Understory vegetation response to thinning disturbance of varying complexity in coniferous stands

Question: Can augmented forest stand complexity increase understory vegetation richness and cover and accelerate the development of late‐successional features? Does within‐stand understory vegetation variability increase after imposing treatments that increase stand structural complexity of the overstory? What is the relative contribution of individual stand structural components (i.e. forest matrix, gaps, and leave island reserves) to changes in understory vegetation richness? Location: Seven study sites in the Coastal Range and Cascades regions of Oregon, USA. Methods: We examined the effects of thinning six years after harvest on understory plant vascular richness and cover in 40‐ to 60‐year‐old forest stands dominated by Douglas‐fir (Pseudotsuga menziesii). At each site, one unthinned control was preserved and three thinning treatments were implemented: low complexity (LC, 300 trees ha^?1), moderate complexity (MC, 200 trees ha^?1), and high complexity (HC, variable densities from 100 to 300 trees ha^?1). Gaps openings and leave island reserves were established in MC and HC. Results: Richness of all herbs, forest herbs, early seral herbs and shrubs, and introduced species increased in all thinning treatments, although early seral herbs and introduced species remained a small component. Only cover of early seral herbs and shrubs increased in all thinning treatments whereas forest shrub cover increased in MC and HC. In the understory, we found 284 vascular plant species. After accounting for site‐level differences, the richness of understory communities in thinned stands differed from those in control stands. Within‐treatment variability of herb and shrub richness was reduced by thinning. Matrix areas and gap openings in thinned treatments appeared to contribute to the recruitment of early seral herbs and shrubs. Conclusions: Understory vegetation richness increased 6 years after imposing treatments, with increasing stand complexity mainly because of the recruitment of early seral and forest herbs, and both low and tall shrubs. Changes in stand density did not likely lead to competitive species exclusion. The abundance of potentially invasive introduced species was much lower compared to other plant groups. Post‐thinning reductions in within‐treatment variability was caused by greater abundance of early seral herbs and shrubs in thinned stands compared with the control. Gaps and low‐density forest matrix areas created as part of spatially variably thinning had greater overall species richness. Increased overstory variability encouraged development of multiple layers of understory vegetation.     

Nonnative plant invasion increases urban vegetation structure and influences arthropod communities

Aim

Ecological theory and empirical evidence indicate that greater structural complexity and diversity in plant communities increases arthropod abundance and diversity. Nonnative plants are typically associated with low arthropod abundance and diversity due to lack of evolutionary history. However, nonnative plants increase the structural complexity of forests, as is common in urban forests. Therefore, urban forests are ideal ecosystems to determine whether structural complexity associated with nonnative plants will increase abundance and diversity of arthropods, as predicted by complexity literature, or whether structural complexity associated with nonnative plants will be depauperate of arthropods, as predicted by nonnative plant literature.

Location

We sampled 24 urban temperate deciduous and mixed forests in two cites, Raleigh, North Carolina and Newark, Delaware, in the eastern United States.

Methods

We quantified ground cover vegetation and shrub layer vegetation in each forest and created structural complexity metrics to represent total, nonnative and native understory vegetation structural complexity. We vacuum sampled arthropods from vegetation and quantified the abundance, biomass, richness and diversity of spiders and non-spider arthropods.

Results

Nonnative plants increase understory vegetation complexity in urban forests. In Raleigh and Newark, we found support for the hypotheses that dense vegetation will increase arthropod abundance and biomass, and against the hypothesis that nonnative vegetation will decrease arthropods. Urban forest arthropod abundance and biomass, but not diversity, increased with greater nonnative and native structural complexity.

Main Conclusions

Invaded urban forests may provide adequate food in the form of arthropod biomass to transfer energy to the next trophic level, but likely fail to provide ecological services and functions offered by diverse species, like forest specialists. Urban land managers should survey urban forests for nonnative and native plant communities and prioritize replacing dense nonnative plants with native species when allocating vegetation maintenance resources.     

Reproduction,foraging and the negative density–area relationship of a generalist rodent

While many species show positive relationships between population density and habitat patch area, some species consistently show higher densities in smaller patches. Few studies have examined mechanisms that may cause species to have negative density–area relationships. We tested the hypothesis that greater reproduction in edge versus interior habitats and small versus large fragments contributes to higher densities of white-footed mice (Peromyscus leucopus) in small versus large forest fragments. We also examined vegetation structure and foraging tray utilization to evaluate if greater reproduction was a result of higher food availability. There were greater number of litters and proportion of females producing litters in the edge versus interior of forest fragments, which may have contributed to greater population growth rates and higher densities in edge versus interior and small versus large fragments. Data on vegetation structure and giving-up densities of seeds in artificial patches suggest that food availability may be higher in edge versus interior habitats and small versus large fragments. These results, in an area with few or no long-tailed weasels, provide a distinct contrast to the findings of Morris and Davidson (Ecology 81:2061, 2000) who observed lower reproduction in forest edge habitat as a result of high weasel predation, suggesting that specialist predators may be important in affecting the quality of edge habitat. While we cannot exclude the potential contributions of immigration, emigration, and mortality, our data suggest that greater reproduction in edge versus interior habitat is an important factor contributing to higher densities of P. leucopus in small fragments.     

Migratory bird community structure in oil palm (Elaies guineensis) plantations and native forest fragments in southern Mexico

Oil palm (Elaies guineensis) plantations are among the fastest growing agroecosystems in the Neotropics, but little is known about how Neotropical birds use oil palm habitats. To better understand the potential value of oil palm as an overwintering habitat for migratory birds, we surveyed birds in oil palm and native forest remnants in Tabasco, Mexico, from 19 December 2017 to 27 March 2018. We collected data on bird abundance and vegetative structure and used generalized linear models and multivariate analysis to assess how oil palm development influenced migrant bird diversity, community assemblages, and abundance. We found that species richness of migratory birds tended to be higher in forest patches than in oil palm, that community assemblages of migratory birds differed between native forest and oil palm plantations, and that differences in migratory bird abundance, and subsequent changes in community assemblages were driven by differences between native forest and oil palm plantations in vegetative structure. The bird community of native forest was characterized by migrant species sensitive to forest loss that forage low in the understory and in the leaf litter, whereas the bird community of oil palm plantations was represented by generalist species that occupy a wider range of foraging niches. Our results suggest that most species of migrant birds responded positively to several forest structural features and that integrating more native trees and increasing the amount of understory vegetation in oil palm plantations may increase the value of working landscapes for migratory birds.     

桉树人工林更新方式对林下植物功能群的影响

该研究综合运用野外调查和室内分析的方法,评估桉树人工林林下植物功能群的组成、分布及更新方式和相关环境因子之间的关系。结果表明:林分更新5 a后,除了非禾本科杂草功能群外,其他林下植物功能群的物种丰富度均呈现不同程度的增加,但与对照组(砍伐迹地)相比,其差异程度均不显著(P0.05);与对照组相比,藤本和蕨类功能群的相对多度也出现增加趋势,但禾本科草本功能群显著减少(P0.05);木本、藤本和蕨类功能群的相对盖度也呈现增加趋势,但禾本科草本功能群与对照组相比显著减少(P0.05);主成分分析(PCA)发现萌芽更新或植苗更新林的林下植物功能群组成和分布与对照组相比均发生了显著的变异,但不同更新方式(萌芽和植苗)下其林下植物功能群组成和分布差异不明显;通过冗余分析(RDA)确定了冠层透光系数、土壤孔隙度、坡向和土壤氮磷比是影响该林地林下植物功能群的主要因子,它们的叠加效应能解释大于75%的林下植物功能群的变异,最终模型通过排序得到冠层透光系数是影响该林地林下植物功能群的最主要因子。短期的研究发现萌芽和植苗这两种不同的更新方式对桉树林下植物功能群的影响有限,这可能与这两种更新方式形成的林冠结构和土壤理化性质差异性较小有关。     

Simplifying understory complexity in oil palm plantations is associated with a reduction in the density of a cleptoparasitic spider,Argyrodes miniaceus (Araneae: Theridiidae), in host (Araneae: Nephilinae) webs

Expansion of oil palm agriculture is currently one of the main drivers of habitat modification in Southeast Asia. Habitat modification can have significant effects on biodiversity, ecosystem function, and interactions between species by altering species abundances or the available resources in an ecosystem. Increasing complexity within modified habitats has the potential to maintain biodiversity and preserve species interactions. We investigated trophic interactions between Argyrodes miniaceus, a cleptoparasitic spider, and its Nephila spp. spider hosts in mature oil palm plantations in Sumatra, Indonesia. A. miniaceus co‐occupy the webs of Nephila spp. females and survive by stealing prey items caught in the web. We examined the effects of experimentally manipulated understory vegetation complexity on the density and abundance of A. miniaceus in Nephila spp. webs. Experimental understory treatments included enhanced complexity, standard complexity, and reduced complexity understory vegetation, which had been established as part of the ongoing Biodiversity and Ecosystem Function in Tropical Agriculture (BEFTA) Project. A. miniaceus density ranged from 14.4 to 31.4 spiders per square meter of web, with significantly lower densities found in reduced vegetation complexity treatments compared with both enhanced and standard treatment plots. A. miniaceus abundance per plot was also significantly lower in reduced complexity than in standard and enhanced complexity plots. Synthesis and applications: Maintenance of understory vegetation complexity contributes to the preservation of spider host–cleptoparasite relationships in oil palm plantations. Understory structural complexity in these simplified agroecosystems therefore helps to support abundant spider populations, a functionally important taxon in agricultural landscapes. In addition, management for more structurally complex agricultural habitats can support more complex trophic interactions in tropical agroecosystems.     

Genetic estimates of immigration and emigration rates in relation to population density and forest patch area in <Emphasis Type="Italic">Peromyscus leucopus</Emphasis>

An emerging pattern is that population densities of generalist rodents are higher in small compared to large forest patches in fragmented landscapes. We used genetically based measures of migration between patches to test two dispersal-based hypotheses for this negative density-area relationship: (1) emigration rates from small patches should be relatively lower compared to large patches (“inhibited dispersal hypothesis”), or (2) immigration rates should be higher into small than large patches (“immigration hypothesis”). Neither hypothesis was supported using data on dispersal inferred from eight microsatellite loci for 12 populations of Peromyscus leucopus in six small (1.3–2.7 ha) and six large (8–150 ha) forest patches. Emigration rates were not lower from and immigration rates were not higher into small than large patches. In fact, contrary to both hypotheses, emigration rates were higher from populations of P. leucopus in small compared to large patches. Based on a combination of genetic and field data, we speculate that higher reproduction in smaller patches resulted in higher densities which led to higher emigration rates from those patches. Rates of reproduction (presumably driven by better habitat conditions in smaller patches), rather than dispersal, seems to drive density differences in forest patches. We conclude that smaller forest patches within an agricultural matrix act as a source of individuals, and that migration rates are fairly high among forest patches regardless of size.     

Cultivation of Non‐timber Forest Products Alters Understory Light Availability in a Humid Tropical Forest in Mexico1

The planting of non‐timber forest products (NTFPs) in the understory of tropical forests is promoted in many regions as a strategy to conserve forested lands and meet the economic needs of rural communities. While the forest canopy is left intact in most understory plantations, much of the midstory and understory vegetation is removed in order to increase light availability for cultivated species. We assessed the extent to which the removal of vegetation in understory plantations of Chamaedorea hooperiana Hodel (Arecaceae) alters understory light conditions. We also examined how any changes in light availability may be reflected by changes in the composition of canopy tree seedlings regenerating in understory plantations. We employed a blocked design consisting of four C. hooperiana plantation sites; each site was paired with an adjacent, unmanaged forest site. Hemispherical canopy photographs were taken and canopy tree seedlings were identified and measured within 12 3 × 2 m randomly placed plots in each site for a total of 96 plots (4 blocks × 2 sites × 12 plots). Plantation management did not affect canopy openness or direct light availability but understory plantations had a higher frequency of plots with greater total and diffuse light availability than unmanaged forest. Comparisons of canopy tree seedling composition between understory plantations and unmanaged forest sites were less conclusive but suggest that management practices have the potential to increase the proportion of shade‐intolerant species of tree seedlings establishing in plantations. Given the importance of advanced regeneration in gap‐phase forest dynamics, these changes may have implications for future patterns of succession in the areas of forest where NTFPs are cultivated.     

What saplings can tell us about forest expansion over natural grasslands

Questions: 1. Do the species composition, richness and diversity of sapling communities vary significantly in differently sized patches? 2. Do forest patches of different sizes differ in woody plant colonization patterns? Location: São Francisco de Paula, Rio Grande do Sul, Brazil, 29°28'S,50°13'W. Methods: Three woody vegetation types, differing in structural development (patch size) and recovering for 10 years from cattle and burning disturbances, were sampled on grassland. We analysed the composition and complexity of the woody sapling communities, through relative abundance, richness and diversity patterns. We also evaluated recruitment status (residents vs. colonizers) of species in communities occurring in different forest patch size classes. Results : 1. There is a compositional gradient in sapling communities strongly associated with forest patch area. 2. Richness and diversity are positively correlated to patch area, but only in poorly structured patches; large patches present richness and diversity values similar to small patches. 3. Resident to colonizer abundance ratio increases from nurse plants to large patches. The species number proportion between residents and colonizers is similar in small and large patches and did not differ between these patch types. 4. Large patches presented a high number of exclusive species, while nurse plants and small patches did not. Conclusions: Woody plant communities in Araucaria forest patches are associated with patch structure development. Richness and diversity patterns are linked to patch colonization patterns. Generalist species colonize the understorey of nurse plants and small patches; resident species cannot recruit many new individuals. In large patches, sapling recruitment by resident adults precludes the immigration of new species into the patches, limiting richness and diversity.     

Core and Satellite Species in Degraded Habitats: an Analysis Using Malagasy Tree Communities

Core-satellite theory predicts that, via the “rescue effect”, widespread, abundant species should have reduced risk of local extinctions. We test this hypothesis in southeastern Malagasy littoral forest using data on distribution and abundance of trees and woody understory vegetation in tropical forest fragments along a disturbance gradient. We partition the mortality risk into two kinds of extinction factors, separately operating at demographic (local) and landscape (regional) scales, contrary to core-satellite predictions, for both trees and woody understory vegetation, that the relative number of core (abundant) species declined significantly with increasing disturbance. In the least-degraded forest fragments there was a strong mode of core species, while in the moderately- and severely-degraded fragments the species distributions were essentially log-normal, lacking a substantial core mode. While the rescue effect mitigates one kind of extinction risk, namely local environmental and demographic stochasticity, it may not counterbalance widespread pervasive sources of mortality. The amount of internal forest fragmentation appears to have a much greater effect on species richness and diversity than either fragment size or shape.     

Light limitation creates patchy distribution of an invasive grass in eastern deciduous forests

Species interactions and their indirect effects on the availability and distribution of resources have been considered strong determinants of community structure in many different ecological systems. In deciduous forests, the presence of overstory trees and shrubs creates a shifting mosaic of resources for understory plants, with implications for their distribution and abundance. Determination of the ultimate resource constraints on understory vegetation may aid management of these systems that have become increasingly susceptible to invasions by non-native plants. Microstegium vimineum (Japanese grass) is an invasive annual grass that has spread rapidly throughout the understory of forests across the eastern United States since it was first observed in Tennessee in 1919. M. vimineum occurs as extensive, dense patches in the understory of eastern deciduous forests, yet these patches often exhibit sharp boundaries and distinct gaps in cover. One example of this distributional pattern was observed relative to the native midstory tree Asimina triloba (pawpaw), whereby dense M. vimineum cover stopped abruptly at the drip line of the A. triloba patch and was absent beneath the A. triloba canopy. We conducted field and greenhouse experiments to test several hypotheses regarding the causes of this observed pattern of M. vimineum distribution, including allelopathy, seed dispersal, light limitations, and soil moisture, texture, and nutrient content. We concluded that light reduction by the A. triloba canopy was the environmental constraint that prevented establishment of M. vimineum beneath this tree. Whereas overstory tree canopy apparently facilitates the establishment of this shade-tolerant grass, the interaction of overstory canopy with midstory canopy interferes with M. vimineum by reducing the availability of sunflecks at the ground layer. It is likely that other midstory species influence the distribution and abundance of other herb-layer species, with implications for management of understory invasive plant species.     

Ecosystem Impacts of a Range Expanding Forest Defoliator at the Forest-Tundra Ecotone

Insect outbreaks in northern-boreal forests are expected to intensify owing to climate warming, but our understanding of direct and cascading impacts of insect outbreaks on forest ecosystem functioning is deficient. The duration and severity of outbreaks by geometrid moths in northern Fennoscandian mountain birch forests have been shown to be accentuated by a recent climate-mediated range expansion, in particular of winter moth (Operophtera brumata). Here, we assess the effect of moth outbreak severity, quantified from satellite-based defoliation maps, on the state of understory vegetation and the abundance of key vertebrate herbivores in mountain birch forest in northern Norway. We show that the most recent moth outbreak caused a regional-scale state change to the understory vegetation, mainly due to a shift in dominance from the allelopathic and unpalatable dwarf-shrub Empetrum nigrum to the productive and palatable grass Avenella flexuosa. Both these central understory plant species responded significantly and nonlinearly to increasing outbreak severity. We further provide evidence that the effects of the outbreak on understory vegetation cascaded to cause strong but opposite impacts on the abundance of the two most common herbivore groups. Rodents increased with defoliation, largely mirroring the increase in A. flexuosa, whereas ungulate abundance instead showed a decreasing trend. Our analyses also suggest that the response of understory vegetation to defoliation may depend on the initial state of the forest, with poorer forest types potentially allowing stronger responses to defoliation.     

Effect of prey availability on the abundance of White‐breasted Wood‐Wrens,insectivorous birds of tropical lowland forests

Some understory insectivorous birds manage to persist in tropical forest fragments despite significant habitat loss and forest fragmentation. Their persistence has been related to arthropod biomass. In addition, forest structure has been used as a proxy to estimate prey availability for understory birds and for calculating prey abundance. We used arthropod biomass and forest structural variables (leaf area index [LAI] and aerial leaf litter biomass) to explain the abundance of White‐breasted Wood‐Wrens (Henicorhina leucosticta), tropical understory insectivorous birds, in six forests in the Caribbean lowlands of Costa Rica. To estimate bird abundance, we performed point counts (100‐m radius) in two old‐growth forests, two second‐growth forests, and two selectively logged forests. Arthropod abundance was the best predictor of wood‐wren abundance (wi = 0.75). Wood‐wren abundance increased as the number of arthropods increased, and the estimated range of bird abundance obtained from the model varied from 0.51 (0.28 – 0.93 [95%CI]) to 3.70 (1.68 – 5.20 [95%CI]) within sites. LAI was positively correlated to prey abundance (P = 0.01), and explained part of the variation in wood‐wren abundance. In forests with high LAI, arthropods have more aerial leaf litter as potential habitat so more potential prey are available for wood‐wrens. Forests with a greater abundance of aerial leaf litter arthropods were more likely to sustain higher densities of wood‐wrens in a fragmented tropical landscape.     

Dryland management regimes alter forest habitats and understory arthropod communities

Dryland forests, those characterised as having low precipitation and soil nutrients, account for over a quarter of forests globally. Increasing their productivity often relies on irrigation and fertilisation, but the impacts on the wider habitat are largely unknown. Understory invertebrates, in particular, play key roles in forest systems (e.g. nutrient cycling), but their responses to dryland forest management practices are untested. We investigated the impacts of irrigation, fertilisation and a combination of both on soil chemistry, understory vegetation, tree growth and understory arthropod communities in a Eucalyptus plantation to establish linkages between dryland management and ecosystem responses. Fertilisation increased all soil nutrients (N, NO₃N, P and K) with similar effects on the chemical composition of understory grasses. Fertilisation also caused declines in foliar silicon concentrations, an important herbivore defence in grasses. Irrigation increased growth of both understory plants (+90%) and trees (+68%). Irrigation increased the abundance of ground‐dwelling arthropods by over 480% relative to control plots, but depressed higher level taxon arthropod diversity by 15%, declining by a further 7% (?22%) in combined treatment plots. Irrigation also caused a surge in the abundance of Collembola (+1300%) and Isopoda (+323%). Fertilisation drove increases in the abundance of Isopoda (+196%) and Diptera (+63%), whereas fertilisation combined with irrigation increased populations of Thysanoptera (+166%) and Acarina (+328%). Airborne arthropods were less affected, but fertilisation increased the abundance of Apocrita (+95%) and depressed populations of Thysanoptera (?77%). Diptera abundance was positively related to understory vegetation growth, whereas the abundance of other groups (Collembola, Isopoda, Thysanoptera and Acarina) correlated positively with tree growth. We proposed that the large increases in populations of key detritivores, Collembola and Isopoda, were linked to increased leaf litter from enhanced tree growth in irrigated and combined treatment plots. Our findings suggest that dryland management can increase both plant productivity and abundance of arthropods, but cause arthropod diversity at the higher taxon level to decline overall.     

Effects of different Rhododendron control methods in eastern beech (Fagus orientalis Lipsky) ecosystems in the western Black Sea region of Turkey

Intensive weed control and plot preparation practices have become a critical and integral part of productive beech forest management in Turkey’s coastal Black Sea region (BSR). This study was conducted in an eastern beech forest of 100+ year old in the BSR to evaluate ecosystem effects of three different experimental Rhododendron ponticum understory control methods with a randomised block design, including manual grubbing, foliar and cut stump spraying with imazapyr (Arsenal) and foliar and cut stump spraying with triclopyr (Garlon). Untreated vegetation plots served as controls. Evaluation of these treatments included their effects on understory and forest floor biomass and nutrients (C, N, P, S, K, Ca and Mg) and effects on soils, including bulk density, pH, soil nutrients (C, N, P and S), exchangeable cations (K, Ca and Mg) and soil cation exchange capacity (CEC). Grubbing and imazapyr treatments had greatly reduced the amount of understory biomass 5 years after application (P = 0.002). Triclopyr treatment also had a major effect on understory vegetation control, but by 5 years later, about 10% of the rhododendron originally present on these plots had gradually re‐sprouted and partially covered the plots. Five years after woody vegetation control treatments, at the 0‐ to 20‐cm depth, treatments did not appear to affect soil bulk density, pH and CEC. For the upper 20‐cm soil depth, the exchangeable soil K concentration at the 10‐ to 20‐cm depth on triclopyr‐treated plots was 33% higher than on grubbing plots, and it was twice that of imazapyr application plots. Imazapyr plots had almost 11 times more dead organic matter on the forest floor than there was on grubbing plots. Forest floor C concentrations on imazapyr plots were 26 and 14% greater than those on grubbing and triclopyr plots, respectively. Total ecosystem (forest floor + understory + soil exchangeable) Ca content was 50% higher on imazapyr plots than that on triclopyr plots, while the ecosystem K pool on imazapyr treatment plots was 27% lower than that on triclopyr plots. Herbicides can be used as an alternative for achieving some forest management objectives when other vegetation control methods are not feasible or economical. It is recommended that vegetation control not be used on steep slopes because of greater risk of soil erosion. There may be benefits in encouraging slash disposal by fire after imazapyr treatments, thus removing recalcitrant understory residues left on the forest floor and releasing the essential nutrients within them.     

Small mammal abundance in Mediterranean post-fire habitats: a role for predators?

We studied patterns of small mammal abundance and species richness in post-fire habitats by sampling 33 plots (225 m² each) representing different stages of vegetation recovery after fire. Small mammal abundance was estimated by live trapping during early spring 1999 and vegetation structure was sampled by visual estimation at the same plots. Recently–burnt areas were characterised by shrubby and herbaceous vegetation with low structural variability, and unburnt areas were characterised by well developed forest cover with high structural complexity. Small mammal abundance and species richness decreased with time elapsed since the last fire (from 5 to at least 50 years), and these differences were associated to the decreasing cover of short shrubs as the post-fire succession of plant communities advanced. However, relationships between vegetation structure and small mammals differed among areas burned in different times, with weak or negative relationship in recently burnt areas and positive and stronger relationship in unburnt areas. Furthermore, the abundance of small mammals was larger than expected from vegetation structure in plots burned recently whereas the contrary pattern was found in unburned areas. We hypothesised that the pattern observed could be related to the responses of small mammal predators to changes in vegetation and landscape structure promoted by fire. Fire-related fragmentation could have promoted the isolation of forest predators (owls and carnivores) in unburned forest patches, a fact that could have produced a higher predation pressure for small mammals. Conversely, small mammal populations would have been enhanced in early post-fire stages by lower predator numbers combined with better predator protection in areas covered by resprouting woody vegetation.     

Effects of microhabitat on palm seed predation in two forest fragments in southeast Brazil

The establishment of plants depends crucially on where seeds are deposited in the environment. Some authors suggest that in forest understory seed predation is lower than in gaps, and higher than at the forest edge. However, most studies have been carried out in large forest patches and very little is known about the effects of microhabitat conditions on seed predation in forest fragments. We evaluated the effects of three microhabitats (gaps, forest edge, and understory) on seed predation of two palm species (Euterpe edulis and Syagrus romanzoffiana) in two semi-deciduous forest fragments (230 and 2100 ha) in southeast Brazil. Our objective was to test two hypotheses: (1) Low rodent abundance in small fragments as a result of meso-predator action levels leads to lower seed predation in small fragments. (2) Most mammal species in small fragments are generalists with respect to diet and habitat, so that seed predation is similar in different microhabitats (gaps, forest edge and understory) in the small fragment, but not in the larger one. The study community of small fragments is usually composed of generalist species (in diet and habitat aspects), so we expected the same rate of seed predation among microhabitats (gaps, forest edge and understory) in the tested smaller fragment. The experiment was carried out in the dry season (for E. edulis) and in the wet season (for S. romanzoffiana) in 1999. We conclude that post-dispersal seed predation in forest fragments can be directly connected with mammal communities, reflecting their historical and ecological aspects.     

Influence of N2-fixing Trifolium on plant species composition and biomass production in alpine tundra

Alpine Trifolium species have high rates of symbiotic N₂-fixation which may influence the abundance and growth of plant species growing near them. The potential for facilitative effects on plant abundance and growth in dry meadow alpine tundra of Niwot Ridge, Colo., characterized by low resource availability, was investigated by measuring soil N, aboveground biomass production, and plant species composition in patches of Trifolium dasyphyllum and surrounding tundra. Extractable inorganic N was more than twofold greater and extractable P was 27% lower in Trifolium patches than in surrounding tundra. Aboveground production was twofold greater in Trifolium patches than in surrounding tundra. However, the difference was largely due to the production of T. dasyphyllum relative to the non-Trifolium component of biomass, which was not different between the Trifolium patches and surrounding tundra. In the Trifolium patches, the proportion of graminoid biomass was lower while the proportion of forb biomass was higher relative to surrounding tundra. Although the abundance of some species was positively associated with the presence of Trifolium, other species were less abundant, possibly due to increased competition for P and differential abilities of alpine species to respond to increased N availability. Trifolium may exert both facilitative and inhibitive effects on dry meadow alpine species and, in the process, substantially influence the spatial heterogeneity in community structure and primary production. Received: 14 October 1997 / Accepted: 2 February 1998     

Temporal variation in the woody understory of an old-growth Fagus-Acer forest and implications for overstory recruitment

Abstract. Changes in woody vegetation were examined over eight years, using a 1.05-ha permanent plot in which the location of every shrub and tree > 1m height was mapped. There was little change in the overstory vegetation, as expected for an old-growth forest. Much greater change occurred in the understory, primarily related to a 40 % increase in density. Differences occurred among species in the under-story, as Acer saccharum and Prunus serotina increased and Fraxinus americana and Fagus grandifolia decreased. Canopy gap dynamics are implicated in differences among species in the establishment and growth of individuals in the understory and their recruitment into the overstory. It is concluded that because understory is temporally variable, overstory recruitment from the understory may take different courses at different times in the same forest.     

Patch size of forest openings and arthropod populations

Summary Five sizes of canopy openings (0.016 ha to 10 ha) were established in the Southern Appalachian Mountains in early 1982 to examine the initial patterns of plant and arthropod establishment across a size range of forest disturbances. Vegetation standing crop after the first growing season was considerably higher in large than small openings in apparent response to greater resource release (e.g., sunlight) in larger openings. Woody stump and root sprouts were the dominant mode of revegetation in each patch size. Forest dominants such as Quercus rubra, Q. prinus and Carya spp. were less important as sprouters in openings than several minor forest components (e.g., Robinia pseudo-acacia, Acer rubrum, Halesia carolina and Cornus florida). Arthropod abundance and community composition varied across the size range of forest openings. Arthropods from the surrounding forest readily utilized the smallest canopy openings (0.016 ha). All feeding guilds were well represented in these small openings and herbivore biomass and load (mg of herbivores/g of foliage) were much higher than in larger patches. In contrast, arthropod abundance and species richness were significantly lower in mid-size than smaller patches. The relatively sparse cover and high sunlight in mid-size openings may have promoted surface heat buildups or soil surface/litter moisture deficits which restricted arthropod entry from the surrounding forest. Arthropod abundance and species richness were higher in large than mid-size patches. The greater vegetation cover in larger openings may have minimized the deleterious effects on arthropod populations. However, the absence of population increases among these arthropod species maintained herbivore loads at very low levels in large patches. Our results suggest that arthropod abundance and diversity in sprout-dominated forest openings are highly dependent on the extent of environmental differences between patch and surrounding forest.