Seedling growth responses to soil resources in the understory of a wet tropical forest

Plant growth responses to resources may be an important mechanism that influences species' distributions, coexistence, and community structure. Irradiance is considered the most important resource for seedling growth in the understory of wet tropical forests, but multiple soil nutrients and species have yet to be examined simultaneously with irradiance under field conditions. To identify potentially limiting resources, we modeled tree seedling growth as a function of irradiance and soil nutrients across five sites, spanning a soil fertility gradient in old-growth, wet tropical forests at La Selva Biological Station, Costa Rica. We measured an array of soil nutrients including total nitrogen (total N), inorganic N (nitrate [NO3-] and ammonium [NH4+]), phosphate (PO4-), and sum of base cations (SBC; potassium, magnesium, and calcium). Shade in the forest understory did not preclude seedling growth correlations with soil nutrients. Irradiance was a significant predictor of growth in 52% of the species, inorganic N in 54% (NO3- in 32%; NH4+ in 34%), total N in 47%, SBC in 39%, and PO4- in 29%. Overall, growth was correlated with both irradiance and soil nutrients in 45% of species and with soil nutrients only in an additional 48%; rarely was irradiance alone correlated with growth. Contrary to expectations, the magnitudes of growth effects, assessed as the maximum growth response to significant resources for each species, were similar for irradiance and most soil nutrients. Among species whose growth correlated with soil nutrients, the rank importance of nutrient effects was SBC, followed by N (total N, NO3-, and/or NH4+) and PO4-. Species' growth responsiveness (i.e., magnitudes of effect) to irradiance and soil nutrients was negatively correlated with species' shade tolerance (survival under 1% full sun). In this broad survey of species and resources, the nearly ubiquitous effects of soil nutrients on seedling growth challenge the idea that soil nutrients are less important than irradiance in the light-limited understory of wet tropical forests.     

Invasibility and Effects of Amur Honeysuckle in Southwestern Ohio Forests

The Asian exotic Amur honeysuckle ( Lonicera maackii [Rupr.] Herder) has become the dominant shrub in many forests in southwestern Ohio and in some other locations in the eastern United States. Our research focused on the invasibility of forest communities and relationships of L. maackii to the abundance of tree seedlings and herbs. We surveyed 93 forest stands near Oxford, Ohio (USA) to determine L. maackii cover, time since invasion, tree canopy cover, tree basal area, and a shade tolerance index. Stepwise multiple regression indicated that greater than one-half of the variation (r ² = 0.550) in Lonicera cover was correlated to five variables (in descending order of importance): tree canopy cover, distance from Oxford, shade tolerance index, tree basal area, and time since invasion. The results suggest that high light levels and proximity to an abundant seed source increase forest invasibility. Tree seedling density, species richness of seedlings, and herb cover were all inversely related to L. maackii cover. When Lonicera becomes abundant, future structure and composition of forests could be affected and local populations of herbs threatened.     

林木根系衰老研究方法与机制

根系衰老研究方法主要有根窗直接观测和挖根取样间接观测。在整株水平上,树木同化碳的能力,碳在不同器官间的分配,尤其是在地上部分和地下部分间的分配比例,对根的萌生和衰老起着重要作用;地上部分的生长过程和健康状况也对根系的生长和寿命有很大影响;当树木的生长环境受到某种胁迫时,树木的抵抗力下降,容易招致病原菌的危害,造成根系衰老和死亡。在生态系统水平上,干旱洪涝、干扰等胁迫和树木一土壤间的养分循环都会引起树木生长环境的改变,对根系的衰老过程发生作用;季节变化使树木的地上部分和地下部分的生理活动处于不同的旺盛期,从而使碳的分配方式有些改变,影响到根系的生长;土壤中养分的存在形式,某些离子的浓度也直接影响到根的寿命;病原菌对根系的危害与土壤中养分含量变化有关。所以,衰老过程受环境条件的影响,伴随着代谢,RNA和蛋白质合成速率的下降和(或)膜与细胞器结构的改变。     

Widespread density-dependent seedling mortality promotes species coexistence in a highly diverse Amazonian rain forest

Negative density-dependent mortality can promote species coexistence through a spacing mechanism that prevents species from becoming too locally abundant. Negative density-dependent seedling mortality can be caused by interactions among seedlings or between seedlings and neighboring adults if the density of neighbors affects the strength of competition or facilitates the attack of natural enemies. We investigated the effects of seedling and adult neighborhoods on the survival of newly recruited seedlings for multiple cohorts of known age from 163 species in Yasuni National Park, Ecuador, an ever-wet, hyper-diverse lowland Amazonian rain forest. At local scales, we found a strong negative impact on first-year survival of conspecific seedling densities and adult abundance in multiple neighborhood sizes and a beneficial effect of a local tree neighborhood that is distantly related to the focal seedling. Once seedlings have survived their first year, they also benefit from a more phylogenetically dispersed seedling neighborhood. Across species, we did not find evidence that rare species have an advantage relative to more common species, or a community compensatory trend. These results suggest that the local biotic neighborhood is a strong influence on early seedling survival for species that range widely in their abundance and life history. These patterns in seedling survival demonstrate the role of density-dependent seedling dynamics in promoting and maintaining diversity in understory seedling assemblages. The assemblage-wide impacts of species abundance distributions may multiply with repeated cycles of recruitment and density-dependent seedling mortality and impact forest diversity or the abundance of individual species over longer time scales.     

Tree species and mycorrhizal associations influence the magnitude of rhizosphere effects

Previous research on the effects of tree species on soil processes has focused primarily on the role of leaf litter inputs. We quantified the extent to which arbuscular mycorrhizal (AM) and ectomycorrhizal (ECM) tree species influence soil microbial activity and nutrient availability through rhizosphere effects. Rhizosphere soil, bulk soil, and fine roots were collected from 12 monospecifc plots (six AM and six ECM tree species) planted on a common soil at the Turkey Hill Plantations in Dryden, New York. Rhizosphere effects were estimated by the percentage difference between rhizosphere and bulk soil samples for several assays. Rhizosphere effects on soil microbes and their activities were significant for ECM species but in only a few cases for AM species. In AM tree species, microbial biomass, net N mineralization, and phosphatase enzyme activity in the rhizosphere were 10-12% greater than in bulk soil. In ECM tree species, rhizosphere effects for microbial biomass, C mineralization rates, net N mineralization, and phosphatase activity were 25-30% greater than bulk soil, and significantly greater than AM rhizosphere effects. The magnitude of rhizosphere effects was negatively correlated with the degree of mycorrhizal colonization in AM tree species (r = -0.83) and with fine root biomass (r = -0.88) in ECM tree species, suggesting that different factors influence rhizosphere effects in tree species forming different mycorrhizal associations. Rhizosphere effects on net N mineralization and phosphatase activity were also much greater in soils with pH < 4.3 for both AM and ECM tree species, suggesting that soil pH and its relation to nutrient availability may also influence the magnitude of rhizosphere effects. Our results support the idea that tree roots stimulate nutrient availability in the rhizosphere, and that systematic differences between AM and ECM may result in distinctive rhizosphere effects for C, N, and P cycling between AM and ECM tree species.     

Plant species differ in their ability to reduce allocation to non-beneficial arbuscular mycorrhizal fungi

Theory suggests that cheaters threaten the persistence of mutualisms, but that sanctions to prevent cheating can stabilize mutualisms. In the arbuscular mycorrhizal symbiosis, reports of parasitism suggest that reductions in plant carbon allocation are not universally effective. I asked whether plant species differences in mycorrhizal responsiveness would affect both their susceptibility to parasitism and their reduction in allocation to non-beneficial arbuscular mycorrhizal fungi (AMF) in high-phosphorus soils. In a greenhouse experiment, I found that two C3 grasses, Bromus inermis and Elymus repens, effectively suppressed root colonization and AMF hyphal abundance. Increases in soil phosphorus did not reduce the degree to which AMF increased plant biomass. In contrast, two C4 grasses, Andropogon gerardii and Schizachyrium scoparium, more weakly reduced root colonization and failed to suppress AMF hyphal abundance. Consequently, they experienced strong declines in their response to AMF, and one species suffered parasitism. Thus, species differ in susceptibility to parasitism and their reduction in allocation to non-beneficial AMF. These differences may affect the distribution and abundance of plants and AMF, as well as the stability of the mutualism.     

Provenance, life span, and phylogeny do not affect grass species' responses to nitrogen and phosphorus

Successful conservation management requires an understanding of how species respond to intervention. Native and exotic species may respond differently to management interventions due to differences arising directly from their origin (i.e., provenance) or indirectly due to biased representations of different life history types (e.g., annual vs. perennial life span) or phylogenetic lineages among provenance (i.e., native or exotic origin) groups. Thus, selection of a successful management regime requires knowledge of the life history and provenance-bias in the local flora and an understanding of the interplay between species characteristics across existing environmental gradients in the landscape. Here we tested whether provenance, phylogeny, and life span interact to determine species distributions along natural gradients of soil chemistry (e.g., soil nitrogen and phosphorus) in 10 upland prairie sites along a 600-km latitudinal transect running from southern Vancouver Island in British Columbia, Canada, to the Willamette Valley in Oregon, USA. We found that soil nitrate, phosphorus, and pH exerted strong control over community composition. However, species distributions along environmental gradients were unrelated to provenance, life span, or phylogenetic groupings. We then used a greenhouse experiment to more precisely measure the response of common grass species to nitrogen and phosphorus supply. As with the field data, species responses to nutrient additions did not vary as a function of provenance, life span, or phylogeny. Native and exotic species differed strongly in the relationship between greenhouse-measured tolerance of low nutrients and field abundance. Native species with the greatest ability to maintain biomass production at low nutrient supply rates were most abundant in field surveys, as predicted by resource competition theory. In contrast, there was no relationship between exotic-species biomass at low nutrient levels and field abundance. The implications of these findings for management of invasive species are substantial in that they overturn a general belief that reduction of nutrient supplies favors native species. The idiosyncratic nature of species response to nutrients in this study suggests that manipulation of nutrient supplies is unlikely to alter the overall balance between native and exotic species, although it may well be useful to control specific exotic species.     

三江源国家公园不同草地土壤微生物功能基因的差异性分析

土壤微生物在陆地生物地球化学循环过程中起着非常重要的作用。为了探索青藏高原高寒草地类型地上植被特性和地下土壤环境与土壤微生物功能基因之间关系,以三江源国家公园高寒草原、高寒沼泽化草甸及高寒草甸3种典型草地类型为研究对象,利用基因芯片(GeoChip 5.0)技术测定其微生物功能基因丰度,并分析它们之间的差异及影响因素。结果表明:(1)3种草地类型地上群落结构和地下土壤环境存在差异性,其中高寒草原物种多样性指数、pH值较高,沼泽化草甸中土壤含水量、微生物量碳、地上生物量、土壤速效氮含量较高,高寒草甸中则是土壤微生物量氮含量较高;(2)3种高寒草地类型的碳循环、氮循环、磷循环、有机修复的土壤微生物功能基因丰度存在显著差异,其中这些功能基因的丰度在高寒沼泽化草甸最高,高寒草甸、高寒草原次之;(3)地上植物物种多样性虽对功能基因丰度变化的解释率(r2)在57.1%-61.2%之间,但统计学上不显著(P>0.05),而微生物基因丰度随地上生物量的增加而增加,且解释率(r2)为77.5%-80.0%(P<0.05)。在pH、土壤含水量、土壤微生物量等地下土壤环境因子中,pH对功能基因丰度存在显著影响(P<0.01)解释率在83.4%-87.5%间,且土壤微生物功能基因丰度随土壤pH的增加而降低;土壤含水量、土壤微生物量对土壤微生物功能基因丰度的解释率分别为81.9%-83.1%(P<0.05)和76.8%-86.2%(P<0.05),微生物功能基因丰度随这两者含量的增加呈上升趋势。进一步运用RDA分析发现,pH、土壤微生物量、地上生物量是影响微生物功能基因丰度的主要因子,其中土壤微生物量是土壤有机质的重要组成部分,土壤有机质又是通过地上植被凋落物沉积所得到的。因此,地上植被特性的自上而下控制因子影响了土壤环境中自下而上的控制因子,间接的影响了微生物功能基因丰度。由此得出,地上植被特性和地下土壤环境因子共同作用控制了微生物功能基因丰度使其出现差异性。     

Effects of Patch Size and Type of Coffee Matrix on Ithomiine Butterfly Diversity and Dispersal in Cloud-Forest Fragments

Abstract: Determining the permeability of different types of landscape matrices to animal movement is essential for conserving populations in fragmented landscapes. We evaluated the effects of habitat patch size and matrix type on diversity, isolation, and dispersal of ithomiine butterflies in forest fragments surrounded by coffee agroecosystems in the Colombian Andes. Because ithomiines prefer a shaded understory, we expected the highest diversity and abundance in large fragments surrounded by shade coffee and the lowest in small fragments surrounded by sun coffee. We also thought shade coffee would favor butterfly dispersal and immigration into forest patches. We marked 9675 butterflies of 39 species in 12 forest patches over a year. Microclimate conditions were more similar to the forest interior in the shade‐coffee matrix than in the sun‐coffee matrix, but patch size and matrix type did not affect species richness and abundance in forest fragments. Furthermore, age structure and temporal recruitment patterns of the butterfly community were similar in all fragments, independent of patch size or matrix type. There were no differences in the numbers of butterflies flying in the matrices at two distances from the forest patch, but their behavior differed. Flight in the sun‐coffee matrix was rapid and directional, whereas butterflies in shade‐coffee matrix flew slowly. Seven out of 130 recaptured butterflies immigrated into patches in the shade‐coffee matrix, and one immigrated into a patch surrounded by sun coffee. Although the shade‐coffee matrix facilitated movement in the landscape, sun‐coffee matrix was not impermeable to butterflies. Ithomiines exhibited behavioral plasticity in habitat use and high mobility. These traits favor their persistence in heterogeneous landscapes, opening opportunities for their conservation. Understanding the dynamics and resource requirements of different organisms in rural landscapes is critical for identifying management options that address both animals’ and farmers’ needs.     

Growth, biomass production and photosynthesis of Cenchrus ciliaris L. under Acacia tortilis (Forssk.) Hayne based silvopastoral systems in semi arid tropics

The growth, biomass production and photosynthesis of Cenchrus ciliaris was studied under the canopies of 17 yr old Acacia tortilis trees in semi arid tropical environment. On an average the full grown canopy of A. tortilis at the spacing of 4 x 4 m allowed 55% of total Photosynthetically Active Radiation (PAR) which in turn increased Relative Humidity (RH) and reduced under canopy temperature to -1.75 degrees C over the open air temperature. C. ciliaris attained higher height under the shade of A. tortilis. The tiller production and leaf area index decreased marginally under the shade of tree canopies as compared to the open grown grasses. C. ciliaris accumulated higher chlorophyll a and b under the shade of tree canopies indicating its shade adaptation potential. The assimilatory functions such as rate of photosynthesis, transpiration, stomatal conductance, photosynthetic water use efficiency (PN/TR) and carboxylation efficiency (PN/CINT) decreased under the tree canopies due to low availability of PAR. The total biomass production in term of fresh and dry weight decreased under the tree canopies. On average of 2 yr C. ciliaris had produced 12.78 t ha(-1) green and 3.72 -t ha(-1) dry biomass under the tree canopies of A. tortilis. The dry matter yield reduced to 38% under the tree canopies over the open grown grasses. The A. tortilis + C. ciliaris maintained higher soil moisture, organic carbon content and available N P K for sustainable biomass production for the longer period. The higher accumulation of crude protein, starch, sugar and nitrogen in leaves and stem of C. ciliaris indicates that this grass species also maintained its quality under A. tortilis based silvopastoral system. The photosynthesis and dry matter accumulation are closely associated with available PAR indicating that for sustainable production of this grass species in the silvopasture systems for longer period about 55% or more PAR is required.     

Untangling positive and negative biotic interactions: views from above and below ground in a forest ecosystem

In ecological communities, the outcome of plant-plant interactions represents the net effect of positive and negative interactions occurring above and below ground. Untangling these complex relationships can provide a better understanding of mechanisms that underlie plant-plant interactions and enhance our ability to predict population, community, and ecosystem effects of biotic interactions. In forested ecosystems, tree seedlings interact with established vegetation, but the mechanisms and outcomes of these interactions are not well understood. To explore such mechanisms, we manipulated above- and belowground interactions among tree seedlings, shrubs, and trees and monitored seedling survival and growth of six species (Pinus banksiana, Betula papyrifera, P. resinosa, Quercus rubra, P. strobus, and Acer rubrum) in mature pine-dominated forest in northern Minnesota, USA. The forest had a moderately open canopy and sandy soils. Understory manipulations were implemented in the forest interior and in large gaps and included removal of shrubs (no interactions), tieback of shrubs (belowground), removal of shrubs with addition of shade (aboveground), and unmanipulated shrubs (both below- and aboveground). We found that shrubs either suppressed or facilitated seedling survival and growth depending on the seedling species, source of interaction (e.g., above- or belowground), and ecological context (e.g., gap or forest interior). In general, shrubs strongly influenced survival and growth in gaps, with more modest effects in the forest interior. In gaps, the presence of shrub roots markedly decreased seedling growth and survival, supporting the idea that belowground competition may be more important in dry, nutrient-poor sites. Shrub shade effects were neutral for three species and facilitative for the other three. Facilitation was more likely for shade-tolerant species. In the forest interior, shrub shade negatively affected seedling survival for the most shade-intolerant species. For several species the net effect of shrubs masked the existence of both positive and negative interactions above and below ground. Our results highlight the complexity of plant-plant interactions, demonstrate that outcomes of these interactions vary with the nature of resource limitation and the ecophysiology of the species involved, and suggest that ecological theory that rests on particular notions of plant-plant interactions (e.g., competition) should consider simultaneous positive and negative interactions occurring above and below ground.     

Life history trade-offs in tropical trees and lianas

It has been hypothesized that tropical trees partition forest light environments through a life history trade-off between juvenile growth and survival; however, the generality of this trade-off across life stages and functional groups has been questioned. We quantified trade-offs between growth and survival for trees and lianas on Barro Colorado Island (BCI), Panama using first-year seedlings of 22 liana and 31 tree species and saplings (10 mm < dbh < 39 mm) of 30 tree species. Lianas showed trade-offs similar to those of trees, with both groups exhibiting broadly overlapping ranges in survival and relative growth rates as seedlings. Life history strategies at the seedling stage were highly correlated with those at the sapling stage among tree species, with all species showing an increase in survival with size. Only one of 30 tree species demonstrated a statistically significant ontogenetic shift, having a relatively lower survival rate at the sapling stage than expected. Our results indicate that similar life history trade-offs apply across two functional groups (lianas and trees), and that life history strategies are largely conserved across seedling and sapling life-stages for most tropical tree species.     

Neutral theory and the evolution of ecological equivalence

Since the publication of the unified neutral theory in 2001, there has been much discussion of the theory, pro and con. The hypothesis of ecological equivalence is the fundamental yet controversial idea behind neutral theory. Assuming trophically similar species are demographically alike (symmetric) on a per capita basis is only an approximation, but it is equivalent to asking: How many of the patterns of ecological communities are the result of species similarities, rather than of species differences? The strategy behind neutral theory is to see how far one can get with the simplification of assuming ecological equivalence before introducing more complexity. In another paper, I review the empirical evidence that led me to hypothesize ecological equivalence among many of the tree species in the species-rich tropical forest on Barro Colorado Island (BCI). In this paper, I develop a simple model for the evolution of ecological equivalence or niche convergence, using as an example evolution of the suite of life history traits characteristic of shade tolerant tropical tree species. Although the model is simple, the conclusions from it seem likely to be robust. I conclude that ecological equivalence for resource use are likely to evolve easily and often, especially in species-rich communities that are dispersal and recruitment limited. In the case of the BCI forest, tree species are strongly dispersal- and recruitment-limited, not only because of restricted seed dispersal, but also because of low recruitment success due to heavy losses of the seedling stages to predators and pathogens and other abiotic stresses such as drought. These factors and the high species richness of the community strongly reduce the potential for competitive exclusion of functionally equivalent or nearly equivalent species.     

Proximity to forests drives bird conservation value of coffee plantations: implications for certification

Widespread loss of primary habitat in the tropics has led to increased interest in production landscapes for biodiversity conservation. In the Western Ghats biodiversity hotspot in India, shade coffee plantations are located in close proximity to sites of high conservation value: protected and unprotected forests. Coffee is grown here under a tree canopy that may be dominated by native tree species or by nonnative species, particularly silver oak (Grevillea robusta). We investigated the influence of properties at the local scale and the landscape scale in determining bird communities in coffee plantations, with particular emphasis on species of conservation priority. We used systematic point counts in 11 coffee plantation sites and analyzed data in a randomized linear modeling framework that addressed spatial autocorrelation. Greater proportion of silver oak at the local scale and distance to contiguous forests at the landscape scale were implicated as factors most strongly driving declines in bird species richness and abundance, while increased basal area of native tree species, a local-scale variable, was frequently related to increased bird species richness and abundance. The influence of local-scale variables increased at greater distances from the forest. Distance to forests emerged as the strongest predictor of declines in restricted-range species, with 92% reduction in the abundance of two commonly encountered restricted-range species (Pompadour Green Pigeon and Yellow-browed Bulbul) and a 43% reduction in richness of bird species restricted to Indian hill forests within 8 km of forests. Increase in proportion of silver oak from 33% to 55% was associated with 91% reduction in the abundance of one commonly encountered restricted-range species (Crimson-fronted Barbet). One conservation strategy is providing incentives to grow coffee in a biodiversity-friendly manner. One implication of our study is that plantations located at varying distances to the forest cannot be compared fairly for biodiversity friendliness by existing certification methodology. Another is that conservation of existing forests at the landscape scale is essential for maintaining higher biodiversity in coffee plantations. Incentive schemes that promote conservation of remnant forests at the landscape scale and biodiversity-friendly practices locally and that relate to coffee communities as a whole rather than individual planters are likely to be more effective.     

Common ectomycorrhizal networks may maintain monodominance in a tropical rain forest

Most tropical rain forests contain diverse arrays of tree species that form arbuscular mycorrhizae. In contrast, the less common monodominant rain forests, in which one tree species comprises more than 50% of the canopy, frequently contain ectomycorrhizal (ECM) associates. In this study, I explored the potential for common ECM networks, created by aggregations of ECM trees, to enhance seedling survivorship near parent trees. I determined the benefit conferred by the common ECM network on seedling growth and survivorship of an ECM monodominant species in Guyana. Seedlings with access to an ECM network had greater growth (73% greater), leaf number (55% more), and survivorship (47% greater) than seedlings without such access, suggesting that the ECM network provides a survivorship advantage. A survey of wild seedlings showed positive distance-dependent distribution and survival with respect to conspecific adults. These experimental and survey results suggest that the negative distance-dependent mechanisms at the seedling stage thought to maintain tropical rain forest diversity are reversed for ECM seedlings, which experience positive feedbacks from the ECM network. These results may in part explain the local monodominance of an ECM tree species within the matrix of high-diversity, tropical rain forest.     

Seed predation by Neotropical rain forest mammals increases diversity in seedling recruitment

Seed dispersal and seedling recruitment (the transition of seeds to seedlings) set the spatiotemporal distribution of new individuals in plant communities. Many terrestrial rain forest mammals consume post-dispersal seeds and seedlings, often inflicting density-dependent mortality. In part because of density-dependent mortality, diversity often increases during seedling recruitment, making it a critical stage for species coexistence. We determined how mammalian predators, adult tree abundance, and seed mass interact to affect seedling recruitment in a western Amazonian rain forest. We used exclosures that were selectively permeable to three size classes of mammals: mice and spiny rats (weighing <1 kg), medium-sized rodents (1-12 kg), and large mammals (20-200 kg). Into each exclosure, we placed seeds of 13 tree species and one canopy liana, which varied by an order of magnitude in adult abundance and seed mass. We followed the fates of the seeds and resulting seedlings for at least 17 months. We assessed the effect of each mammalian size class on seed survival, seedling survival and growth, and the density and diversity of the seedlings that survived to the end of the experiment. Surprisingly, large mammals had no detectable effect at any stage of seedling recruitment. In contrast, small- and medium-sized mammals significantly reduced seed survival, seedling survival, and seedling density. Furthermore, predation by small mammals increased species richness on a per-stem basis. This increase in diversity resulted from their disproportionately intense predation on common species and large-seeded species. Small mammals thereby generated a rare-species advantage in seedling recruitment, the critical ingredient for frequency dependence. Predation by small (and to a lesser extent, medium-sized) mammals on seeds and seedlings significantly increases tree species diversity in tropical forests. This is the first long-term study to dissect the effects of various mammalian predators on the recruitment of a diverse set of tree species.     

Deer herbivory alters forest response to canopy decline caused by an exotic insect pest.

Hemlock woolly adelgid (HWA; Adelges tsugae) infestations have resulted in the continuing decline of eastern hemlock (Tsuga canadensis) throughout much of the eastern United States. While the initial impacts of HWA infestations have been documented, our understanding of forest response to this disturbance remains incomplete. HWA infestation is not occurring in isolation but within a complex ecological context. The role of potentially important interacting factors, such as elevated levels of white-tailed deer herbivory, is poorly understood. Despite the potential for herbivory to alter forest successional trajectories following a canopy disturbance, little is known about herbivory-disturbance interactions, and herbivory is rarely considered in assessing forest response to a co-occurring disturbance. We used repeated censuses of deer exclosures and paired controls (400 paired plots) to quantify the impact of deer herbivory on tree seedling species abundance in 10 eastern hemlock ravines that span a gradient in HWA-induced canopy decline severity. Use of a maximum likelihood estimation framework and information theoretics allowed us to quantify the strength of evidence for alternative models developed to estimate the impacts of herbivory on tree seedling abundance as a function of varying herbivore density and canopy decline severity. The exclusion of deer herbivory had marked impacts on the abundance of the studied seedling species: Acer rubrum, Acer saccharum, Betula lenta, Nyssa sylvatica, Quercus montana, and Tsuga canadensis. For all six species, the relationship between seedling abundance and deer density was either exponential or saturating. Although the functional form of the response varied among seedling species, the inclusion of both deer density and canopy decline severity measures consistently resulted in models with substantially greater support. Canopy decline resulted in higher proportional herbivory impacts and altered the ranking of herbivory impacts by seedling species. Our results suggest that, by changing species' competitive abilities, white-tailed deer herbivory alters the trajectory of forest response to this exotic insect pest and has the potential to shift future overstory composition.     

Bird Populations in Shade and Sun Coffee Plantations in Central Guatemala

We studied the avifauna of sun and shade coffee plantations and associated mid-elevation habitats during the dry season of 1995. The three plantation types (Inga, Gliricidia, and sun) showed high faunistic similarities with each other and were both distinct and depauperate compared to matorral and forest patch habitats. Of all the coffee plantation habitats, Inga shade had the highest diversity. Species associated with wooded vegetation were more common in shade plantations, particularly in Inga. A second census showed a decline in bird numbers that was more pronounced in sun and Gliricidia than in Inga plantations. Overall, differences between the plantation types were small and all coffee plantations were less diverse than traditional coffee farms previously studied in nearby Chiapas, México. The relatively low bird diversity was probably due to the low stature, low tree species diversity, and heavy pruning of the canopy. These features reflect management practices that are common throughout Latin America. The most common species of birds in all coffee plantation habitats were common second-growth or edge species; more specialized forest species were almost completely absent from plantations. Furthermore, many common matorral species were rare or absent from coffee plantations, even sun plantations with which matorral shares a similar superficial structure. Coffee plantations probably will only be important for avian diversity if a tall, taxonomically and structurally diverse canopy is maintained. We suggest this is most likely to occur on farms that are managed for a variety of products rather than those designated entirely for the production of coffee.     

Constraints on tree seedling establishment in montane grasslands of the Valles Caldera, New Mexico

Montane and subalpine grasslands are prominent, but poorly understood, features of the Rocky Mountains. These communities frequently occur below reversed tree lines on valley floors, where nightly cold air accumulation is spatially coupled with fine soil texture. We used field experiments to assess the roles of minimum temperature, soil texture, grass competition, and ungulate browsing on the growth, photosynthetic performance, and survival of transplanted ponderosa pine (Pinus ponderosa) seedlings at 32 sites straddling such reversed tree lines in the Valles Caldera National Preserve (VCNP) of the Jemez Mountains, New Mexico (USA). Seedling growth increased most strongly with increasing nighttime minimum temperatures away from the valley bottoms; seedlings experiencing the coldest temperatures on the caldera floor exhibited stunted needles and often no measurable height growth. Based on the chlorophyll fluorescence ratios PhiPSII and Fv/Fm, we found that low minimum temperatures, low soil moisture, and fine soil texture all contributed to photoinhibition. Neighboring herbs had only minor negative effects on seedlings. We found no effect of ungulates, but golden-mantled ground squirrels (Spermophilus lateralis) caused substantial seedling mortality. Second-year seedling survival was highest on sandy soils, and third-year survival was highest at sites with higher minimum temperatures. We conclude that differential tree seedling establishment driven by low minimum temperatures in the valley bottoms is the primary factor maintaining montane grasslands of the VCNP, although this process probably operated historically in combination with frequent surface fire to set the position of the tree line ecotone. As at alpine tree lines, reversed tree lines bordering montane and subalpine grasslands can represent temperature-sensitive boundaries of the tree life form.     

Facilitation drives local abundance and regional distribution of a rare plant in a harsh environment

The importance of facilitation to local community dynamics is becoming increasingly recognized. However, the predictability of positive interactions in stressful environments, the balance of competition and facilitation along environmental gradients, and the scaling of local positive interactions to regional distributions are aspects of facilitation that remain unresolved. I explored these questions in a habitat specialist, Delphinium uliginosum, and a moss, Didymodon tophaceus, both found in small serpentine wetlands. I tested three hypotheses: (1) moss facilitates germination, growth, and/or fecundity of D. uliginosum; (2) facilitation is stronger at the harsher ends of gradients in soil moisture, toxicity, and/or biomass; and (3) facilitation is reflected in positive associations at the levels of local abundance and regional occurrence. Although considerable competitive interactions occurred in later life stages, moss strongly facilitated D. uliginosum seedling emergence. There was no evidence that this facilitative effect weakened, or switched to competition, in benign environments. D. uliginosum was more locally abundant and more frequently present, across a large portion of its range, with than without moss, indicating a net facilitative effect in the face of competitive influences. Facilitated recruitment, possibly by seed retention, was found to be an important control on abundance and distribution in this rare species.