Old-field seedling responses to insecticide,seed addition,and competition

The controls of seedling emergence and survival determine the potential distribution of adult plants and, thereby, plant community structure. Seed availability, competition from established neighbors, and seedling predation may all limit seedling recruitment. In this field experiment, we followed the emergence and survival of seedlings of three perennial forbs, Achillea millefolium, Hypericum perforatum, and Monarda fistulosa, in old-fields in southeastern Michigan, USA. As adults, all three have aromatic foliage that may deter herbivory, but seedlings may be more susceptible than adults. To establish the relative importance of potential controls on seedling numbers, we manipulated seed availability through seed additions, the influence of competitors by neighbor-removals, and the influence of insect herbivores with insecticide in a fully factorial field experiment. Seed addition and insecticide never affected seedling emergence for any species. Competition from established neighbors controlled seedling emergence for all three species and decreased Achillea survival. Insecticide significantly increased Monarda seedling survival in competition plots, significantly increased Hypericum survival in open plots, and had no effect on Achillea. Notably, insecticide increased survival of the native Monarda fistulosa more than the two introduced species. While neighbors strongly reduced emergence and survival of all three species, herbivores acted on a species-specific basis. These results suggest the differential effects of insects may contribute more to the seedling species composition and abundance patterns than the less-selective influence of competition.     

Salinity and light interactively affect neotropical mangrove seedlings at the leaf and whole plant levels

We have studied the interactive effects of salinity and light on Avicennia germinans mangrove seedlings in greenhouse and field experiments. We hypothesized that net photosynthesis, growth, and survivorship rates should increase more with an increase in light availability for plants growing at low salinity than for those growing at high salinity. This hypothesis was supported by our results for net photosynthesis and growth. Net daily photosynthesis did increase more with increasing light for low-salinity plants than for high-salinity plants. Stomatal conductance, leaf-level transpiration, and internal CO₂ concentrations were lower at high than at low salinity. At high light, the ratio of leaf respiration to assimilation was 2.5 times greater at high than at low salinity. Stomatal limitations and increased respiratory costs may explain why, at high salinity, seedlings did not respond to increased light availability with increased net photosynthesis. Seedling mass and growth rates increased more with increasing light availability at low than at high salinity. Ratios of root mass to leaf mass were higher at high salinity, suggesting that either water or nutrient limitations may have limited seedling growth at high salinity in response to increasing light. The interactive effects of salinity and light on seedling size and growth rates observed in the greenhouse were robust in the field, despite the presence of other factors in the field—such as inundation, nutrient gradients, and herbivory. In the field, seedling survivorship was higher at low than at high salinity and increased with light availability. Interestingly, the positive effect of light on seedling survivorship was stronger at high salinity, indicating that growth and survivorship rates are decoupled. In general, this study demonstrates that environmental effects at the leaf-level also influence whole plant growth in mangroves.     

Simulated effects of herb competition on planted Quercus faginea seedlings in Mediterranean abandoned cropland

Abstract. We tested simulated effects of herb competition on the performance of planted seedlings of Quercus faginea ssp. faginea in Mediterranean abandoned cropland. We produced three types of environment with respect to herb competition: absence of competition (AC), below‐ground competition (BGC), and total competition (TC). We assessed the performance of Q. faginea seedlings in each treatment in five ways: (1) seedling mortality, (2) leaf length and total plant leaf area, (3) water potential, (4) total biomass and biomass allocation, and (5) non‐structural carbohydrate storage in different plant organs. We also measured (6) soil moisture at different depths and (7) biomass production of herbs. The TC treatment reduced water availability more than the BGC treatment, in agreement with the most pronounced water stress in seedlings under TC conditions. BGC and TC treatments showed a high and similar seedling mortality, which was one order of magnitude higher than that in the AC treatment. Competition treatments affected glucose concentration in both shoots and roots, and followed the rank TC > BGC > AC. Q. faginea seedlings might compensate a lower water availability through glucose accumulation in leaves to reduce the osmotic potential. There was a maximum starch concentration in the BGC treatment that hints that a moderate resource limitation would limit tissue growth but not carbon assimilation. We conclude that the negative effects of herbs on Q. faginea seedlings are mostly a result of competition for water, and that this competition is noticeable since the earliest stages of the establishment. Complete weed removal is a technique that would strongly improve seedling survivorship.     

Herbivory and plant competition reduce mountain beech seedling growth and establishment in New Zealand

The effect of herbivory and resource availability on the competitive ability of different plant species has been an area of intense debate amongst plant ecologists for at least two decades, but the interactive effects of herbivory and plant competition between woody and herbaceous plants are rarely studied and theory is poorly developed. This study used experimental manipulations on transplanted and naturally occurring mountain beech (Nothofagus solandri var. cliffortioides) seedlings to show the effects of deer browsing and competition from deer-induced, herbaceous turf communities on mountain beech regeneration in New Zealand. Differences in the species composition of these turfs had little effect on mountain beech seedling establishment, but turf removal increased seedling growth and survivorship, showing that competition with other plants had direct effects on mountain beech regeneration. Deer browsing reduced the establishment and growth of seedlings, but the size of this effect did not vary with light and nutrient availability. There was no immediate compositional response of turf communities following the removal of deer browsing. The addition of nutrients appeared to reduce the intensity of belowground competition (stem growth increased relative to root growth) and increase seedling mortality, but there was no effect of changing levels of light. These results showed simple and direct negative effects of deer browsing on mountain beech regeneration. Indirect negative effects on regeneration were caused by deer-induced turf communities. We found little evidence for interactive effects between herbivory, plant competition and the availability of light or nutrients on seedling regeneration, which suggests that these factors acted independently. Nomenclature: Beever et al. (1992); Parsons et al. (1995); Edgar and Connor (2000); and Brownsey and Smith-Dodsworth (2000). Raukaua simplex is described by Mitchell et al. (1997). Coprosma “taylorae” is referred to by Eagle (1986) and Halocarpus biformis, Phyllocladus alpinus, Podocarpus hallii and Podocarpus nivalis by Wilson and Galloway (1993).     

Refuse dumps from leaf-cutting ant nests reduce the intensity of above-ground competition among neighboring plants in a Patagonian steppe

In arid environments, the high availability of sunlight due to the scarcity of trees suggests that plant competition take place mainly belowground for water and nutrients. However, the occurrence of soil disturbances that increase nutrient availability and thereby promote plant growth may enhance shoot competition between neighboring plants. We conducted a greenhouse experiment to evaluate the influence of the enriched soil patches generated by the leaf-cutting ant, Acromyrmex lobicornis, on the performance of the alien forb Carduus thoermeri (Asteraceae) under different intraspecific competition scenarios. Our results showed that substrate type and competition scenario affected mainly aboveground plant growth. As expected, plants growing without neighbors and in nutrient-rich ant refuse dumps showed more aboveground biomass than plants growing with neighbors and in nutrient-poor steppe soils. However, aboveground competition was more intense in nutrient-poor substrates: plants under shoot and full competition growing in the nutrient-rich ant refuse dumps showed higher biomass than those growing on steppe soils. Belowground biomass was similar among focal plants growing under different substrate type. Our results support the traditional view that increments in resource availability reduce competition intensity. Moreover, the fact that seedlings in this sunny habitat mainly compete aboveground illustrates how limiting factors may be scale-dependent and change in importance as plants grow.     

Recovery patterns of three chaparral shrub species after wildfire

Summary In a mature, even aged stand of mixed chaparral, Rhus laurina (facultative resprouter) had consistently higher water potentials and deeper roots than Ceanothus spinosus (facultative resprouter) and Ceanothus megacarpus (obligate seeder). For two years following a wildfire, the same stand of chaparral had resprouts with higher survivorships, predawn water potentials, stomatal conductances, photosynthetic rates and shoot elongation rates than seedlings. Supplemental irrigation of seedlings during summer months removed differences between resprouts and seedlings suggesting that the cause of such differences was limited water availability to the shoot tissues of seedlings. After two years of postfire regrowth, mean seedling survivorship for the obligate seeder (C. megacarpus) was 42%, whereas seedling survivorship for facultative resprouters was only 18% (C. spinosus) and 0.01% (R. laurina). Our results are consistent with the hypothesis that lack of resprouting ability among obligate seeders is offset by an enhanced ability to establish seedlings after wildfire, allowing obligate seeders to maintain themselves in mixed populations through many fire cycles.     

Coexistence and interference between a native perennial grass and non-native annual grasses in California

Little is known about the potential for coexistence between native and non-native plants after large-scale biological invasions. Using the example of native perennial bunchgrasses and non-native annual grasses in California grasslands, we sought to determine the effects of interference from non-native grasses on the different life stages of the native perennial bunchgrass Nassella pulchra. Further, we asked whether N. pulchra interferes with non-native annual grasses, and whether competition for water is an important component of these interspecific interactions in this water-limited system. In a series of field and greenhouse experiments employing neighbor removals and additions of water, we found that seedling recruitment of N. pulchra was strongly seed-limited. In both field and greenhouse, natural recruitment of N. pulchra seedlings from grassland soil was extremely low. In field plots where we added seeds, addition of water to field plots increased density of N. pulchra seedlings by 88% and increased total aboveground N. pulchra seedling biomass by almost 90%, suggesting that water was the primary limiting resource. In the greenhouse, simulated drought early in the growing season had a greater negative effect on the biomass of annual seedlings than on the seedlings of N. pulchra. In the field, presence of annuals reduced growth and seed production of all sizes of N. pulchra, and these effects did not decrease as N. pulchra individuals increased in size. These negative effects appeared to be due to competition for water, because N. pulchra plants showed less negative pre-dawn leaf water potentials when annual neighbors were removed. Also, simply adding water caused the same increases in aboveground biomass and seed production of N. pulchra plants as removing all annual neighbors. We found no evidence that established N. pulchra plants were able to suppress non-native annual grasses. Removing large N. pulchra individuals did not affect peak biomass per unit area of annuals. We conclude that effects of interference from non native annuals are important through all life stages of the native perennial N. pulchra. Our results suggest that persistence of native bunchgrasses may be enhanced by greater mortality of annual than perennial seedlings during drought, and possibly by reduced competition for water in wet years because of increased resource availability. Received: 12 November 1998 / Accepted: 4 August 1999     

Capture and allocation of nitrogen byQuercus douglasii seedlings in competition with annual and perennial grasses

Summary The spatial overlap of woody plant root systems and that of annual or perennial grasses promotes competition for soil-derived resources. In this study we examined competition for soil nitrogen between blue oak seedlings and either the annual grassBromus mollis or the perennial grassStipa pulchra under controlled outdoor conditions. Short-term nitrogen competition was quantified by injecting¹⁵N at 30 cm depth in a plane horizontal to oak seedling roots and that of their neighbors, and calculating¹⁵N uptake rates, pool sizes and¹⁵N allocation patterns 24 h after labelling. Simultaneously, integrative nitrogen competition was quantified by examining total nitrogen capture, total nitrogen pools and total nitrogen allocation.Stipa neighbors reduced inorganic soil nitrogen content to a greater extent than didBromus plants. Blue oak seedlings responded to lower soil nitrogen content by allocating lower amounts of nitrogen per unit of biomass producing higher root length densities and reducing the nitrogen content of root tissue. In addition, blue oak seedlings growing with the perennial grass exhibited greater rates of¹⁵N uptake, on a root mass basis, compensating for higher soil nitrogen competition inStipa neighborhoods. Our findings suggest that while oak seedlings have lower rates of nitrogen capture than herbaceous neighbors, oak seedlings exhibit significant changes in nitrogen allocation and nitrogen uptake rates which may offset the competitive effect annual or perennial grasses have on soil nitrogen content.     

Direct and indirect interactions among plants explain counterintuitive positive drought effects on an eastern Mediterranean shrub species

Coexistence of woody and herbaceous plants may be governed by a complex set of direct and indirect interactions, whose relative importance have been rarely assessed. We experimentally studied woody species establishment in a mixed plant community by disentangling the potential role of such biotic interactions and the effect of environmental variations on them. Seedling establishment of the common eastern Mediterranean shrub species Sarcopoterium spinosum was investigated under different rainfall and light conditions, combined with the effect of the presence of adult shrubs and annual neighbors. We predicted that seedlings will be directly affected by competition with annuals with increasing water availability, while direct effects of adult shrubs will be positive via amelioration of water stress. Indirect effects were expected beneath shrub canopies due to reduced water stressed and light availability for both annuals and shrub seedlings, which may intensify competition between annuals and shrub seedlings. To test these predictions we performed field and garden experiments in which we combined manipulation of shrub and annual presence with manipulations of water availability and light conditions to simulate the effect of shrub canopy. In contrast to our prediction, shrub seedling establishment was not facilitated but inhibited by adult shrubs because of light limitation. As expected, annuals had direct negative effects on shrub seedlings under wet conditions, which shifted to neutral or positive effects under dry conditions. Thus, interactions among shrubs and annuals, and in particular the release from competition during drought years, leads to a counterintuitive positive effect of drought on shrub seedling establishment. Our findings point to the importance of experimentally studying multidimensional interactions for coexistence of different life forms and to the underestimated role of light for success in water‐limited ecosystems.     

Tissue water relations of three chaparral shrub species after wildfire

Summary We compared the tissue water relations among resprouts and seedlings of three chaparral species during the first summer drought after wildfire. Two of the species, Rhus laurina and Ceanothus spinosus recover after fire by a combination of resprouting and seedling establishment (facultative resprouters), whereas a third species, Ceanothus megacarpus recovers by seedling establishment alone (obligate seeder). Our objectives were to document any differences in tissue water characteristics that might arise between resprouts and seedlings and to test the hypothesis that seedlings of obligate seeders develop more drought tolerant characteristics of their tissues than seedlings of facultative resprouters. We found that resprouts had much higher predawn values of water potential, osmotic potential, and turgor potentials than seedlings. Predawn turgor potentials of resprouts were 1.5 MPa through July and August when turgor potentials for seedlings remained near 0 MPa. During summer months, midday water potentials were 2 to 3 MPa higher for resprouts than seedlings and midday conductances of resprouts were two to five fold greater than those of seedlings. Even though resprouts did not experience severe water stress like seedlings, their tissue water characteristics, as determined by pressure-volume curve analyses, were similar by the peak of the drought in August. Further-more, the tissue water characteristics of seedlings from the obligate seeder, C. megacarpus, were similar to those of facultative resprouters — R. laurina, and C. spinosus. We attribute the observed differences in plant water status between resprouts and seedlings to differences in rooting depths and access to soil moisture reserves during summer drought. We conclude that the higher growth rates, photosynthetic performance, and survivorship of postfire resprouts are primarily a result of higher water availability to resprouting tissues during summer months. It appears that the greater seedling survivorship during summer drought observed for the obligate seeder, C. megacarpus, is not associated with more favorable tissue water characteristics.     

Seedling performance in a trioecious cactus,Pachycereus pringlei: Effects of maternity and paternity

We studied seed germination and the growth and survivorship of seedlings of females and hermaphrodites ofPachycereus pringlei (cardon), a Mexican columnar cactus whose geographically variable breeding system includes trioecy and gynodioecy. Results of a two-year field experiment conducted near Bahia Kino, Sonora, Mexico and a ten-month laboratory experiment were similar and did not support the hypothesis that seedlings of females outperform those of hermaphrodites. In the field, percent seed germination and 2-yr seedling survivorship averaged 66% and 95%, respectively and did not differ among six treatment classes. Seedlings of hermaphrodites generally were larger than those of females at the end of both experiments. Selfed seedlings of hermaphrodites did not grow more slowly than outcrossed seedlings of hermaphrodites or females. Hermaphrodite seedlings performed best when pollinated with hermaphrodite pollen; female seedlings performed best with male pollen. We conclude that superior seedling performance cannot explain why females are able to coexist with hermaphrodites in populations of this cactus. Instead, we postulate that greater annual seed production, which averaged 1.6 times higher in females than in hermaphrodites in two years, may be sufficient to allow females to co-occur with hermaphrodites in this large, longlived plant, especially if sex determination involves cytoplasmic-nuclear inheritance.     

Limitations to seedling establishment in a mesic Hawaiian forest

While invasive species may be visible indicators of plant community degradation, they may not constitute the only, or even the primary, limitation to stand regeneration. We used seed-augmentation and grass-removal experiments under different canopy conditions to assess the relative importance of dispersal limitation, resource availability, and competition on seedling establishment in the understory shrubs Sophora chrysophilla, Dodonea viscosa, and Pipturus albidus in a montane mesic forest in Hawaii. The study location was an Acacia koa-Metrosideros polymorpha forest at 1000–1500 m elevation on the leeward side of Hawaii Island; it is a closed-canopy forest historically subject to logging and grazing by cattle and sheep and currently dominated by the exotic grass, Ehrharta stipoides, in the herb layer. Seedling establishment after 1 and 2 years was strongly dispersal limited in Sophora and Dodonea, but not in Acacia, a non-augmented species in which abundant seedlings established, nor in Pipterus, in which only one seedling established in 2 years. Grass cover reduced seedling establishment in Acacia, Sophora, and Dodonea and, for the latter two species, seedling establishment was substantially greater in the warmer, more moist forest at the lowest elevation. Light, moisture, and resin-captured N and P were more strongly affected by elevation and canopy composition than by grass cover, but in most cases seedling establishment was not positively correlated with resource availability. Limitations to the establishment of woody seedlings in this forest-grassland mixture vary among species; however, both dispersal limitation and competition from a shade-tolerant grass are important deterrents to regeneration in these forests.     

Early establishment of planted Retama sphaerocarpa seedlings under different levels of light,water and weed competition

Large amounts of former cropland are being abandoned in developedregions. To formulate guidelines for land reclamation programmes, we exploredthe effects of artificial shading, irrigation, and removal of weed competitionon the performance of Retama sphaerocarpa (L.) Boiss.seedlings in a factorial experiment located in an abandoned cropland in CentralSpain. R. sphaerocarpa is of interest for revegetationbecause it is a drought tolerant leguminous shrub that is a major structuralcomponent of the native plant community. Seedling performance was evaluated inthree ways: seedling survivorship, growth, and photochemical efficiency. Wealsomeasured soil moisture and weed biomass production and found that bothincreasedunder artificial shading conditions. Soil moisture increased very slightlywhereweeds were removed. Thus, increased transpiration from weeds outweighed reducedevaporation from soils due to shading by weeds. Artificial shading was the mosteffective treatment for seedling survivorship, followed by removal ofcompetition by weeds. After summer, 34 % of the seedlings survived in the mostfavourable conditions (artificially shaded plots where weeds were removed),compared to ca. 1 % in full-light plots with no removal of weed competition. Apositive effect of irrigation was found for growth of seedling cover and heightin shaded plots. The analysis of photochemical efficiency pointed out therelevance of weed competition removal, and confirmed the usefulness of fastfluorescence transient techniques for the quantification of seedlingperformance. The data suggest that competition between seedlings and weeds wasprimarily for water rather than for light. We conclude that i) artificialshading improved seedling performance, but this is a little practical techniquebecause of its cost; ii) as weeds compete with, rather than facilitate, plantedseedlings, weed clipping around the seedlings is a feasible technique thatwouldimprove seedling survival; and iii) seedling performance could alsoconsiderablyimprove with a higher irrigation than was used in this experiment(75lm^–2 per growth period), provided that weedsare removed.     

Establishment and growth of seedlings of Eucalyptus obliqua: Interactive effects of litter,water, and pathogens

We studied how leaf litter, water and fungal pathogens affect the establishment, survivorship, and growth of Eucalyptus obliqua using glasshouse and field experiments. In a glasshouse experiment, the presence of leaf litter of E. obliqua increased the establishment of seedlings when the seeds were placed beneath the litter, but dense litter layers reduced seedling emergence when seeds were placed on top. Results from a complementary experiment suggest that the effect of litter may be mediated by the high humidity microenvironment it creates, which may enhance imbibition of unburied seeds. In another glasshouse experiment, litter in combination with excessive water applications reduced seedling emergence and increased seedling mortality, but the addition of fungicide reverted these effects. Litter with lower water application or fungicide added increased seedling emergence. In all cases higher biomass in seedlings growing in litter treatments was associated with earlier emergence. In two recently burnt sites, the density of seedlings of E. obliqua was positively correlated to the presence of litter. Experimental addition of litter in the field reduced soil temperature during the summer, and together with watering, increased soil water content, and the water potential of established seedlings. However, neither treatment had any positive effect on their growth or survivorship. On the contrary, litter addition reduced the height and number of leaves, probably as a result of increased herbivory. We conclude that in Mediterranean-type systems litter may produce various effects on seedling establishment depending on their developmental stage, and the season.     

Spatial patterns of tree seedling establishment and their relationship to environmental variables in a cold-temperate deciduous forest of eastern North America

Patterns of seedling recruitment may have persistent effects on population and community processes. Assuming seed availability is not limiting, the environmental sieve (i.e., the suite of factors influencing seed germination and seedling emergence and survival) determines how many seedlings establish and, most importantly, where they do so. In this study, we identify the spatial structure of some resources and abiotic conditions known to be significant for tree seedling emergence and survival and determine how these environmental factors influence the establishment of Fagus grandifolia, Acer saccharum, Fraxinus americana, and Ostrya virginiana in a deciduous forest of southern Québec (Canada). We expect an increase from Fagus, through Acer and Fraxinus, to Ostrya in the control of environmental variables on seedling emergence and survival, because of differences in the seed size of these species. Density of newly-emerged seedlings of all four species showed positive spatial autocorrelation at distances of up to ca. 10 m. Environmental variables were also structured at the same spatial scale, except for soil moisture. Acer seedling emergence pattern was positively correlated to photosynthetic photon flux density (PPFD), and the pattern of Fraxinus to soil N and moisture. Seedling survival was not spatially autocorrelated for any of the four species, although it was positively density-dependent in Acer and Fagus. In only Ostrya was seedling survival correlated (positively) to one of the environmental variables studied, i.e., PPFD. Overall, environmental variables were spatially less heterogeneous than seedling emergence and survival. Either seed availability was not saturating or factors not considered here, such as competition and predation (the intensity of which often varies with resources and/or abiotic conditions), modified the influence that the physical environment had on patterns of seedling establishment. Our prediction of a greater environmental control on seedling emergence and survival in small-seed species was not totally confirmed.     

Evaluation of different approaches for modelling individual tree seedling height growth

We compared different approaches for modelling height growth of individual beech seedlings in a controlled factorial experiment as well as in field data from naturally regenerated beech seedlings under the canopy of overstorey mature beech trees. Several competition indices, a model of overstorey fine root density, relative photosynthetically active radiation (PAR) values, and soil water values were used in these approaches. In the factorial experiment relative PAR and soil water content were measured and used for the prediction of seedlings height growth. In the field experiment this was done by using relative PAR and estimated fine root biomass as a surrogate for below ground resource availability. The latter approach was compared with a model where we used various competition indices representing the impact of overstorey trees on beech seedlings. Our results suggested that (1) models which combine resource based growth functions are suitable for the prediction of individual height growth of beech seedlings. Resource based models offer the opportunity to investigate on the independent multiplicative effect of irradiance and water supply and their interactions on tree seedlings. It was (2) shown that a combined model could be used not only to predict individual height growth of beech seedlings in a controlled experiment but also in the field. The model parameters of a pure light response function for the controlled factorial experiment are comparable to those obtained in the field study. The results showed (3) that the precision of predicting beech seedlings height growth is comparable between the model types tested within this study. Approximately half of the observed variation in seedlings relative height growth rate could be explained. However, the simple competition index approach provides no information on the environmental factors constraining tree seedlings growth; whereas the multiplicative combined models can be used to get a better understanding of growth dynamics in the field.     

EXPERIMENTAL STUDIES OF THE EVOLUTIONARY SIGNIFICANCE OF SEXUAL REPRODUCTION. IV. EFFECT OF NEIGHBOR RELATEDNESS AND APHID INFESTATION ON SEEDLING PERFORMANCE

To investigate the effect of neighbor relatedness in seedling populations, propagules of six Anthoxanthum odoratum parents, produced from a reciprocal diallel cross, were planted into the parental source population, a mown field. The propagules were either surrounded by four unrelated neighbors in a 1 cm square, four sibling neighbors, or no neighbors. About 45% of the emerging seedlings were attacked by aphids (Schizaphis graminum). Aphid infestation significantly reduced seedling survivorship and did not differ with parental genotype or neighbor relatedness; plants without neighbors had a slightly higher infestation risk. Plants without neighbors had lower survivorship than those surrounded by unrelated neighbors although this difference was only significant for plants infested by aphids. When infested by aphids, plants surrounded by siblings had lower survivorship than plants surrounded by non-relatives, suggesting the operation of minority advantage. These results are consistent with the frequency-dependent selection hypothesis for the evolution and maintenance of sexual reproduction.     

Positive density dependence in seedlings of the neotropical tree species Garcinia macrophylla and Xylopia micans

Question: The Janzen‐Connell hypothesis predicts that herbivores and pathogens prevent seedlings from establishing in dense patches near adult conspecifics. Although many studies have investigated the Janzen‐Connell hypothesis, the environmental context – local or regional – in which juveniles establish is often overlooked. The objectives of this study were: (1) to evaluate Janzen‐Connell effects in contrasting environments, and (2) to incorporate microsite variation into the study of this hypothesis. Location: Pacaya‐Samiria Reserve, Peru. Methods: I assessed seedling performance of two tree species, Garcinia macrophylla and Xylopia micans, during one growing season. In an observational study, mortality and growth rates were regressed against distance to the nearest adult conspecific, conspecific seedling density, heterospecific plant density, and several abiotic variables in upland and floodplain forests. Field and shadehouse experiments were used to isolate distance‐ and density‐dependent effects. Results: Contrary to predictions, seedling survivorship increased in the presence of conspecific seedlings (Garcinia) and heterospecific understory plants (Garcinia and Xylopia) in the observational study. Survivorship in the field experiment, however, was unaffected by conspecific seedling density or adult proximity. In the shadehouse, Garcinia growth rates were highest in floodplain soils collected near adult conspecifics, but mortality was unrelated to the soil's habitat or proximity to an adult. Conclusions: The positive density dependence found in this study could have been produced by: (1) environmental factors that increase both density and survivorship, or (2) interspecific facilitation, if heterospecifics reduce herbivore or pathogen pressure on the focal species. Such interactions could help explain species coexistence in tropical forests.     

Comparative habitat susceptibility to invasion by Chinese ash (<Emphasis Type="Italic">Fraxinus chinensis</Emphasis>: Oleaceae) in a tropical Andean landscape

Chinese ash (Fraxinus chinensis) is an exotic tree species that has been used in non-commercial monospecific plantations for revegetation programs in the Central Andes of Colombia. At the Otún river watershed, these plantations occur in patches intermixed with old pastures, oak forests, and successional forests. In this heterogeneous landscape, the ash has been able to invade some of its surrounding habitats. This study evaluates the invasion patterns of ash to each of these habitats and experimentally quantifies seed and seedling survivorship and seedling growth as three processes that could determine ash establishment. Of the four habitats examined, old pastures were the most vulnerable to invasion, followed by oak plantations, and successional forest. Ash plantations exhibited recruitment levels intermediate between pastures and oak plantations. Abandoned pastures showed the highest seed germination, seedling survivorship, and seedling growth. In the ash plantations, recruitment seemed negatively affected by the low number of germinated seeds, high mortality of seedlings, and low growth. Invasion in oak plantations was constrained by high seed mortality and burial of seedlings by leaf litter, although the seedlings that did survive grew fast and produced a high number of leaves. Within the successional forest, Chinese ash seeds germinated but establishment was constrained by a reduced seedling survivorship and low growth. This research offers new evidence of how different processes affecting the establishment of an invader may differ among landscape elements, and it also yields important information for the management and control of ash in these Andean landscapes.     

An experimental test of limits to tree establishment in Arctic tundra

1 Five treeline species had low seed germination rates and low survivorship and growth of seedlings when transplanted into Alaskan tundra. Seed germination of all species increased with experimental warming, suggesting that the present treeline may in part result from unsuccessful recruitment under cold conditions.
2 Growth, biomass and survivorship of seedlings of treeline species transplanted into tundra were largely unaffected by experimental warming. However, transplanted seedlings of three species ( Betula papyrifera , Picea glauca and Populus tremuloides ) grew more when below‐ground competition with the extant community was reduced. All three measures of transplant performance were greater in shrub tundra than in the less productive tussock or heath tundra. Establishment of trees in tundra may thus be prevented by low resource availability and competition.
3 Two species ( Alnus crispa and Populus balsamifera ) had low seed germination and survivorship of germinated seeds; transplants of these species did not respond to the manipulations and lost biomass following transplanting into tundra. Isolated populations of these two species north of the present treeline in arctic Alaska probably became established during mid‐Holocene warming rather than in recent times.
4 Of all the species studied here, Picea glauca was the most likely to invade intact upland tundra. Its seeds had the highest germination rates and it was the only species whose seedlings survived subsequently. Furthermore, transplanted seedlings of Picea glauca had relatively high survivorship and positive growth in tundra, especially in treatments that increased air temperature or nutrient availability, two factors likely to increase with climate warming.