Does understory vegetation reflect the history of fluvial disturbance in a riparian forest?

We have examined the relationship between the history of fluvial disturbance and understory vegetation in a riparian forest. The study site was divided into three sites, by use of aerial photographs and topographical maps, with different histories of fluvial disturbance: (1) Fagus-type on land that has not been flooded for the last 39 years, at least; (2) Populus-type on land that has not been flooded since debris flow occurred 34 years ago; and (3) Salix-type on land that has been flooded periodically from an abandoned channel since debris flow occurred 34 years ago. Species richness in the Salix-type was significantly higher than in the other types. Detrended correspondence analysis revealed obvious floristic differences among the three canopy types. Canonical correspondence analysis showed that herbaceous species were mainly found on lower plots with high moss cover, implying that moss layers may capture seeds transported by the stream. Tall herbs occurred in less shaded plots and on higher plots, suggesting that their rapid growth prevented the occurrence of other species. Fagus-type was dominated by species with ingested fruits which depended on animals for their dispersal. Populus and Salix-types were dominated by species with wind dispersal or no dispersal mechanism, which depended on physical phenomena for dispersal. Attributes of current understory vegetation were connected with historical events, suggesting that riparian vegetation reflects the history of fluvial disturbance.     

Changes in forest understorey vegetation in Norway related to long‐term soil acidification and climatic change

Question: Does the understorey vegetation of Norwegian boreal forests change in relation to broad‐scale, long‐term changes? Location: Norway. Methods: Permanently marked 1‐m² vegetation plots from 17 monitoring reference areas in forests dominated by Picea abies (11 areas, 620 plots) and Betula spp. (six areas, 300 plots) were analysed twice, at the start in 1988–1997 and 5 yr later (1993–2002). Species subplot frequency data were analysed separately for each area by univariate and multivariate statistical methods; 5‐yr changes in single species abundances, species number per plot and species composition were tested. Results: Two distinct patterns of change were found: 1. Abundance of several vascular plant species decreased in SE Norwegian Picea forests, most noticeably of species with a preference for richer soils, such as Oxalis acetosella. 2. Abundance of many bryophyte species as well as bryophyte species number per plot increased in forests of both types over most of Norway. Conclusions: The pattern of vascular plant changes is probably a time‐delayed response of long‐lived, mainly clonal, populations to acidified soils resulting from deposition of long‐distance airborne pollutants. The pattern bryophyte changes, with reference to the close link between climatic conditions for growth and abundance changes for Hylocomium splendens established in previous demographic studies, is related to climatic conditions favourable for bryophyte growth. We conclude that many forest understorey plants are sensitive indicators of environmental change, and that the concept used for intensive monitoring of Norwegian forests enables early detection of changes in vegetation brought about by broad‐scale, regional, impact factors.     

Disturbance effects on herbaceous layer vegetation and soil nutrients in Populus forests of northern lower Michigan

Abstract. Recent disturbance models have identified changes in resource availability as factors that control plant community response. Soil nutrient resources typically are assumed to change following forest disturbance, usually with nutrient availablity increasing initially and subsequently decreasing through later stages of succession. We examined the effects of disturbance (clearcut harvesting with a brief recovery period) on soil organic matter, pH and extractable soil nutrients in successional aspen forests of northern lower Michigan to determine relationships of these variables to changes in herbaceous layer vegetation. Two site types were identified: drymesic (glacial outwash sands, low in organic matter) and mesic (calcareous clay till, high in organic matter). Extractable nutrient concentrations were 1.5 to 3 times higher in the A₁ horizon of mesic sites than those of dry-mesic sites. Soil pH and cations increased after disturbance on mesic sites, but not on dry-mesic sites. Patterns of change with disturbance were less pronounced in lower horizons on both site types. Herblayer species diversity increased after disturbance on mesic sites, but with decreases in the importance of shade-tolerant tree species and Maianthemum canadense. Species characteristic of open habitats (e.g. Pteridium aquilinum, Rubus spp., Fragaria virginiana, and Diervilla lonicera), increased in importance. Soil factors, species composition and diversity on dry-mesic sites changed little after disturbance, with Pteridium aquilinum and ericaceous species remaining dominant in both mature (55–82 yr) and disturbed (≤ 15 yr) stands. These results suggest that soil nutrient resources do not always change through secondary succession and that patterns of change can be distinctly site-dependent. Disturbance response patterns in the herbaceous layer of these aspen forests are also site-dependent.     

Structure and spatial patterns of trees in old-growth northern hardwood and mixed forests of northern Maine

Stand structure including spatial patterns was studied in northern hardwood and mixed forest types in the 2000-ha old-growth Big Reed Forest Reserve in northern Maine using complete stem mapping, dendrochronology, and spatial analyses on 0.5 plots. The inclusion of saplings, dead wood, age distributions, spatial pattern, and interactions provided some idea of underlying processes and temporal change. Structural characteristics were most comparable to spruce-northern hardwood forests of northern New England and New York, and most characteristics matched expected patterns for old-growth forests of the region.Results indicated smaller maximum-tree sizes, lower basal areas (26–34 ) and downed-wood volumes (29–64), higher densities (475–649), but similar species longevities compared to other mesic old-growth forests further south and in the Lake States. The stands were dominated by very shade-tolerant tree species, including Fagus grandifolia Ehrh., Acer saccharum Marsh.,Picea rubens Sarg. and Abies balsamea(L.) Mill, with each species found in many crown positions and age and size classes. The sapling layer was dominated by Fagus grandifolia followed by Picea rubens. Most species had reverse-J shaped diameter distributions, but age distributions were indicative of synchronous, episodic recruitment. In most plots, Acer saccharum diameter distributions were skewed towards the mid-larger size classes. Lack of young and small Acer saccharum stems suggested change in forest composition towards Fagus grandifolia dominance. Most species formed small-scale clusters (15 ) perhaps in response to small gap disturbances. Snags were the dominant dead wood type and were randomly to regularly distributed in most plots. Logfall directions were unrelated to hurricane paths. Recent small-scale disturbance events and topographic position appear to be important in explaining current structure and dynamics of the hardwood and mixed forests of Big Reed Forest Reserve in northern Maine. The continued effects of beech bark disease had a greater effect on hardwood plots, whereas a recent spruce budworm outbreak had a greater effect on plots with higher conifer density.The dominance of very shade tolerant tree species in small-scale clusters, and randomly distributed snags rather than clustered uproots were indicative of the prevalence of small scale gap disturbance regimes in the hardwood and mixed forests of Big Reed Forest Reserve in northern Maine. Varying topographic position may allow for slight changes in disturbance regime leading to consequent variation in structure and dynamics. H1, a more open plot on upper exposed slopes, had distinctly different characteristics such as lower live and dead tree and sapling densities than the other plots, but more uprooted trees and Acer saccharum saplings. Such small scale gap disturbance regimes operating on an episodic basis, and effects of slight variations in this regime on stand composition and structure have significant implications for silvicultural interventions and management of these forest types.     

Overstory species composition of naturally regenerated clearcuts in an ecological classification framework

Multifactor ecological classification systems are being developed for many regions. An element of these systems not yet well understood is how disturbances, clearcutting in this instance, may alter the vegetative component of the classification units at the stand and landscape levels. We sampled 1,096 plots in 21–35 year old naturally regenerated clearcuts on the Hoosier National Forest (HNF) in south-central Indiana, USA. We examined overstory species composition of clearcut plots in comparison to reference plots (80+ years old), both within and among six Ecological Landtype Phases (ELTPs) of two ecological sections using non-metric multidimensional scaling and non-metric multi-response permutation procedures. Clearcutting drastically changed species composition in comparison to reference plots within ELTPs ranging from mixed oak-dominated ridges and slopes to bottomland, cove hardwood communities; Quercus species on ridges and slopes were replaced by Liriodendron tulipifera L. and, to a lesser degree, Prunus serotina Ehrh. and Acer rubrum L., in ELTPs of both sections. Contrasts of overstory species composition of reference plots exhibited differences among ELTPs, but clearcut plots showed mixed results and indicated very similar species composition across all ELTPs. Autogenic factors are likely the main drivers of overstory composition of clearcut sites. Species composition of ELTPs will continue to develop in response to autogenic and allogenic factors over time, and differences among ELTPs may emerge in later stages of stand development as the effects of allogenic factors accumulate. It is expected that L. tulipifera, a long-lived species, will be a dominant species in terms of basal area and density of all ELTPs in mature stands. Classification systems not designed to deal with changes related to disturbance and a failure to predict successional pathways after disturbance may limit their usefulness as a management tool in terms of overstory vegetation. For ecological classification systems to be fully effective, we must better understand the role of disturbance in ecosystem function at many different scales and integrate that knowledge into our decision-making and planning regimes to establish realistic and attainable objectives at multiple scales.     

Diversity of vascular plants on Ssese islands in Lake Victoria, central Uganda

Diversity and distribution of trees [≥5 cm diameter at breast height (dbh)], shrubs and herbs was assessed in thirty 0.05‐ha (10 × 50 m) plots of a tropical high forest in the Ssese islands of Lake Victoria, central Uganda. The aim was to determine the floristic richness and composition of the forests. We recorded 179 species belonging to 70 families and 146 genera. Of these, nine families had five species or more. Rubiaceae was the richest with fourteen species followed by Euphorbiaceae (thirteen), Apocynaceae (ten) and Moraceae (nine). The majority of the families (35) were represented by one species each. Fifty‐eight herbaceous species, 39 lianas, ten shrubs and 72 species of trees were recorded. The commonest species recorded in the forest included: Uapaca guineensis Mull. Arg., Tabernaemontana pachysiphon Stapf., and Aframomum luteoalbum (K Schum.) K. Schum. Among the rare species encountered were Ficus densistipulata De Willd., Englerophytum oblanceolatum (S. Moore) Pennington, and Afromomum zambeziacum (Bak.) K. Schum. The present study has shown that the Ssese islands are floristically rich in species and compare well with other mainland forests. Species richness, rarity and uniqueness of habitats can be considered as approaches in the prioritization of conservation sites within the fragmented forests of Ssese islands.     

Little evidence for niche partitioning among ectomycorrhizal fungi on spruce seedlings planted in decayed wood versus mineral soil microsites

Ectomycorrhizal fungal (EMF) communities vary among microhabitats, supporting a dominant role for deterministic processes in EMF community assemblage. EMF communities also differ between forest and clearcut environments, responding to this disturbance in a directional manner over time by returning to the species composition of the original forest. Accordingly, we examined EMF community composition on roots of spruce seedlings planted in three different microhabitats in forest and clearcut plots: decayed wood, mineral soil adjacent to downed wood, or control mineral soil, to determine the effect of retained downed wood on EMF communities over the medium and long term. If downed and decayed wood provide refuge habitat distinct from that of mineral soil, we would expect EMF communities on seedlings in woody habitats in clearcuts to be similar to those on seedlings planted in the adjacent forest. As expected, we found EMF species richness to be higher in forests than clearcuts (P ≤ 0.01), even though soil nutrient status did not differ greatly between the two plot types (P ≥ 0.05). Communities on forest seedlings were dominated by Tylospora spp., whereas those in clearcuts were dominated by Amphinema byssoides and Thelephora terrestris. Surprisingly, while substrate conditions varied among microsites (P ≤ 0.03), especially between decayed wood and mineral soil, EMF communities were not distinctly different among microhabitats. Our data suggest that niche partitioning by substrate does not occur among EMF species on very young seedlings in high elevation spruce-fir forests. Further, dispersal limitations shape EMF community assembly in clearcuts in these forests.     

Ungulate browsers promote herbaceous layer diversity in logged temperate forests

Ungulates are leading drivers of plant communities worldwide, with impacts linked to animal density, disturbance and vegetation structure, and site productivity. Many ecosystems have more than one ungulate species; however, few studies have specifically examined the combined effects of two or more species on plant communities. We examined the extent to which two ungulate browsers (moose [Alces americanus]) and white‐tailed deer [Odocoileus virginianus]) have additive (compounding) or compensatory (opposing) effects on herbaceous layer composition and diversity, 5–6 years after timber harvest in Massachusetts, USA. We established three combinations of ungulates using two types of fenced exclosures – none (full exclosure), deer (partial exclosure), and deer + moose (control) in six replicated blocks. Species composition diverged among browser treatments, and changes were generally additive. Plant assemblages characteristic of closed canopy forests were less abundant and assemblages characteristic of open/disturbed habitats were more abundant in deer + moose plots compared with ungulate excluded areas. Browsing by deer + moose resulted in greater herbaceous species richness at the plot scale (169 m²) and greater woody species richness at the subplot scale (1 m²) than ungulate exclusion and deer alone. Browsing by deer + moose resulted in strong changes to the composition, structure, and diversity of forest herbaceous layers, relative to areas free of ungulates and areas browed by white‐tailed deer alone. Our results provide evidence that moderate browsing in forest openings can promote both herbaceous and woody plant diversity. These results are consistent with the classic grazing‐species richness curve, but have rarely been documented in forests.     

Post-fire regeneration in a Mediterranean pine forest with historically low fire frequency

Species of Mediterranean vegetation are known to regenerate directly after fire. The phenomenon of autosuccession (direct regeneration) has been found to be often combined with an increase of species richness during the first years after fire due to the high abundance of short-lived herbaceous plants facilitated by plentiful nutrients and light. The high degree of vegetation resilience, which is expressed in terms of autosuccession, has been explained by the selective pressure of fire in historic times. According to existing palaeoecological data, however, the Pinus halepensis forests in the Ricote Mountains (Province of Murcia, SE Spain) did not experience substantial fire impact before the presence of man nor are they especially fire-prone today. Therefore, we studied post-fire regeneration to find out if direct succession is present or if species from pre-fire vegetation are absent during the post-fire regeneration stages. Patterns of succession were deduced from observations made in sample plots on sites of a known regeneration age as well as in adjacent unburnt areas. The results of the vegetation analyses, including a Detrended Correspondence Analysis, indicate that Pinus halepensis forest regeneration after fire resembles autosuccession. As regards the presence of woody species, there is a high percentage similarity on north (83%) and south (70%) facing slopes during the first year after fire vs. reference areas which is due, for example, to direct regeneration of the resprouting Quercus coccifera or seeders like Pinus halepensis or Fumana laevipes. However, if herbaceous species are included in the comparison, the similarity on north-facing sites decreases (to 53%) with the presence of additional species, mainly ruderals like Anagallis arvensis or Reseda phyteuma, and even woody species on the burnt plots. This effect indicates “enhanced autosuccession”, which was not found on south-facing sites where overall species richness was very high irrespective of the impact of fire. Locally we found limited regeneration of some species, for example Pinus halepensis at high altitudes (1000 m), even 22 years after fire. As we assume that historical fires did not play an important role in the area and direct succession is present nevertheless, our results support the theory that autosuccession is not a process restricted to fire-prone areas. Fire has been only one of several selective forces since human settlement that probably led to a set of species pre-adapted against recurrent disturbance.     

Strong influence of long-distance edge effect on herb-layer vegetation in forest fragments in an agricultural landscape

In agricultural landscapes in central Europe, species richness of the herbaceous plant community may be compromised by processes associated with forest fragmentation, habitat loss, and management practices. We examined variability in species richness and composition of the herbaceous layer in 229 plots located in 23 forest fragments (0.1 to 255 ha), in a representative upland agricultural landscape in central Bohemia, in relation to the most important site environmental factors, edge effects, and site history. The influence of environmental factors on the composition of vegetation in the herb layer was evaluated using generalized additive models, which enabled us to analyze highly non-linear and non-monotonic relationships. Total species richness and number of red-listed and ancient forest species were significantly influenced by type of forest vegetation, light quality, soil pH, slope aspect, and distance from the forest edge. Implications of the significant explanatory variables corresponded well to previous findings, with the exception of distance from the forest edge, for which we found a positive relationship with species richness for distances up to 200 m toward the forest interior. Plant species with low colonization ability occupied plots with increasing frequency from edge to forest interior, while fast-colonizing species showed the opposite trend. Apart from the edge effect, forest continuity should be considered for its important contribution to the richness of ancient forest and red-listed species, whereas the effect of forest fragment size appeared to be generally weak. These results do not negate the importance of large forest fragments for the maintenance of herb layer species richness, but specifically emphasize the essential contribution of the core habitats of these forests. In summary, we showed that the negative effects of habitat fragmentation on the richness of ancient forest and red-listed species and on herb layer species in total can be largely attributed to either the edge effect itself or to aggregate effects of forest edge and forest continuity.     

“Ordinary biodiversity” in western Burkina Faso (West Africa): what vegetation do the state forests conserve?

The role of state forests in the conservation of the savannah environment of western Burkina Faso (West Africa) was assessed by considering the customary and fundamental biological components of plant diversity. The focus is on the “ordinary biodiversity” which constitutes the current environment of human populations and which generates their resources. Overall floristic composition, α, β, γ diversities and species accumulation curves of landscape units were compared inside and outside protected areas. Common species were identified according to their frequencies in landscape units and their local abundance. The occurrences of functional (life form, dispersion, etc.) and human-valued (current uses, potential threats, etc.) traits were compared. Current land uses outside protected areas have an unequal effect on the various components of plant diversity. Species richnesses and specific structures were all the more altered for woody species as agricultural pressure is high; conversely, they were maintained for herbaceous communities for which the proportion of wide-ranging species increased. The preservation of useful plant species on cultivated lands was obvious for woody species with edible fruits but less tangible for timber and service wood. It was deficient for species that furnish other non-wood products (e.g. leaves used as food or fodder), the conservation of which may be jeopardized where anthropic pressure is high. Given current land uses and dynamics, maintaining conservation areas in land use planning is crucial in order to preserve ordinary biodiversity. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Habitat availability in forests and clearcuts for saproxylic beetles associated with aspen

To maintain biodiversity in managed, boreal forest in Scandinavia, aspen trees (Populus tremula) are often retained at clearcutting. In this study, the habitat availability for beetles associated with aspen coarse woody debris (CWD) was predicted for forests and clearcuts with a model of CWD dynamics. Habitat requirements of eight beetle species (Agathidium bicolor, Cerylon ferrugineum, Cyphaea curtula, Endomychus coccineus, Homalota plana, Mycetophagus fulvicollis, Ptilinus fuscus and Xylotrechus rusticus) were obtained from their occurrence patterns in relation to characteristics of CWD objects in forest and on clearcuts in a study landscape in central Sweden. Three species were more frequent in forest and three at clearcuts. Five species increased with increasing girth of the CWD. Three were more frequent on standing CWD, and two on lying CWD. From the same study area, we also obtained field data on the recruitment of CWD (i.e., tree mortality) and amounts of different types of CWD. Annual tree mortality of aspen was higher for recent clearcuts (6.3%) compared with older clearcuts (1.1%). For all species, the habitat availability was higher on clearcuts, because enhanced tree mortality increased the amount of recently dead CWD. As a conclusion, green-tree retention of aspen is a conservation effort that is beneficial for species associated with aspen CWD.     

Stand dynamics of a yellow-poplar (Liriodendron tulipifera L.) forest in the Appalachian Mountains, Virginia, USA

Yellow-poplar (Liriodendron tulipifera L.) is a large, rapidly growing, shade-intolerant tree species common after disturbances on moist sites in the Appalachian Mountains. The species is typically scattered throughout old-growth mesophytic forests, where periodic gap formation creates conditions favorable for yellow-poplar establishment and growth. On abandoned agricultural fields, however, it is common for nearly monospecific forests of yellow-poplar to develop.This study examines stand dynamics of a yellow-poplar forest in western Virginia, USA that was established on agricultural fields abandoned in the late 1940s. Increment cores were collected from yellow-poplar trees growing on exposed ridgetops and in a more sheltered hollow. Tree-ring data show that the forest is even-aged. Tree establishment began about 5 years earlier on the ridgetops than in the hollow. Major ice storms disturbed the forest in 1978 and 1994, with two separate events in 1994. Ice storms disturb forests by depositing heavy loads of freezing rain on trees, breaking or uprooting them. The dendroecological data collected for this study provide little support for the hypothesis that ice storm disturbance promotes the establishment of new yellow-poplar cohorts. However, the data show that radial growth of some trees increased after ice storm disturbance, a pattern that reflects the increased availability of light following disturbance. Radial growth declined in some other trees as a consequence of severe injury during the storms.Radial growth responses following the 1978 ice storm were stronger on the ridgetops than in the hollow, suggesting that tree damage was more severe on the higher, more exposed sites. Growth responses were relatively mild following the storms of 1994, and did not exhibit pronounced topographic variations.     

A multi-scale assessment of the occurrence of exotic plants on the Olympic Peninsula,Washington

Abstract. Exotic plants were surveyed in 208 plots within the Dungeness and Hoh river watersheds on the Olympic Peninsula, Washington, USA. Landscape patch types included uplands (clearcuts, young and mature forests) and riparian zones (cobble bars, shrub patches, riparian forests, and alder flats). Patterns of exotic plant invasions were assessed between watersheds, between riparian and upland areas, among patch types, and within clearcuts. 52 exotic plant species were encountered, accounting for 23% of the flora in each watershed. In both watersheds, exotic species richness was approximately 33% greater in riparian zones than in uplands, and mean number and cover of exotic species were > 50% greater in riparian zones than in uplands. Among landscape patch types, richness and mean number and cover of exotics was highest in young riparian patches, intermediate in clearcuts and riparian forests, and lowest in young and mature forests. The exception to this was Hoh alder flats, which had the highest mean cover of exotic plants. Cover of exotic plants peaked in uplands 3 to 7 yr after clearcutting, then decreased with increased canopy closure. Disturbance type and time since disturbance were major factors influencing invasibility. Landscape patch size, position within watershed (distance from patch to human population centers, major highway, or river mouth), and environmental variables (slope, aspect, and elevation) were not important indicators of landscape patch invasibility. Riparian zones facilitated movement of exotic plants through landscapes, but did not appear to act as sources of exotic plants for undisturbed upland areas.     

Productivity and Subordinate Species Response to Dominant Grass Species and Seed Source during Restoration

Grasses can be important regulators of species diversity and ecosystem processes in prairie systems. Although C₄ grasses are usually assumed to be ecologically similar because they are in the same functional group, there may be important differences among species or between seed sources that could impact restorations. I tested whether C₄ grass species identity, seed source, or grass species richness scales to influence aboveground net primary productivity (ANPP), resistance to weed invasion, or establishment of subordinate prairie species during restoration. Plots in western Iowa, United States, were planted with equal‐sized transplants of one of five common grass species (Panicum virgatum L., Sorghastrum nutans (L.) Nash, Andropogon gerardii Vitman, Schizachyrium scoparium (Michx.) Nash, and Bouteloua curtipendula (Michx.) Torrey) either from local seed or from cultivar seed sources. These plots were compared to plots containing all five species in mixture and to nonplanted plots. Differences in ANPP were found among species but not between cultivars and noncultivars or between monocultures and mixtures. Panicum virgatum, S. nutans, and S. scoparium were more productive than A. gerardii and B. curtipendula. Weed invasion was much higher when plots were not planted with grasses. Schizachyrium scoparium allowed greater establishment of subordinant prairie species than all other focal grass species. There were two separate mechanisms by which grasses suppressed prairie species establishment either (1) by growing tall and capturing light or (2) by quickly filling in bare space by spreading horizontally through rhizome growth in short species. These results suggest that high ANPP can be found with noncultivar plantings during the first 2 years after planting and that subordinate species establishment is most likely when shorter bunchgrasses such as S. scoparium are dominant.     

Impact of pocket gopher disturbance on plant species diversity in a shortgrass prairie community

Summary We examined the impact of pocket gopher disturbances on the dynamics of a shortgrass prairie community. Through their burrowing activity, pocket gophers (Thomomys bottae) cast up mounds of soil which both kill existing vegetation and create sites for colonization by competitively-inferior plant species. Three major patterns emerge from these disturbances: First, we show that 10 of the most common herbaceous perennial dicots benefit from pocket gopher disturbance; that is, a greater proportion of seedlings are found in the open space created by pocket gopher disturbance than would be expected based on the availability of disturbed habitat. Additionally, these seedlings exhibited higher growth rates than adjacent seedlings of the same species growing in undisturbed habitat. Second, we tested two predictions of the Intermediate Disturbance Hypothesis and found that species diversity was greatest for plots characterized by disturbances of intermediate age. However, we did not detect significant differences in diversity between plots characterized by intermediate and high levels of disturbance, indicating that many species are adapted to or at least tolerant of high levels of disturbance. Third, we noted that the abundance of grasses decreased with increasing disturbance, while the abundance of dicots increased with increasing disturbance.     

The herb community of a tropical forest in central Panamá: dynamics and impact of mammalian herbivores

Mammals are hypothesized to either promote plant diversity by preventing competitive exclusion or limit diversity by reducing the abundance of sensitive plant species through their activities as browsers or disturbance agents. Previous studies of herbivore impacts in plant communities have focused on tree species and ignored the herbaceous community. In an experiment in mature-phase, tropical moist forest sites in central Panamá, we studied the impact of excluding ground-dwelling mammals on the richness and abundance of herbs in 16, 30×45-m plots. Within each plot, we censused the herbaceous community in 28, 2×2-m subplots (1,792 m² total area sampled). We identified over 54 species of herbs averaging 1.21 ramets m⁻² and covering approximately 4.25% of the forest floor. Excluding mammals for 5 years had no impact on overall species richness. Within exclosures, however, there was a significant two-fold increase in the density of rare species. Overall herbaceous density and percent cover did not differ between exclosures and adjacent control plots, although cover did increase over time. Mammalian exclusion significantly increased the total cover of three-dominant herb species, Pharus latifolius, Calathea inocephala, and Adiantum lucidum, but did not affect their density. This study represents one of the most extensive herbaceous community censuses conducted in tropical forests and is among a few that quantify herbaceous distribution and abundance in terms of both density and cover. Additionally, this work represents the first community level test of mammalian impacts on the herbaceous community in a tropical forest to date. Our results suggest that ground dwelling mammals do not play a key role in altering the relative abundance patterns of tropical herbs in the short term. Furthermore, our results contrast sharply with prior studies on similar temporal and spatial scales that demonstrate mammals strongly alter tree seedling composition and reduce seedling density. Thus, we question the pervasiveness of top–down control on tropical plant communities and the paradigm that defaunation will inexorably lead to widespread, catastrophic shifts in plant communities.     

Diversity and abundance of macro‐invertebrates on abandoned cattle kraals in a semi‐arid savanna

Abandoned cattle (Bos taurus) kraals are sources of habitat heterogeneity in dystrophic semi‐arid African savannas with a strong positive effect on soil nutrients and plant productivity. However, little is known regarding how macro‐invertebrate assemblages vary between abandoned kraals and the surrounding savanna matrix. We tested whether herbaceous biomass and basal and aerial covers and soil nutrients have an effect on aboveground and belowground macro‐invertebrate assemblages. Twelve abandoned kraals were contrasted with their paired control plots for soil characteristics, herbaceous productivity, and macro‐invertebrate assemblages in Save Valley Conservancy, Zimbabwe. Abandoned kraals had significantly higher concentrations of soil nitrogen (N), phosphorus (P), potassium (K), and calcium (Ca) as well as herbaceous biomass and basal and aerial covers than control plots. Both aboveground and belowground macro‐invertebrate species richness were higher on abandoned kraals. However, only belowground macro‐invertebrate diversity (Shannon H′ and Hill number 1) was significantly higher on abandoned kraals. Soil nutrients and herbaceous productivity had positive and significant correlations with the dominant taxa (Coleoptera, Hymenoptera, Hemiptera, Isoptera, and Myriapoda) on abandoned kraals. These results add to the growing body of evidence that abandoned kraals exert significant effects on savanna spatial heterogeneity years later, with implications on ecosystem processes and functioning.     

The population biology of Bromus tectorum in forests: effect of disturbance,grazing, and litter on seedling establishment and reproduction

Summary The effect of tree canopy, understory, herbivores, and litter depth on seedling establishment, survival, and reproduction of the alien grass, Bromus tectorum (cheatgrass), was examined in a series of experiments in four forest habitat types in western North America. Higher recruitment, survival, and reproduction on clearcuts, which would be expected if the overstory alone is limiting the distribution of cheatgrass in forests, were not observed. Removing the understory in an otherwise undisturbed Pinus ponderosa forest did, however, increase the emergence of B. tectorum, but plants in these experimentally-created openings were more vulnerable to grazing by small mammals. In contrast, removing the sparse understory in an Abies forest neither enhanced recruitment nor increased the incidence of grazing of B. tectorum seedlings. Regardless of the forest habitat, most grazed plants died before maturity; even fewer grazed plants produced seeds. Litter depth influenced both recruitment and biomass production: both the rate of germination and the size of resultant seedlings were lower on thick litter (6 cm) compared to results on thin litter (1.5 cm). In the more open Pinus ponderosa and Pseudotsuga menziesii forests, cheatgrass colonization may often occur in openings in the understory alone. Colonization in the more shady A. grandis and Thuja plicata forests is unlikely, however, unless the opening extends through both the understory and the overstory. As a result, cheatgrass is unlikely to increase in any of these forests unless the scale and incidence of disturbance increases substantially.     

Predicting yellow toadflax infestations in the Flat Tops Wilderness of Colorado

Understanding species–environment relationships is important to predict the spread of non-native species. Yellow toadflax (Linaria vulgaris Mill.) is an invasive perennial recently found in the Flat Tops Wilderness of the White River National Forest on the western slope of the Colorado Rocky Mountains. We hypothesized yellow toadflax occurrence could be predicted from easily measured site characteristics. We used logistic regression with stepwise selection to generate a model to predict yellow toadflax occurrence on a particular plot based on that site’s physical characteristics. The experimental design was a paired-plot study in two locations using circular 1,018-m² plots. Sixty-eight plots that did not contain yellow toadflax and 65 plots that contained yellow toadflax were sampled at the Ripple Creek site in 1999. In 2000, 54 non-toadflax plots and 55 toadflax-containing plots were sampled in the Marvine Creek site. Site characteristics sampled included: vegetation type; under-canopy light level; slope; aspect; soil properties; presence of disturbance, trails, and/or water; and total species richness. A model that correctly classified >90% of the 242 plots sampled included two vegetation type parameters, the presence of trails, and total species richness. Yellow toadflax is most often found in areas that were open-canopy sites, along trails, and with higher species diversity plots (>23 species). This approach can be used for other species in other areas to rapidly identify areas vulnerable to invasion.