清除优势草本植物对西双版纳飞机草-马唐群落中木本幼苗的影响

在农业弃耕地上通常会出现由草本植物占优势的演替早期群落。为研究优势草本植物对群落中木本幼苗及群落微环境的影响以及外来入侵植物和本地植物对木本幼苗的影响有何差异,在西双版纳地区选取由外来入侵植物飞机草和本地植物马唐共同占优势的演替早期群落,实施物种清除实验,观测样地中木本幼苗的高增长及死亡补充情况,同时对群落微环境进行观测。结果显示,清除优势种显著提高了木本幼苗的高增长,降低了其死亡率,并且使新增幼苗数量有所提高。清除优势种对0-50cm幼苗高增长和死亡率的影响均大于50-100(或200)cm幼苗。清除处理显著提高了样地的冠层下可见天空比例,但对土壤含水量和土壤养分的影响并不显著。飞机草和马唐均能形成浓密的冠层,通过对光的竞争抑制木本幼苗的生长,且这两者的抑制作用没有显著差异。该地区次生林中常见的先锋树种在研究样地内均有幼苗存在,但其生长却受到优势草本植物的强烈抑制。因此,对演替早期群落中草本植物的控制与管理应同时注意外来入侵物种与本地杂草。     

Rain forest expansion mediated by successional processes in vegetation thickets in the Western Ghats of India

Aim The objective of this study was to document succession from grassland thickets to rain forest, and to provide evidence for their potential as restoration tools. Location The Linganamakki region (State of Karnataka) of the Central Western Ghats of India. Method We selected thirty vegetation thickets ranging from 4 to 439 m² in area in the vicinity of rain forest. The area of each small thicket was estimated as an oval using its maximum length and its maximum width. When the shape was irregular (mostly in large thickets) the limits of the thicket were mapped and the area calculated from the map. Plant species were identified, the number of individuals was estimated and their heights measured. Results There was a progression in the thickets from early to late successional species. Small thickets were characterized by ecotone species and savanna trees such as Catunaregam dumetorum. Savanna trees served as a nucleus for thicket formation. Colonizing species were mostly bird‐dispersed. As succession proceeded in larger thickets, the proportion of evergreen, late‐successional rain forest trees increased. The species composition of the large thickets differed depending on the species composition of reproductive adults in the nearby forested areas. The species within small thickets were also found in the large thickets. The nestedness in species composition suggested that species turnover was deterministic based on thicket size. Human disturbance (leaf and wood collection by the local populations) affected the species composition and the species–area relationship of thickets. Main conclusions Vegetation thickets are nodal centres for rain forest colonization within grasslands. They expand and replace savanna. Early successional bird‐dispersed species established around savanna trees followed by late‐successional rain forest trees dispersed from the nearby forest by birds. Restoration programmes that reproduce natural successional processes such as those observed in thickets will be more successful and less expensive than the methods currently being employed, where trees are individually planted in grassland. Wood harvesting is the only factor that prevents thicket growth and coalescence and hampers forest expansion.     

Intraspecific divergence and evolution of a life‐history trade‐off along a successional gradient in Hawaii's Metrosideros polymorpha

The importance of environmental gradients in the diversification of long‐lived tree species is poorly understood. Two morphologically distinct varieties of the endemic Hawaiian tree, 'ōhi'a lehua (Metrosideros polymorpha), are the canopy dominants at alternate extremes of a successional gradient formed by the recurring disturbance of lava flows on east Hawai'i Island. The maintenance of these varieties despite hybridization may be due to disruptive selection at either end of the successional gradient. To test this hypothesis, seeds from three, replicate monotypic stands of each variety on east Hawai'i Island were germinated and the resulting seedlings grown under four combinations of light and nitrogen levels in a greenhouse, and at early‐ and late‐successional field sites. Growth and survivorship measures revealed differential fitness of these varieties in high‐ and low‐light environments in the greenhouse with corresponding differential fitness in early‐ and late‐successional field sites. Unique light‐by‐nitrogen interaction effects on growth were observed in each variety, and only the late‐successional variety appeared to be nitrogen limited. These two varieties exhibit the classic plant life‐history trade‐off between fast growth in high light and high survivorship in shade, but notably within a single tree species. These findings strongly implicate a role for Hawaii's striking environmental heterogeneity in the emergence of at least two endemic forms of this woody genus.     

Nitrogen-fixers <Emphasis Type="Italic">Alnus</Emphasis> and <Emphasis Type="Italic">Lupinus</Emphasis> influence soil characteristics but not colonization by later successional species in primary succession on Mount St. Helens

Changes to the primary successional environment caused by colonizing plants that present symbiotic associations with nitrogen-fixing bacteria were investigated at two areas on Mount St. Helens. One area was occupied by alder (Alnus viridis) thickets and old lupine (Lupinus lepidus) patches and the other area by young lupine patches and pumice barrens. Alder thicket soils had higher levels for a few soil nutrients and had greater cover by other pioneer species as compared to old lupine patches. Many soil nutrients, including nitrogen and soil organic matter, were below detection limits in old lupine patches but not in alder thicket soils. Young lupine patch soils were generally not different from barren site soils but had greater cover by other pioneer species. Below detection nitrogen and soil organic matter levels also occurred in many barren soil samples but not in young lupine patch soils. Barren soils were moister than were the other sites. The apparent increase in soil fertility has not led to invasion by later successional species, perhaps due to dry conditions or to other inhibitory factors. Seedbanks, composed of early successional species, appear to be developing in these areas.     

Comparisons of Mycorrhizal Responsiveness with Field Soil and Commercial Inoculum for Six Native Montane Species and Bromus tectorum

Reestablishing native perennial plants and reducing invasive species are pivotal for many ecological restoration projects. The interactions among plant species, arbuscular mycorrhizal fungi, and soil P availability may be critical determinants of the success of native and non‐native plants in restoration and species invasions. Here we assessed mycorrhizal responsiveness for three late‐successional and three early‐successional plant species native to Rocky Mountain National Park and for the non‐native Downy brome, cheatgrass (Bromus tectorum L.) using field soil and commercial inoculum. Factorial greenhouse experiments were conducted to compare biomass of plant species with and without field soil and commercial inoculum treatments along a phosphorus (P) gradient, which ranged from ambient field levels to 12% of field levels, using dilutions of native soils. The two field soil inoculum treatments resulted in significant biomass differences for all species studied. Late‐successional species responded positively to field inoculum, whereas early‐successional species responded negatively. The two commercial inocula had low colonization rates (14 of 166 inoculated plants). The commercial inocula substrates had significant treatment effects on five of seven species included in the study in the apparent absence of mycorrhizal symbiosis. Soil P levels influenced mycorrhizal responsiveness in only one species, Smooth blue aster (Aster laevis L.). Our results show that, at least for the species studied here, locally collected field inoculum is the best choice for reestablishment of late‐successional native plant species.     

Species Trials of Short Rotation Woody Crops on Two Wastewater Application Sites in North Carolina,USA

Forty-two Populus spp. clones, Eucalyptus benthamii, and seven tree species native to North Carolina were evaluated for survival and height growth through the establishment phase at two municipal wastewater application sites. Groundwater was monitored at each site to determine if establishment of the species trials resulted in exceedances of nutrient mitigation requirements. At the Gibson Wastewater Treatment Facility, 26 Populus clones had 100 % survival, with mean height growths ranging between 152 to 260 cm, and basal diameters ranging between 11.4 and 28.8 mm. Green ash, planted in 2011 and 2012, had high survivorship (>95 %) with first year mean height growth of 30?±?28 cm (2012) and second year mean height growth of 101?±?52 cm (2011). Basal diameter for green ash was 33.3?±?12.6 mm. E. benthamii had moderate survivorship (>77 %) and first year mean height growth of 47?±?27 cm. At the Jacksonville Wastewater Treatment Facility, green ash and bald cypress had high survivorship (>96 %), first year mean height growths of 14?±?25 cm and 27?±?16 cm, and basal diameters of 13.1?±?3.9 mm and 11.6?±?4.8 mm, respectively. Survivorship for 12 Populus clones ranged from 50 and 94 % with mean first year height growths between 58 to 121 cm, and basal diameters between 6.8 and 12.5 mm. E. benthamii had low survivorship (43 %) with mean first year height growths of 17?±?17 cm and basal diameters of 12.0?±?7.7 mm. Groundwater concentrations of NO₃?+?NO₂ and N-NH₄ remained below regulatory requirements at both sites with one exceedance in February 2012 in Jacksonville, NC. The results show that some Populus clones are excellent candidates for woody biomass production on municipal wastewater application fields. Native green ash and bald cypress are also good candidates, but these trees may require longer rotations than Populus to achieve similar biomass yields.     

Identifying Native Vegetation for Reducing Exotic Species during the Restoration of Desert Ecosystems

There is currently much interest in restoration ecology in identifying native vegetation that can decrease the invasibility by exotic species of environments undergoing restoration. However, uncertainty remains about restoration's ability to limit exotic species, particularly in deserts where facilitative interactions between plants are prevalent. Using candidate native species for restoration in the Mojave Desert of the southwestern U.S.A., we experimentally assembled a range of plant communities from early successional forbs to late‐successional shrubs and assessed which vegetation types reduced the establishment of the priority invasive annuals Bromus rubens (red brome) and Schismus spp. (Mediterranean grass) in control and N‐enriched soils. Compared to early successional grass and shrub and late‐successional shrub communities, an early forb community best resisted invasion, reducing exotic species biomass by 88% (N added) and 97% (no N added) relative to controls (no native plants). In native species monocultures, Sphaeralcea ambigua (desert globemallow), an early successional forb, was the least invasible, reducing exotic biomass by 91%. However, the least‐invaded vegetation types did not reduce soil N or P relative to other vegetation types nor was native plant cover linked to invasibility, suggesting that other traits influenced native‐exotic species interactions. This study provides experimental field evidence that native vegetation types exist that may reduce exotic grass establishment in the Mojave Desert, and that these candidates for restoration are not necessarily late‐successional communities. More generally, results indicate the importance of careful native species selection when exotic species invasions must be constrained for restoration to be successful.     

Disturbance Type and Plant Successional Communities in Bahamian Dry Forests

Different disturbances in similar habitats can produce unique successional assemblages of plants. We collected plant species composition and cover data to investigate the effects of three common types of disturbances—fire, anthropogenic clearing (‘cleared’), and clearing followed by goat grazing (‘cleared‐and‐grazed’)—on early‐successional coppice (dry forest) community structure and development on Eleuthera, Bahamas. For each disturbance type, both the ground layer (<0.5 m height) and shrub layer (>0.5 m height) were sampled in eight patches (>1 ha) of varying age (1–28 yr) since large‐scale mature coppice disturbance. Overall, plant communities differed among disturbance types; several common species had significantly higher cover in the shrub layer of fire patches, and cleared‐and‐grazed patches exhibited higher woody ground cover. Total percent cover in the shrub layer increased in a similar linear fashion along the investigated chronosequence of each disturbance type; however, cover of the common tree species, Bursera simaruba, increased at a notably slower rate in cleared‐and‐grazed patches. The pattern of increase and subsequent decrease in cover of Lantana spp. and Zanthoxylum fagara in the shrub layer was characterized by longer persistence and higher covers, respectively, in cleared‐and‐grazed patches, which also exhibited low peak cover and fast decline of nonwoody ground cover. Our results suggest that goats may accelerate some aspects of succession (e.g., quickly removing nonwoody ground cover) and retard other aspects (e.g., inhibiting growth of tree species and maintaining early‐successional shrubs in the shrub layer). These effects may lead to different successional trajectories, and have important conservation implications.     

Successional trends in standing dead biomass in Mediterranean basin species

Question: Landscape models of fire occurrence in ecosystems assume that the time since the last fire determines vegetation flammability by enabling the accumulation of dead biomass. In this study we ask if Mediterranean basin shrublands respond to these models or, on the contrary, if initial successional stages in these ecosystems could be more flammable than later stages. Location: Mediterranean shrubland in the Valencia region, eastern Spain. Methods: Using different stages of vegetation development (5, 9, 14 and 26 years since the last fire), we first study the structural comiosition of the above‐ground biomass in 375 individuals of nine woody species. Then, we measure how the standing dead biomass varies during succession, taking into account the surface cover of each species and the quantity of total dead biomass accumulated in different successional stages (3, 9, 14 and 26 years since the last fire). Results: The largest amount of standing dead biomass at the plant community level is observed in the middle stages of the succession. Early successional species, such as Cistus spp., Ulex parviflorus and Pinus halepensis, have a higher percentage of standing dead biomass at earlier stages in the succession than species typical of later successional stages, e.g. Juniperus oxycedrus, Quercus coccifera and Quercus ilex. Conclusions: The results suggest that monotonic increase in fire hazard with increasing stand age is not necessarily the rule in Mediterranean basin shrublands, since early successional species may accumulate large amounts of standing dead biomass and thus promote fire at early successional stages.     

Predispersal seed predation on five Piper species in tropical rainforest

Absolute number of seeds lost to predispersal seed predators and proportion of total seeds lost per infructescence were compared among five Costa Rican Piper species of different annual fecundities. Mean seed number and mean seed size in the five species were negatively correlated. The impact of predation on these species was inversely related to the number of seeds they produced. The two early successional species had very high fecundities, a combination of many seeds per infructescence, many infructescences per plant, and, in one species, year-round reproduction. Although seed predators destroyed as many or more seeds of these early successional species than they did of the less fecund, late successional species, this loss accounted for a relatively minor proportion (9 and 12%) of the seeds of the early successional species. In contrast, late successional species produced fewer, larger seeds in a smaller number of infructescences and were not continually in fruit. One of these species, which produced intermediate numbers of intermediately sized seeds, lost 30% of the seeds in each infructescence on average. Seed predators destroyed a larger proportion (65 and 76%) of the seeds per infructescence in the two species with fewest seeds per infructescence. High levels of insect damage in these late successional species caused many of their infructescences to abort prematurely. Taken together these factors resulted in annual fecundities several orders of magnitude smaller in shade-tolerant Piper species than the annual fecundities of shade-intolerant, early successional species. Seedlings of the two early successional species were common in large gaps and other sunny clearings and seedlings of the species with 30% seed loss were occasional, whereas no seedlings were seen of the two species with the highest proportional seed loss, suggesting that seed predation on the latter species may limit seedling recruitment.     

Restoration of a Forest Understory After the Removal of an Invasive Shrub, Amur Honeysuckle (Lonicera maackii)

The recruitment of native seedlings is often reduced in areas where the invasive Amur honeysuckle (Lonicera maackii) is abundant. To address this recruitment problem, we evaluated the effectiveness of L. maackii eradication methods and restoration efforts using seedlings of six native tree species planted within eradication and unmanipulated (control) plots. Two eradication methods using glyphosate herbicide were evaluated: cut and paint and stem injection with an EZ‐Ject lance. Lonicera maackii density and biomass as well as microenvironmental characteristics were measured to study their effects on seedling growth and survivorship. Mean biomass of Amur honeysuckle was 361 ± 69 kg/ha, and density was 21,380 ± 3,171 plants/ha. Both eradication treatments were effective in killing L. maackii (≥ 94%). The injection treatment was most effective on large L. maackii individuals (>1.5 cm diameter), was 43% faster to apply than cutting and painting and less fatiguing for the operator, decreased operator exposure to herbicide, and minimized impact to nontarget vegetation. Deer browse tree protectors were used on half of the seedlings, but did not affect survivorship or growth. After 3 years, survival of native seedlings was significantly less where L. maackii was left intact (32 ± 3%) compared with the eradication plots (p < 0.002). Seedling survival was significantly different between cut (51 ± 3%) and injected (45 ± 3%) plots. Species had different final percent survival and rates of mortality. Species survival differed greatly by species (in descending order): Fraxinus pennsylvanica > Quercus muehlenbergii ≥ Prunus serotina≥ Juglans nigra > Cercis canadensis > Cornus florida. Survivorship and growth of native seedlings was affected by a severe first‐year drought and by site location. One site exhibited greater spring soil moisture, pH, percent open canopy, and had greater survivorship relative to the other site (55 ± 2 vs. 30 ± 2%). Overall, both L. maackii eradication methods were successful, but restorationists should be aware of the potential for differential survivorship of native seedlings depending on species identity and microenvironmental conditions.     

Survivorship of juvenile barnacles and mussels: spatial dependence and the origin of alternative communities

Mussels, barnacles, and rockweeds often form a distinct mosaic of patches on rocky intertidal shores, and it has been suggested that these communities may represent alternative community states. One way that alternative community states can arise is if early successional events are scale-dependent, but it is not known if juvenile survivorships of mussels and barnacles are, in fact, scale-dependent. Scale-dependence of barnacles (Semibalanus balanoides (L.)) and mussels (Mytilus edulis L.) was tested in the Gulf of Maine, USA. In winter 1997, clearings of 1, 2, 4 and 8 m in diameter and uncleared controls were made in stands of the rockweed Ascophyllum nodosum (L.) Le Jolis at 12 sites spread evenly across four bays on Swan's Island, ME. Summer and fall-winter survivorship of barnacles, which recruited in spring 1997, were estimated by tracking the 1997 cohort until late winter 1998. Survivorship of mussels was estimated from following the fate of transplanted juveniles over 4 days in late August 1997. Both barnacles and mussels showed better survival in 4 and 8 m clearings than in small clearings and controls. There was also significant variation in survivorship among sites. Densities of gastropods in the clearings did not reflect survivorship patterns of barnacles and mussels. Barnacle survivorship increased in fall and winter, and in large clearings was comparable to survivorship in barnacle-dominated habitats. Mussel survivorship was low in all clearing sizes suggesting that mussel beds develop slowly.     

Changes in woody vegetation abundance and diversity after natural disturbances causing different levels of mortality

Questions: How does woody vegetation abundance and diversity differ after natural disturbances causing different levels of mortality? Location: Abies balsamea–Betula papyrifera boreal mixed‐wood stands of southeast Quebec, Canada. Methods: Woody vegetation abundance and diversity were quantified and compared among three disturbance‐caused mortality classes, canopy gap, moderate‐severity disturbances, and catastrophic fire, using redundancy analysis, a constrained linear ordination technique, and diversity indices. Results: Substantial changes in canopy tree species abundance and diversity only occurred after catastrophic fire. Shade‐tolerant, late‐successional conifer species remained dominant after canopy gap and moderate‐severity disturbances, whereas shade‐intolerant, early‐successional colonizers dominated canopy tree regeneration after catastrophic fire. Density and diversity of mid‐tolerant and shade‐intolerant understory tree and shrub species increased as the impact of disturbance increased. Highest species richness estimates were observed after catastrophic fire, with several species establishing exclusively under these conditions. Relative abundance of canopy tree regeneration was most similar after canopy gap and moderate‐severity disturbances. For the sub‐canopy tree and shrub community, relative species abundances were most similar after moderate‐severity disturbances and catastrophic fire. Vegetation responses to moderate‐severity disturbances thus had commonalities with both extremes of the disturbance‐caused mortality gradient, but for different regeneration layers. Conclusions: Current spatio‐temporal parameters of natural disturbances causing varying degrees of mortality promote the development of a complex, multi‐cohort forest condition throughout the landscape. The projected increase in time intervals between catastrophic fires may lead to reduced diversity within the system.     

Lack of tree layer control on herb layer characteristics in a subtropical forest,China

Question: Knowledge of the interaction between understorey herb and overstorey tree layer diversity is mostly restricted to temperate forests. How do tree layer diversity and environmental variables affect herb layer attributes in subtropical forests and do these relationships change in the course of succession? Do abundance and diversity of woody saplings within the herb layer shift during succession? Location: Subtropical broad‐leaved forests in southeast China (29°8′18″‐29°17′29″N, 118°2′14″118°11′12″E). Methods: A full inventory of the herb layer including all plants below 1‐m height was done in 27 plots (10 × 10 m) from five successional stages (<20, <40, <60, <80 and ≥80 yr). We quantified the contribution of different life forms (herbaceous, woody and climber species) to herb layer diversity and productivity and analysed effects of environmental variables and tree layer diversity on these attributes. Results: Herb layer composition followed a successional gradient, as revealed by non‐metric multidimensional scaling (NMDS), but diversity was not correlated to the successional gradient. There was no correlation of diversity across layers. Herb layer productivity was neither affected by tree layer diversity nor by herb layer diversity. Although abundance of woody species in the herb layer decreased significantly during succession, woody species contributed extraordinarily to herb layer species diversity in all successional stages. All environmental factors considered had little impact on herb layer attributes. Conclusions: The subtropical forest investigated displays an immense richness of woody species in the herb layer while herbaceous species are less prominent. Species composition of the herb layer shows a clear successional pattern, however, the presence or absence of certain species appears to be random.     

Aggregated recruitment patterns under adult crowns in Photinia glabra,a bird‐dispersed tree species

To clarify recruitment patterns of Photinia glabra, which is an evergreen, broad‐leaved, bird‐dispersed tree species, we analyzed spatial distribution in P. glabra recruits at each growth stage and demography of current‐year seedlings with respect to distributions of adults in a warm‐temperate secondary forest, western Japan. Although individuals ≥ 5 cm diameter at breast height (DBH) that had nearly produced fruits showed a random distribution, seedlings (≥ 1 year old, < 10‐cm stem length [SL]), small saplings (10 ≤ SL < 30 cm) and large saplings (≥ 30‐cm SL, < 5‐cm DBH) were clumped and associated with reproductive adults at approximately 2–3‐m scales, nearly equal to their average crown radius. Based on monitoring the demography of current‐year seedlings, emerged seedling density profoundly decreased, and no seedlings survived at longer than an adult's crown scales, with distance‐dependent mortality as a result of disease and herbivory not greatly affecting the current‐year seedling mortality. Thus, aggregated seed dispersal under the crown of adult P. glabra would directly influence the distribution of recruits for P. glabra in this forest. Of the bird‐dispersed tree species in this forest, P. glabra produced the highest amount of fruits during large crop years, and their fruits ripened during the late seasonal period (early January), suggesting that birds might be strongly attracted to these species, in turn leading to seeds being deposited mostly under the tree crowns. We propose that dispersal limitation would occur, even in a bird‐dispersed tree species such as P. glabra, owing to plant–bird interactions in the forest.     

Incorporating Restoration in Sustainable Forestry Management: Using Pine-Bark Mulch to Improve Native Species Establishment on Tephra Deposits

Native plant establishment is limited by harsh environmental conditions in areas affected by tephra deposition following volcanic eruptions. Late‐successional species might be lacking even decades after the disturbance. We assessed the effectiveness of pine‐bark mulch, a by‐product of sustainable timber production in the study area, in promoting the establishment and survival of a late‐successional species (Pinus pseudostrobus) and a nitrogen‐fixing legume (Lupinus elegans). We established a factorial experiment in areas covered with tephra during the eruption of the Paricutín volcano in the state of Michoacán, Mexico. After 1 year, P. pseudostrobus survival was significantly higher (p < 0.001) in plots with pine‐bark mulching (46.5%) than in plots without mulching (21.8%). After 2 years, surviving pines with mulching were significantly taller (p= 0.03) than pines without mulching (45.3 ± 3.8 cm and 31.2 ± 3.7 cm, respectively). Lupinus elegans plants survived longer when grown in plots with pine‐bark mulching than without mulching. Mulching reduced tephra temperatures during the dry season (when temperatures can reach up to 58°C 4 cm below the surface of bare tephra). Lupinus elegans plants were affected by herbivory by small rodents, run‐off, and frost at the end of the growing season. Our results suggest that mulching can ameliorate harsh environmental conditions on sites covered with tephra while incorporating a by‐product of sustainable forestry into restoration practice.     

Patterns of woody plant abundance,recruitment, mortality,and growth in a 65 year chronosequence of old-fields

We surveyed vegetation along forest margins in a 65-year chronosequence of old-fields at the Cedar Creek Natural History Area in east-central Minnesota, USA, to identify successional patterns of woody plants and to determine if these were correlated with soil nitrogen. We predicted that shrub and tree abundance, size, and distance of occurrence from the forest edge would be correlated with field age or soil nitrogen. Instead we did not find successional trends in the abundance or composition of woody species. Even in the oldest field the abundance of trees and shrubs was low and concentrated in areas close to the forest. Though trees were larger and present further from the forest edges in older fields, average tree height was less than 126 cm in all fields.Since we did not find successional trends we looked at various local factors (local seed sources, deer browsing, and forest edge aspect) and their relation to recruitment, mortality, or growth to explain variation among fields in abundance of trees or shrubs. The three most common tree species (Quercus rubra, Q. macrocarpa,and Populus tremuloides) all had a higher relative abundance of seedlings, and two (Q. rubra and Q. macrocarpa) had a higher relative abundance of large trees adjacent to forests with a high abundance of conspecific adults. Most trees taller than 20 cm were browsed by deer and were shorter in 1995 than they were in 1993. Mortality was higher for trees less than 30 cm indicating that mortality was size-dependent. Forest edge aspect did not significantly influence the abundance or demography of any species. Our results suggest that the patterns of seedling recruitment were largely determined by the proximity of seed sources and that these patterns may persist so that tree communities in old-fields resemble adjacent forests. Deer may be a significant factor in the suppression of tree populations in old-fields through repeated browsing which reduces tree growth and elevates tree mortality by prolonging the period of time trees remain susceptible to size-dependent mortality.     

Leaf Litter Decomposition and Substrate Chemistry of Early Successional Species on Landslides in Puerto Rico1

Decomposition is a critical process for nutrient release and accumulation of soil organic matter in disturbed soils, such as those found on landslides. I conducted a decomposition experiment on five landslides in the Luquillo Mountains of Puerto Rico as part of an investigation of the successional roles of two of the most common plant colonists to landslides, Cecropia schreberiana Miq. (Cecropiaceae) a pioneer tree species, and Cyathea arborea (L.) Sm. (Cyatheaceae) a pioneer tree fern. I compared leaf litter decomposition over one year and the initial and 1‐yr chemistry for both species. Initial litter chemistry differed between the two species, as Cecropia had slightly higher nitrogen (9.2 mg/g) than Cyathea (8.2 mg/g) and higher lignin (28.6%) than Cyathea (26.0%), but water‐soluble carbon and nonpolar extractable carbon (fats and oils, waxes, chlorophylls) were higher in Cyathea than Cecropia. Total carbon, acid‐soluble carbon, total phosphorus, and pH did not differ significantly between leaf litter species. Across all five landslides, Cyathea (k= 0.93 ± 0.06) leaves decomposed significantly faster than Cecropia (k= 0.68 ± 0.06). The differences in these species leaf litter decomposition rates and chemical composition could potentially influence organic matter dynamics and nutrient cycling rates in these early successional systems.     

Seedling assemblages and the alternative successional pathways experienced by Atlantic forest fragments

Background: Habitat loss and fragmentation have been argued to drastically alter the composition of tree assemblages inhabiting small forest fragments but the successional trajectory experienced by such edge-affected habitats remains controversial.

Aims: Here we examine whether small fragments (3.4–91.2 ha) support seedling assemblages more similar to those in 10–70-year-old secondary forests than to those in mature forests, in order to infer to what extent fragments move toward early successional systems.

Methods: Using 59 0.1-ha plots distributed in a fragmented landscape of Brazilian Atlantic forest, we evaluated species richness and functional and taxonomic composition of seedling assemblages in 20 small forest fragments, 19 stands of secondary forest and 20 stands of mature forests in the interior of an exceptionally large fragment (ca. 3500 ha).

Results: Small fragments presented the least species-rich seedling assemblages (17.2 ± 5.7 species), followed by secondary (22.5 ± 5.3), and mature forest (28.4 ± 5.3). Small fragments had seedling assemblages with functional and taxonomic composition more similar to those in secondary than in mature forest. Small fragments had a greater relative richness and abundance of pioneer trees (ca. 40% more), vertebrate-dispersed (6–25%), and those bearing medium-sized seeds (30–70%), while large-seeded species and individuals were reduced (>50% decrement) in comparison to seedling assemblages in mature forest.

Conclusions: By comparing seedlings across a wide range of successional habitats we offer evidence that small forest fragments are experiencing an alternative successional pathway towards an early-successional system with reduced plant diversity.     

Colonization dynamics and facilitative impacts of a nitrogen‐fixing shrub in primary succession

Abstract. Whether nitrogen‐fixing plants facilitate or inhibit species change in primary succession is best resolved by examining their impacts throughout the plant's entire life cycle from arrival to senescence. We experimentally examined two aspects of the successional impacts of a nitrogen‐fixing shrub, Coriaria arborea, on Mt. Tarawera, a volcano in New Zealand: factors limiting Coriaria colonization and impacts of Coriaria‐induced soil changes on a later successional tree, Griselinia littoralis. Coriaria germination was promoted by artificial wind protection and by the presence of heath shrubs. Transplanted Coriaria seedlings survived only if nodulated with Frankia, and the addition of Coriaria‐enriched soils slowed Coriaria seedling growth and did not improve seedling survival. This explained why Coriaria seedlings were found mostly in protected habitats away from adult Coriaria, and suggested that Coriaria thickets are not self‐replacing. Coriaria increased soil fertility by developing a 4 cm thick organic soil horizon that was richer in nitrogen (tenfold) and phosphorus (threefold) than pre‐Coriaria stages. These soil changes resulted in three‐ to sixfold increases in growth of Griselinia when it was grown in Coriaria‐enriched soils in a glasshouse. Coriaria's net effect on primary succession is facilitative, but the establishment of Coriaria is itself facilitated by the amelioration of the physical habitat by earlier colonists, suggesting facilitation is important throughout the life cycle of Coriaria. Sequential facilitative events determine the order of species replacements in this study but inhibition, linked to the developmental stages of Coriaria, may determine the rate of species change.