Differences in nitrogen use strategies between native and exotic tree species: predicting impacts on invaded ecosystems

Background and aim

Exotic plant species can alter the nitrogen cycle in invaded ecosystems. We assess the differences in nitrogen use strategies and litter production and dynamics among three native riparian trees (Fraxinus angustifolia, Populus alba and Ulmus minor) and three co-occurring exotics (Ailanthus altissima, Robinia pseudoacacia and Ulmus pumila), currently spreading throughout river banks in inner Spain. We aim to predict the ecological consequences of a replacement of the natives by the exotics.

Methods

We compared the leaf lifespan, nitrogen resorption efficiency in leaves, nitrogen mean residence time, amount and timing of litter production and amount of nitrogen returned to soils between these native and exotic species.

Results

We found differences among species in all the variables measured, but not between native or exotic origins. Species were ranked from the most to the least conservative nitrogen use strategy as follows: U. pumila was the most conservative species, followed by the three natives (with an intermediate strategy), A. altissima and finally by the nitrogen-fixer R. pseudoacacia. The studied exotic species would produce contrasting impacts on the nitrogen cycle upon invasion.

Conclusions

On the basis of our results, we predict little impacts on the nitrogen cycle if U. pumila dominates the landscape. Despite being nitrogen-fixer R. pseudocacia would not increase soil nitrogen availability in the study area due to its low litter production and litter decomposition rates. In contrast, we predict an increase in nitrogen availability of soils upon A. altissima invasion, as this species produces a high amount of nitrogen rich and labile litter. This study offers a striking example of the contingencies involved in predicting the ecosystem impacts of exotic plant invasion.     

Effects of exotic and native tree leaf litter on soil properties of two contrasting sites in the Iberian Peninsula

Aims

We assessed the effects of native and exotic tree leaf litter on soil properties in two contrasting scenarios. The native Quercus robur and Pinus pinaster tree species coexist with the aliens Eucalyptus globulus and Acacia dealbata in acid soils of NW Spain. The native trees Fraxinus angustifolia and Ulmus minor coexist with the aliens Ailanthus altissima, Robinia pseudoacacia and Ulmus pumila in eutrophic basic riparian soils in Central Spain.

Methods

Four plastic trays per species were filled with homogenized top-soil of the site and covered with leaf litter. Before and after 9?months of incubation, litter mass, soil pH, organic matter, mineral and total N were measured. Available mineral N (NO ₃ ^? -N and NH ₄ ⁺ -N) was assessed every 2?months.

Results

Soil biological activity was higher in the basic than in the acid soil. Litter of the exotic trees tended to decompose less than litter of native species, probably due to the presence of secondary metabolites in the former. Soil pH, mineral and total N responded differently to different litter types, irrespective of their exotic or native origin (acid soil), or was similar across litter treatments (basic riparian soil). The similar response of the basic soil to the addition of different litter types may be due to the low contrast of litter quality between the species. E. globulus litter inhibitied soil microbial activity much more than the rest of the studied litter types, leading to a drastic impoverishment of N in soils.

Conclusion

Litter of exotic N-fixing trees (A. dealbata and R. pseudoacacia) did not increase soil N pools because of the inhibition of microbial activity by secondary compounds. Therefore, secondary metabolites of the litter played a major role explaining exotic litter impact on soil properties.     

Comparison of leaf decomposition and macroinvertebrate colonization between exotic and native trees in a freshwater ecosystem

One of the most important sources of energy in aquatic ecosystems is the allochthonous input of detritus. Replacement of native tree species by exotic ones affects the quality of detritus entering freshwater ecosystems. This replacement can alter nutrient cycles and community structure in aquatic ecosystems. The aims of our study were (1) to compare leaf litter decomposition of two widely distributed exotic species (Ailanthus altissima and Robinia pseudoacacia) with the native species they coexist with (Ulmus minor and Fraxinus angustifolia), and (2) to compare macroinvertebrate colonization among litters of the invasive and native species. Litter bags of the four tree species were placed in the water and collected every 2, 25, 39, 71, and 95 days in a lentic ecosystem. Additionally, the macroinvertebrate community on litter bags was monitored after 25, 39, and 95 days. Several leaf chemistry traits were measured at the beginning (% lignin; lignin:N, C:N, LMA) and during the study (leaf total nitrogen). We detected variable rates of decomposition among species (k values of 0.009, 0.008, 0.008, and 0.005 for F. angustifolia, U. minor, A. altissima and R. pseudoacacia, respectively), but we did not detect an effect of litter source (from native/exotic). In spite of its low decay, the highest leaf nitrogen was found in R. pseudoacacia litter. Macroinvertebrate communities colonizing litter bags were similar across species. Most of them were collectors (i.e., they feed on fine particulate organic matter), suggesting that leaf material of either invasive or native trees was used as substrate both for the animals and for the organic matter they feed on. Our results suggest that the replacement of the native Fraxinus by Robinia would imply a reduction in the rate of leaf processing and also a slower release of leaf nitrogen to water.     

Allelopathic potentials of exotic invasive and native trees over coexisting understory species: the soil as modulator

Ailanthus altissima and Robinia pseudoacacia are two aggressive exotic tree species invading riparian ecosystems in Central Spain. We explored their allelopathic potentials as a possible mechanism explaining their success in these ecosystems. Specifically, we aimed (1) to compare the phytotoxic effects of the exotic and native (Fraxinus angustifolia and Populus alba) trees on the fitness of several understory plants coexisting in riparian ecosystems, and (2) to assess the capacity of the riparian soil to modulate the phytotoxic effects. In laboratory bioassays, aqueous leaf litter extracts from the donor tree species at field-realistic concentrations were tested on different fitness indicators of 13 understory target species, using germination paper and soil as substrates in petri dishes. Using germination paper, we found species-specific effects between donor and target species, but the phytotoxicity of the exotic trees as a group was not greater than that of the natives. Nevertheless, the exotic R. pseudoacacia was the most effective donor species reducing the radicle growth of the target species. Over riparian soil substrate, the aqueous leaf litter extracts did not produce any phytotoxic effect on the target species, except in one case. Altogether, our results highlight the importance of using both a native control when assessing the phytotoxicity of nonnative plants and also the natural soil in the modulation of phytotoxic effects. Ignoring both factors in laboratory bioassays would have led to the overestimation of the phytotoxicity of the exotic species as a mechanism contributing to their invasion success.     

Effects of exotic invasive trees on nitrogen cycling: a case study in Central Spain

We assess the hypothesis that rates of nitrogen transformations in the soil are altered upon replacement of native by exotic trees, differing in litter properties. Ailanthus altissima and Robinia pseudoacacia, two common exotic trees naturalized in the Iberian Peninsula, were compared with the native trees Ulmus minor and Fraxinus angustifolia, respectively. Naturally senesced leaves of each species were collected and C:N ratio, N and lignin content assessed. We prepared 64 litter bags per species and left them to decompose, below the canopy of the same species and below the canopy of the paired species. Dry mass, N concentration and N pool of the remaining litter were assessed after 5 and 7 months. Soil samples were collected three times during the experiment to assess soil moisture, organic matter, pH, potential mineralization rates and mineral N pools. Mineral N availability was assessed three times in the field by using ion-exchange resin-impregnated membranes. Ailanthus litter decomposed faster than Ulmus litter, probably due to the higher toughness of the latter. In spite of its high N content, Robinia litter decomposed slower than Fraxinus one, probably due to its high lignin content. In both cases, litter decomposition was faster below the exotic than the native canopies. The release of N per unit of initial litter mass was higher under both invaded situations (Ailanthus below Ailanthus and Robinia below Robinia) than under the native ones. However, soils collected below native and exotic trees neither differed in potential N mineralization rate nor in mineral N. This may be attributed to a quick plant uptake of released N and/or to a high organic matter accumulation in the soil previous to invasion that can exert a tighter control on soil N transformations than the current exotic litter.     

A last stand in the Po valley: genetic structure and gene flow patterns in Ulmus minor and U. pumila

Background and Aims Ulmus minor has been severely affected by Dutch elm disease (DED). The introduction into Europe of the exotic Ulmus pumila, highly tolerant to DED, has resulted in it widely replacing native U. minor populations. Morphological and genetic evidence of hybridization has been reported, and thus there is a need for assessment of interspecific gene flow patterns in natural populations. This work therefore aimed at studying pollen gene flow in a remnant U. minor stand surrounded by trees of both species scattered across an agricultural landscape.Methods All trees from a small natural stand (350 in number) and the surrounding agricultural area within a 5-km radius (89) were genotyped at six microsatellite loci. Trees were morphologically characterized as U. minor, U. pumila or intermediate phenotypes, and morphological identification was compared with Bayesian clustering of genotypes. For paternity analysis, seeds were collected in two consecutive years from 20 and 28 mother trees. Maximum likelihood paternity assignment was used to elucidate intra- and interspecific gene flow patterns.Key Results Genetic structure analyses indicated the presence of two genetic clusters only partially matching the morphological identification. The paternity analysis results were consistent between the two consecutive years of sampling and showed high pollen immigration rates (∼0·80) and mean pollination distances (∼3 km), and a skewed distribution of reproductive success. Few intercluster pollinations and putative hybrid individuals were found.Conclusions Pollen gene flow is not impeded in the fragmented agricultural landscape investigated. High pollen immigration and extensive pollen dispersal distances are probably counteracting the potential loss of genetic variation caused by isolation. Some evidence was also found that U. minor and U. pumila can hybridize when in sympatry. Although hybridization might have beneficial effects on both species, remnant U. minor populations represent a valuable source of genetic diversity that needs to be preserved.     

黄土高原不同树种枯落叶混合分解效应

混交林中不同树种枯落物混合分解是否产生促进或抑制作用是评价种间关系和混交适宜性的重要依据之一。以黄土高原主要树种为对象,通过室内枯落叶混合分解模拟试验,结果表明:(1)沙棘、白榆、柠条和小叶杨枯落叶分解最快(周转期1 a左右),其次为旱柳、侧柏和白桦枯落叶(周转期略大于1 a),紫穗槐、辽东栎和刺槐枯落叶分解稍慢(周转期1.5 a左右),而樟子松、落叶松和油松枯落叶分解最慢(周转期略大于2 a)。(2)对于针叶树,与油松枯落叶混合,存在明显促进分解作用的是侧柏、落叶松,其次是白桦、沙棘和刺槐;与樟子松枯落叶混合,存在明显促进分解作用的是落叶松、侧柏、沙棘、白榆,其次是柠条、紫穗槐和小叶杨,而存在明显抑制分解作用的是刺槐,其次是白桦和辽东栎;与落叶松枯落叶混合,存在较明显促进分解作用的是白榆、白桦和辽东栎,存在较明显抑制作用的是刺槐;紫穗槐与侧柏枯落叶混合存在较明显的抑制分解作用。(3)对于阔叶树,与小叶杨枯落叶混合,存在较明显促进分解作用的是紫穗槐,其次是辽东栎和刺槐;与刺槐枯落叶混合,存在较明显促进分解作用的是白榆和沙棘,存在明显抑制分解作用的是柠条,其次是辽东栎和白桦;与白桦枯落叶混合,存在较明显促进分解作用的是辽东栎和紫穗槐,存在较明显抑制分解作用的是柠条;白榆与辽东栎、旱柳枯落叶混合均存在较明显的促进分解作用,而白榆与柠条枯落叶混合存在较明显的抑制分解作用;紫穗槐与旱柳、沙棘枯落叶混合均存在较明显的促进分解作用,而紫穗槐与柠条枯落叶混合有较明显的抑制作用。     

Hybridization and introgression between the exotic Siberian elm, Ulmus pumila, and the native Field elm, U. minor, in Italy

In response to the first Dutch elm disease (DED) pandemic, Siberian elm, Ulmus pumila, was planted to replace the native elm, U. minor, in Italy. The potential for hybridization between these two species is high and repeated hybridization could result in the genetic swamping of the native species and facilitate the evolution of invasiveness in the introduced species. We used genetic markers to examine the extent of hybridization between these two species and to determine the pattern of introgression. We quantified and compared the level of genetic diversity between the hybrids and the two parental species. Hybrids between U. pumila and U. minor were common. The pattern of introgression was not as strongly biased towards U. pumila as was previously observed for hybrids between U. rubra and U. pumila in the United States. The levels of heterozygosity were similar between U. minor and the hybrids and both groups had higher levels of heterozygosity relative to U. pumila. The programs Structure and NewHybrids indicated the presence of first- (F₁) and second- generation (F₂) hybrids and of backcrosses in the hybrid population. The presence of healthy DED resistant U. minor individuals combined with the self-compatibility of U. minor could help explain the presence of F₂ individuals in Italy. The presence of F₂ individuals, where most of the variability present in the hybrids will be released, could facilitate rapid evolution and the potential evolution of invasiveness of U. pumila in Italy.     

Intra- and interspecific hybridization in invasive Siberian elm

Hybridization creates unique allele combinations which can facilitate the evolution of invasiveness. Frequent interspecific hybridization between the Siberian elm, Ulmus pumila, and native elm species has been detected in the Midwestern United States, Italy and Spain. However, Ulmus pumila also occurs in the western United States and Argentina, regions where no native elm species capable of hybridizing with it occurs. We examined whether inter- or intraspecific hybridization could be detected in these regions. Nuclear markers and the program STRUCTURE helped detect interspecific hybridization and determine the population genetic structure in both the native and the two non-native ranges. Chloroplast markers identified sources of introduction into these two non-native ranges. No significant interspecific hybridization was detected between U. pumila and U. rubra in the western United States or between U. pumila and U. minor in Argentina and vice versa. However, the genetic findings supported the presence of intraspecific hybridization and high levels of genetic diversity in both non-native ranges. The evidence presented for intraspecific hybridization in the current study, combined with reports of interspecific hybridization from previous studies, identifies elm as a genus where both inter- and intraspecific hybridization may occur and help maintain high levels of genetic diversity potentially associated with invasiveness.     

Analysis of interactions between the invasive tree-of-heaven (<Emphasis Type="Italic">Ailanthus altissima</Emphasis>) and the native black locust (<Emphasis Type="Italic">Robinia pseudoacacia</Emphasis>)

Invasive exotic plants can persist and successfully spread within ecosystems and negatively affect the recruitment of native species. The exotic invasive Ailanthus altissima and the native Robinia pseudoacacia are frequently found in disturbed sites and exhibit similar growth and reproductive characteristics, yet each has distinct functional roles such as allelopathy and nitrogen fixation, respectively. A four-month full additive series in the greenhouse was used to analyze the intraspecific and interspecific interference between these two species. In the greenhouse experiment, the inverse of the mean total biomass (g) response per plant for each species was regressed on the density of each species and revealed that the performance of the plants was significantly reduced by interspecific interference and not by intraspecific interference (p < 0.05). Other biomass traits such as root dry weight, shoot dry weight, stem dry weight, and leaf dry weight were also negatively affected by interspecific interference. Competition indices such as Relative Yield Total and Relative Crowding Coefficient suggested that A. altissima was the better competitor in mixed plantings. Ailanthus altissima consistently produced a larger above ground and below ground relative yield while R. pseudoacacia generated a larger aboveground relative yield in high density mixed species pots.     

Girdling of young Robinia pseudoacacia trees on the Tama River terrace,central Japan

Robinia pseudoacacia L. has been widely planted worldwide for a variety of purposes, but it is an aggressive invader in many regions. To control invasion by the species, we examined the effectiveness of girdling young trees to ground level to kill trees and preclude sprout development through comparison among girdling, clearcut and control treatments on the Tama river terrace, Tokyo. More than 99% of R. pseudoacacia trees were killed by girdling. The number of sprouts that developed from roots and stumps after girdling was distinctly reduced compared with that observed after clearcutting. We also conducted girdling of young trees of Ailanthus altissima (Mill.) Swingle, which shows similar developmental and ecological characteristics to R. pseudoacacia, and the two species are often sympatric. More than 98% of A. altissima trees were killed, but a higher number of sprouts developed from A. altissima stumps compared with that from R. pseudoacacia stumps. These results indicate that girdling young trees to ground level is an effective method of controlling R. pseudoacacia by killing the stem and precluding sprout development. The girdling method is simple to apply and imposes a low burden on the surrounding environment.

     

Interactions between seedlings of the invasive tree Ailanthus altissima and the native tree Robinia pseudoacacia under low nutrient conditions

The allelopathic effect of A. altissima and the nitrogen fixing ability of R. pseudoacacia make the interaction between these two species important to community dynamics. A replacement series greenhouse experiment was used to investigate the type of interaction between seedlings of A. altissima and R. pseudoacacia at high and low soil nutrition states. Also, seeds of A. altissima from its native (China) and invasive ranges (USA) were used to compare the effect of the different seed sources on the interaction with R. pseudoacacia. Robinia pseudoacacia was the better competitor. The presence of A. altissima significantly inhibited nodulation of R. pseudoacacia roots. In summary, in low nutrient early succession sites, seedlings of R. pseudoacacia and A. altissima will coexist and R. pseudoacacia will be the better competitor. However, A. altissima can increase its competitive ability by inhibiting nitrogen fixation by R. pseudoacacia. Differences between the competitive ability of the Chinese and US seed lots of A. altissima did not support the rapid evolution theory for invasive species success.     

Exotic species as modifiers of ecosystem processes: Litter decomposition in native and invaded secondary forests of NW Argentina

Invasions of exotic tree species can cause profound changes in community composition and structure, and may even cause legacy effect on nutrient cycling via litter production. In this study, we compared leaf litter decomposition of two invasive exotic trees (Ligustrum lucidum and Morus sp.) and two dominant native trees (Cinnamomum porphyria and Cupania vernalis) in native and invaded (Ligustrum-dominated) forest stands in NW Argentina. We measured leaf attributes and environmental characteristics in invaded and native stands to isolate the effects of litter quality and habitat characteristics. Species differed in their decomposition rates and, as predicted by the different species colonization status (pioneer vs. late successional), exotic species decayed more rapidly than native ones. Invasion by L. lucidum modified environmental attributes by reducing soil humidity. Decomposition constants (k) tended to be slightly lower (−5%) for all species in invaded stands. High SLA, low tensile strength, and low C:N of Morus sp. distinguish this species from the native ones and explain its higher decomposition rate. Contrary to our expectations, L. lucidum leaf attributes were similar to those of native species. Decomposition rates also differed between the two exotic species (35% higher in Morus sp.), presumably due to leaf attributes and colonization status. Given the high decomposition rate of L. lucidum litter (more than 6 times that of natives) we expect an acceleration of nutrient circulation at ecosystem level in Ligustrum-dominated stands. This may occur in spite of the modified environmental conditions that are associated with L. lucidum invasion.     

Allelopathy: a tool for weed management in forest restoration

Forest restoration uses active management to re-establish natural forest habitat after disturbance. However, competition from early successional species, often aggressively invasive exotic plant species, can inhibit tree establishment and forest regeneration. Ideally, restoration ecologists can plant native tree species that not only establish and grow rapidly, but also suppress exotic competitors. Allelopathy may be a key mechanism by which some native trees could reduce the abundance and impact of exotic species. Allelopathy is a recognized tool for weed management in agriculture and agroforestry, but few studies have considered how allelopathic interactions may aid restoration. Here we introduce the “Homeland Security” hypothesis, which posits that some naïve exotic species may be particularly sensitive to allelochemicals produced by native species, providing a tool to reduce the growth and impacts of invasive exotic species on reforestation. This article explores how exploiting allelopathy in native species could improve restoration success and the re-establishment of natural successional dynamics. We review the evidence for allelopathy in agroforestry systems, and consider its relevance for reforestation. We then illustrate the potential for this approach with a case study of tropical forest restoration in Panama. C₄ grasses heavily invade deforested areas in the Panama Canal watershed, especially Saccharum spontaneum L. We measured the effect of leaf litter from 17 potential restoration tree species on the growth of invasive C₄ grasses. We found that leaf litter from legume trees had a greater inhibitory effect on performance of S. spontaneum than did litter from non-legume trees. However, allelopathic effects varied greatly among species within tree functional groups. Further evaluation of intra- and inter-specific interactions will help to improve our selection of restoration species.     

Vessel length and conductivity of Ulmus branches: ontogenetic changes and relation to resistance to Dutch elm disease

Changes in age of the hydraulic architecture of Ulmus minor and U. minor × U. pumila juvenile wood were studied and related to tolerance to Dutch elm disease (DED). The xylem vessel dimensions and the conductivity to air of 2- to 7-year-old branches were analyzed and quantified. No obvious differences in vessel length distribution and conductivity were found to explain differences in DED tolerance among the U. minor clones, or, at the taxon level, the higher DED tolerance of U. minor × U. pumila. Among the U. minor clones, the more susceptible one had wider vessels and a higher maximum vessel diameter than the more tolerant clone. Relations between vessel lengths, vessel diameters and branch sizes were highly significant, and varied between taxa. The diameter and length of vessels increased with age, and average values stabilized 1–2 years earlier for U. minor than for U. minor × pumila. Mean maximum vessel length was significantly higher in U. minor and increased more with age and maximum vessel diameter than in U. minor × pumila. With each 0.2 m increase in height up the stem, conductivities for U. minor and U. minor × U. pumila decreased by 59 and 50 %, respectively, probably due to shortening of the vessels. The implications of xylem structure for the means of pathogen movement and resistance to DED are discussed.     

Arthropod diversity of exotic vs. native Robinia species in northern Arizona

1 Arthropods were collected on native locust, Robinia neomexicana A. Gray, and exotic Robinia pseudoacacia L. in northern Arizona over a 2‐year period to determine the number of arthropod species and number of individuals present. 2 More arthropod species were found on the native (251) than on the exotic Robinia (174). 3 Greater species diversity was likewise found on the native than the exotic. The five most numerous insects collected each year accounted for 81% to 91% of the total number collected on the exotic and native Robinia in 1997 and 1998. Only 12 species occurred on both the native and exotic Robinia in both years. 4 These findings are discussed in the context of using exotic trees in plantations and ecological theory regarding rates of arthropod species accumulation on exotic hosts.     

Trees in the desert: Reproduction and genetic structure of fragmented Ulmus pumila forests in Mongolian drylands

The potential natural distribution of deciduous forests in Central Asia is the subject of ongoing discussions. Ulmus pumila (Siberian elm) is the only tree species occurring in southern and south-eastern Mongolia. In the semi-arid Mongolian Gobi, the species is restricted to ravines and beds of semi-temporary rivers. Compared to zonal occurrences in moister northern Mongolia, elm trees in the Gobi were found to be larger, in spite of their slower growth. Recruitment was very rare in the field although germination studies revealed that seeds were viable, survived osmotic stress, and were tolerant of repeated cycles of moistening and drying. Thus, they should be capable of germination in episodically flooded river beds. Fingerprinting revealed that clonal growth is of negligible importance in the Gobi as almost all U. pumila individuals studied constituted separate genets. Given that many trees were <100 years old and must have become established under current climatic conditions, we infer that the current lack of recruitment is likely to be caused by grazing impact. Our data imply that Ulmus pumila could potentially be much more common in the drylands of southern Mongolia and northern China.     

Revegetation patterns and seedbank structure on abandoned pastures in northern Japan

To compare revegetation patterns between native and exotic species on abandoned pastures in northern Japan, we surveyed the vegetation structure and the seedbank using a flotation technique in 140 1 1 m plots. Previously introduced grasses such as Poa pratensis, Phleum pratense, and Dactylis glomerata were abundant 20 yr after the pasture abandonment, while Sasa senanensis, native shrub species, regenerated from propagation that had spread from the surrounding forests. S. senanensis shrublands and P. pratensis/P. pratense grasslands established on deep soils while D. glomerata grasslands established on shallow soils. Trees rarely became established on abandoned pastures. The seed density in the seedbank, representing 19 species, ranged from 542 to 2957 seeds/m². The dominant species in the vegetation (P. pratensis, Trifolium repens, and Rumex acetosella) were also common in the seedbank, whereas Cerastium holosteoides var. angustifolium, Chenopodium album var. centrorubrum, and Erigeron canadensis were widespread in the seedbank but did not occur in the extant vegetation. S. senanensis regenerated by vegetative propagation, and P. pratensis and P. pratense developed a seedbank. We concluded that for native species, particularly S. senanensis, vegetative reproduction has an important role on revegetation rather than regeneration from the seedbank, and the dwarf bamboo may be a keystone species in the ecosystems.     

Dispersal and recruitment limitation in native versus exotic tree species: life‐history strategies and Janzen‐Connell effects

Life‐history traits of invasive exotic plants are typically considered to be exceptional vis‐à‐vis native species. In particular, hyper‐fecundity and long range dispersal are regarded as invasive traits, but direct comparisons with native species are needed to identify the life‐history stages behind invasiveness. Until recently, this task was particularly problematic in forests as tree fecundity and dispersal were difficult to characterize in closed stands. We used inverse modelling to parameterize fecundity, seed dispersal and seedling dispersion functions for two exotic and eight native tree species in closed‐canopy forests in Connecticut, USA. Interannual variation in seed production was dramatic for all species, with complete seed crop failures in at least one year for six native species. However, the average per capita seed production of the exotic Ailanthus altissima was extraordinary: > 40 times higher than the next highest species. Seed production of the shade tolerant exotic Acer platanoides was average, but much higher than the native shade tolerant species, and the density of its established seedlings (≥ 3 years) was higher than any other species. Overall, the data supported a model in which adults of native and exotic species must reach a minimum size before seed production occurred. Once reached, the relationship between tree diameter and seed production was fairly flat for seven species, including both exotics. Seed dispersal was highly localized and usually showed a steep decline with increasing distance from parent trees: only Ailanthus altissima and Fraxinus americana had mean dispersal distances > 10 m. Janzen‐Connell patterns were clearly evident for both native and exotic species, as the mode and mean dispersion distance of seedlings were further from potential parent trees than seeds. The comparable intensity of Janzen‐Connell effects between native and exotic species suggests that the enemy escape hypothesis alone cannot explain the invasiveness of these exotics. Our study confirms the general importance of colonization processes in invasions, yet demonstrates how invasiveness can occur via divergent colonization strategies. Dispersal limitation of Acer platanoides and recruitment limitation of Ailanthus altissima will likely constitute some limit on their invasiveness in closed‐canopy forests.     

黄土高原樟子松和落叶松与其他树种枯落叶混合分解对土壤的影响

通过采集树木枯落叶与土壤进行室内混合分解培养试验,研究了黄土高原常见的樟子松和落叶松与其他树种枯落叶混合分解对土壤性质的影响及存在的相互作用,从而为不同树木种间关系的探索和该地区人工纯林的混交改造提供科学指导。结果表明:12种枯落叶单一分解均明显提高了土壤脲酶(54%—110%)、脱氢酶(85%—288%)和磷酸酶(81%—301%)活性以及有机质(29%—55%)和碱解N(12%—49%)含量,但对土壤速效P含量和CEC的影响存在较大差异。综合而言,樟子松分别与白桦、刺槐、白榆、柠条和落叶松枯落叶混合分解在对土壤性质的影响中存在相互促进作用,而分别与小叶杨、沙棘、紫穗槐、侧柏和辽东栎枯落叶混合分解在对土壤性质的影响中存在相互抑制作用;落叶松分别与刺槐、白桦、小叶杨和紫穗槐枯落叶混合分解在对土壤性质的影响中存在相互促进作用,而分别与柠条、侧柏、辽东栎、沙棘、油松和白榆枯落叶混合分解在对土壤性质的影响中存在相互抑制作用。