species.     

The Effect of Spatial Structure of Pasture Tree Cover on Avian Frugivores in Eastern Amazonia

The movement of frugivores between remnant forests and successional areas is vital for tropical forest tree species to colonize successional habitats. The response of these species to the spatial structure of pasture tree cover is largely unknown. We studied avian frugivores that were found in primary forest edges and large pastures in eastern Amazonia, Brazil. We determined how the small‐scale spatial structure of pasture trees at forest edges affects five response variables: bird presence, visitation rate, duration of visit, species richness, and an index accounting for species’ level of frugivory and abundance in forests. We used hierarchical linear models to estimate the effect of four predictor variables on response variables: (1) clustering of pasture trees; (2) percent canopy cover of pasture trees; (3) distance of pasture tree to forest edge; and (4) tree crown area. The study species, many of which are widely distributed in the Neotropics, were generally insensitive to percent cover and clustering of trees. Frugivore visitation to individual trees remained constant as cover increased. Visitation was positively correlated with focal tree distance to forest edge and crown area. The positive relationship between distance and visitation rates may be due to the increased abundance of some resource further from forests. If pastures were abandoned the distance from forest edges would not likely limit frugivore visitation and seed deposition under large pasture trees in our study (i.e., up to 200 m distant).     

Seed Dispersal by Birds and Bats in Lowland Philippine Forest Successional Area

In the tropical forests of SE Asia, only a few studies have dealt with the role animal dispersal plays in early forest succession and rehabilitation, and a comparison of bird and bat dispersal is even rarer. We investigated seed dispersal by birds and bats in a successional area in the lowland dipterocarp forest of the Subic Watershed Forest Reserve (SWFR) in Luzon Island, Philippines. Using pairs of day and night traps, we collected seeds during 3 mo of wet season and 3 mo of dry season in a 1.2-ha study site. Bird-dispersed seeds predominated over those dispersed by bats in terms of both seed abundance and number of seed species. The most abundant endozoochorous seed species were significantly biased toward either bird or bat dispersal. Birds and bats appeared to compete more strongly for fruit resources during the dry season than during the wet season, and bats responded more to changes in the seasons than birds did. GLM analyses showed that the factor that had the strongest influence on overall seed distribution was the number of fleshy-fruited trees surrounding the traps, and that the distribution pattern of day-dispersed seeds was affected by more physical factors (number of trees, size of trees, presence of fleshy-fruited and conspecific trees) in the study site than the pattern of night-dispersed seeds were. Given that birds are the more important dispersers in the study site, restoration efforts in SWFR might benefit by focusing on attracting these dispersers into its degraded habitats.     

Average body length of arboreal and aerial beetle (Coleoptera) assemblages from remnant and plantation Eucalyptus forests in southwestern Australia

We examined body length differences between the beetle fauna of Eucalyptus plantation forests and remnant Eucalyptus forest in the same region using aerial and arboreal sampling. Mean body length of plantation-using species was greater than for remnant forest species, and the same pattern was apparent in the mean of all individuals collected regardless of species. This pattern was true for both Malaise-trap-collected beetles (aerial) and canopy-collected (arboreal) beetles. The tendency for plantation-restricted species to have longer bodies is significant even if clade is treated as a random factor in the analysis. Greater body length among plantation-using species is consistent with a few other studies that have found body size of insect species in early successional environments is typically greater than in late successional environments. Some studies suggest that larger species are better dispersers, which can rapidly colonise early successional habitats. In this study, however, there was little relationship between body length and the number of sites occupied; suggesting dispersal was not a major determinant of community membership. Two different patterns in this study support the hypothesis that body size differences are linked to trophic structure of the communities. First, the body length shift comparing remnant users to plantation users was greater among phytophages than for predators or saprophages. Second, saprophages were typically smaller than phytophages, and constituted a larger fraction of the remnant forest using fauna, driving down the mean body length in the saprophage-rich remnant forest community.     

鸟类栖木在森林植被恢复中的生态意义

鸟类传播种子对于植物种群自然更新及群落演替具有重要的生态意义,鸟类栖息的树木具有富集种子雨功能,在山体滑坡区及废弃矿区等干扰生境中人工设置鸟类栖木,可以增加鸟类传播的种子雨输入,丰富土壤种子库,尤其食果鸟类可以传播那些处于群落演替中期至后期的乔、灌木树种,能够促进受干扰地区森林植被的演替进程.5·12汶川大地震对地震灾区造成了大面积的山体滑坡和严重的植被破坏,在地震灾区森林植被的生态恢复工作中,一些地区可以采用人工设置鸟类栖木方法,充分利用山体滑坡区周围残余森林的丰富种子源及众多食果鸟类,促进灾区森林植被的自然恢复.     

Frugivoría por Aves en un Paisaje Fragmentado: Consecuencias en la Dispersión de Semillas1

Few data exist on seed dispersal by frugivorous birds in fragmented landscapes, originating from tropical dry forests, in contrast to more abundant data from tropical rain forests. In this study, we assessed the effect of frugivorous birds in a fragmented landscape of Veracruz, Mexico, now occupied by remnant fragments of tropical semi‐deciduous forest and dry deciduous forest, grassland, and shrubby patches on sand dunes. We determined four characteristics related to seed dispersal by birds: the interacting species of plants and birds, the characteristics of these species, spatio‐temporal variation in the dispersal system, and the outcome of the process. During one year, we recorded 54 frugivorous bird species and 33 ornithochorous plant species, which engaged in 176 different bird‐plant species interactions. Similarity (Sorensen index) of frugivorous bird communities using different vegetation types was high (>70%), suggesting that many bird species used all of the vegetation types. In contrast, the similarity of ornithochorous plant communities among vegetation types commonly was low (<37%), suggesting that most plant species were restricted to particular sites in this landscape. At the landscape level, as well as for tropical deciduous forest, we detected a significant positive relationship (Spearman's correlation of rank coefficient >0.65, P <0.05) among richness per month of frugivorous birds and plant species bearing fleshy fruits. Seeds of many plant species previously detected in studies of seed rain at the site were eaten by birds during this study. Most seeds of zoochorous species, which are deposited in the dry and decidous tropical forests patches, are produced within these vegetation types (i.e., they are autochthonous species), whereas bird‐dispersed seeds arriving in grassland and shrubby patches are produced outside (i.e., allochthonous) and are mostly woody species. Birds are important seed dispersers among vegetation types in this landscape but they have different effects in each one. The four characteristics studied, as well as the landscape approach of this research, allowed us to detect spatial and temporal patterns that otherwise would have remained undetected.     

Landscape and Vegetation Effects on Avian Reproduction on Bottomland Forest Restorations

ABSTRACT Forest restoration has been undertaken on >200,000 ha of agricultural land in the Mississippi Alluvial Valley, USA, during the past few decades. Decisions on where and how to restore bottomland forests are complex and dependent upon landowner objectives, but for conservation of silvicolous (forest-dwelling) birds, ecologists have espoused restoration through planting a diverse mix of densely spaced seedlings that includes fast-growing species. Application of this planting strategy on agricultural tracts that are adjacent to extant forest or within landscapes that are predominately forested has been advocated to increase forest area and enhance forested landscapes, thereby benefiting area-sensitive, silvicolous birds. We measured support for these hypothesized benefits through assessments of densities of breeding birds and reproductive success of 9 species on 36 bottomland forest restoration sites. Densities of thamnic (shrub-scrub dwelling) and silvicolous birds, such as yellow-breasted chat (Icteria virens), indigo bunting (Passerina cyanea), and white-eyed vireo (Vireo griseus) were positively associated with 1) taller trees, 2) greater stem densities, and 3) a greater proportion of forest within the landscape, whereas densities of birds associated with grasslands, such as dickcissel (Spiza americana) and red-winged blackbird (Agelaius phoeniceus), were negatively associated with these variables. Vegetation structure, habitat edge, and temporal effects had greater influence on nest success than did landscape effects. Taller trees, increased density of woody stems, greater vegetation density, and more forest within the landscape were often associated with greater nest success. Nest success of grassland birds was positively related to distance from forest edge but, for thamnic birds, success was greater near edges. Moreover, nest success and estimated fecundity of thamnic species suggested their populations are self-sustaining on forest restoration sites, whereas these sites are likely population sinks for grassland and open-woodland species. We recommend restoration strategies that promote rapid development of dense forest stands within largely forested landscapes to recruit breeding populations of thamnic and silvicolous birds that have reproductive success sufficient to sustain their populations.     

Dynamics of vesicular-arbuscular mycorrhizae during old field succession

Summary The species composition of vesicular-arbuscular mycorrhizal (VAM) fungal communities changed during secondary succession of abandoned fields based on a field to forest chronosequence. Twenty-five VAM fungal species were identified. Seven species were clearly early successional and five species were clearly late successional. The total number of VAM fungal species did not increase with successional time, but diversity as measured by the Shannon-Wiener index tended to increase, primarily because the community became more even as a single species, Glomus aggregatum, became less dominant in the older sites. Diversity of the VAM fungal community was positively correlated with soil C and N. The density of VAM fungi, as measured by infectivity and total spore count, first increased with time since abandonment and then decreased in the late successional forest sites. Within 12 abandoned fields, VAM fungal density increased with increasing soil pH, H₂O soluble soil C, and root biomass, but was inversely related to extractable soil P and percent cover of non-host plant species. The lower abundance of VAM fungi in the forest sites compared with the field sites agrees with the findings of other workers and corresponds with a shift in the dominant vegetation from herbaceous VAM hosts to woody ectomycorrhizal hosts.     

Patterns of woody plant abundance,recruitment, mortality,and growth in a 65 year chronosequence of old-fields

We surveyed vegetation along forest margins in a 65-year chronosequence of old-fields at the Cedar Creek Natural History Area in east-central Minnesota, USA, to identify successional patterns of woody plants and to determine if these were correlated with soil nitrogen. We predicted that shrub and tree abundance, size, and distance of occurrence from the forest edge would be correlated with field age or soil nitrogen. Instead we did not find successional trends in the abundance or composition of woody species. Even in the oldest field the abundance of trees and shrubs was low and concentrated in areas close to the forest. Though trees were larger and present further from the forest edges in older fields, average tree height was less than 126 cm in all fields.Since we did not find successional trends we looked at various local factors (local seed sources, deer browsing, and forest edge aspect) and their relation to recruitment, mortality, or growth to explain variation among fields in abundance of trees or shrubs. The three most common tree species (Quercus rubra, Q. macrocarpa,and Populus tremuloides) all had a higher relative abundance of seedlings, and two (Q. rubra and Q. macrocarpa) had a higher relative abundance of large trees adjacent to forests with a high abundance of conspecific adults. Most trees taller than 20 cm were browsed by deer and were shorter in 1995 than they were in 1993. Mortality was higher for trees less than 30 cm indicating that mortality was size-dependent. Forest edge aspect did not significantly influence the abundance or demography of any species. Our results suggest that the patterns of seedling recruitment were largely determined by the proximity of seed sources and that these patterns may persist so that tree communities in old-fields resemble adjacent forests. Deer may be a significant factor in the suppression of tree populations in old-fields through repeated browsing which reduces tree growth and elevates tree mortality by prolonging the period of time trees remain susceptible to size-dependent mortality.