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.     

Recruitment limitation of forest communities in a degraded Mediterranean landscape

Question: How does habitat degradation affect recruitment limitation and its components (seed limitation versus establishment limitation) of woody plant communities in a Mediterranean landscape? Location: 1600‐1900 m a.s.l. in the Sierra Nevada National Park, southern Spain. The landscape is a mosaic composed of native forest and two degraded landscape units: reforestation stands and shrubland. Methods: We evaluated fruit production, seed rain, seedling emergence and seedling survival in two consecutive years with contrasting rainfall patterns. Seed and seedling data were used to calculate values of seed and establishment limitation. Results: In general, the woody community was both severely seed‐ and establishment‐limited. Species were less seed‐limited in the landscape units with higher adult density (i.e. shrub species in shrubland, Pinus spp. in reforestation stands). In contrast, degradation did not exacerbate establishment limitation, which was severe in all landscape units. This general pattern was modulated by the biogeographical distribution, dispersal type, and life form of the species. Boreo‐alpine species were more limited in establishment than species with a typical Mediterranean distribution. Zoochorous species were less seed‐limited in the landscape units preferred by dispersers (i.e. native forest). Tree species were more establishment‐limited than shrub species, irrespective of the landscape unit. Seed limitation, and especially establishment limitation, varied among years, with establishment being almost nil in the very dry year. Conclusion: In the case of Mediterranean landscapes, when degradation from human impact involves a reduction in the adult abundance of the woody plant community (trees and shrubs), seed limitation increases, although establishment limitation is generally high in all landscape units, especially for boreo‐alpine species. Conservation and restoration strategies should take into account our results showing that tree species were unable to recruit in an extremely dry year, because more aridity is expected under a climatic change scenario in Mediterranean ecosystems.     

Seed dispersal and seedling recruitment of trees at different successional stages in a temperate forest in northeastern China

Aims Spatial distribution of adult trees in a forest community is determined by patterns of both seed dispersal and seedling recruitment. The objectives of our study were to understand the processes of seed dispersal and seedling recruitment of dominant tree species in a temperate forest of northeastern China and to identify the factors constraining seed dispersal and seedling establishment at different stages of forest succession.Methods During three summer and autumn sessions between 2006 and 2008, altogether 113080 seeds from 22 different tree species were collected in three large field plots representing different forest types in the Changbai Mountain region of northeastern China. The spatial distribution of seed abundance was analyzed using a Syrjala test. Regeneration success of nine major tree species was assessed using variables defining 'limitations' in 'seeds' and 'seedling establishment'.Important findings We found that seed production fluctuated between years and varied greatly with forest types. Four tree species, Acer spp., Fraxinus mandshurica, Tilia amurensis and Betula spp., had the greatest seed production and the widest range of seed dispersal, whereas Quercus mongolica showed the most sustained seed production pattern. The spatial patterns of seed abundance differed significantly among forest types and years. The tree species investigated in this study differed in the degree of seed limitation, as well as in limitation of seedling establishment. There were both negative and positive correlations between seed density and seedling density, depending on site and parental tree density. Seeds of 16 tree species were found in the Populus davidiana–Betula platyphylla forest (PBF) plot, 11 in the conifer and broad-leaved mixed forest (CBF) plot but only 8 in the broad-leaved-Korean pine mixed forest (BKF) plot. The number of seed-contributing species was not only greater in the secondary forests (CBF and PBF plots) than in the primary forest (BKF plot) but was also more variable during the 3 years of assessment. Results from the correlations between seed density and seedling occurrence and that between parental tree density or seed weight and dispersal limitation confirm our intuitive expectations, i.e. heavy seeds had greater dispersal limitation but higher establishment success than light seeds.     

子午岭三种生境下辽东栎幼苗定居限制

辽东栎(Quercus wutaishanica)是子午岭地区的乡土乔木树种,也是该地区气候顶极群落的建群种,其幼苗的补充更新影响着森林群落的结构及物种组成。在3种典型生境(辽东栎林、人工油松林、灌草丛)中,设置3因素(种子、干扰、遮荫)两水平的野外播种实验(随机区组设计),记录辽东栎幼苗出苗量,并监测幼苗同生群在3种生境中3a间的生长状况,以确定种子及微生境在辽东栎幼苗补充过程中的限制作用。播种实验样方大小30cm×30cm,共计216个样方。结果显示,在辽东栎林及油松林内,增加种子和干扰强度(去除枯落物),能引起出苗量和幼苗补充量的显著增加,且2种处理方式间存在交互作用,表明在郁闭林冠下,辽东栎幼苗的补充受到了种子和微生境的双重限制,枯落物是导致微生境限制的主要因素之一。在灌草丛生境,各种处理方式均不能增加幼苗的补充量,表明辽东栎无法在开阔生境(强光照、干旱)中完成实生幼苗的补充更新。3种生境中的幼苗同生群生存分析表明,辽东栎幼苗在森林群落中存活率显著高于灌草丛群落。根据幼苗生长指标判断,在3种生境中,人工油松林是辽东栎幼苗定居的最佳场所。与实验预期相反,灌木对辽东栎幼苗的补充无显著影响。     

Dispersal limits natural recruitment of African mahoganies

The abundance of forest tree species may be locally limited by the inability of species to disperse to all sites suitable for germination and establishment. This phenomenon of "dispersal limitation" has been suggested to promote the maintenance of high species diversity in tropical forests by slowing down competitive exclusion. We present the first direct experimental evaluation of dispersal limitation in tropical forests, and of its importance relative to other factors affecting recruitment. Seed addition, litter removal and light availability effects on seedling establishment were evaluated in a full factorial experiment on two African mahogany species, Entandrophragma utile and Khaya anthotheca . Seedling recruitment after 18 months, was ∼22 times greater in seed addition treatments (with addition rates corresponding to the tail of seed shadows of large reproductive trees) compared to controls. This seed addition effect was an order of magnitude greater than either gap or litter removal effects, with similar results observed in both logged and primary forest locations. We conclude that dispersal limitation strongly limits seedling establishment for the two species studied. An important "applied corollary" of this result is that seed supplementation may substantially increase local abundance of valuable or rare species in disturbed or managed tropical forests.     

Consequences of habitat disturbance on seed fate of a Brazilian tropical dry forest tree Cavanillesia arborea (Malvaceae)

We compared seed fate (survival, mortality by rodent predators, desiccation and other causes) of the tree Cavanillesia arborea (Malvaceae) in preserved old‐growth tropical dry forests and in nearby abandoned pasturelands in Brazil. For this purpose, we performed an experiment where 15 seeds were placed in the surroundings of 15 parental individuals in each habitat. These 450 seeds were monitored over four months. At the end of the experiment, seed predation by rodents was higher in forest (56.5%) than in pasture (8.9%) areas, but seed desiccation showed the opposite pattern (8.9% vs. 80.4%). Mortality by desiccation was also faster in pasture than in forest areas, probably reducing their attractiveness to predators in these areas. None of the seeds placed in the pasture survived, whereas 26.2% of seeds became seedlings in forest areas. The absence of seedling recruitment of Cavanillesia arborea in pastures is likely a consequence of their incapacity to tolerate the harsh abiotic conditions in this habitat. Thus, forest conversion imposes a strong limitation to the long‐term population viability of this species. However, restoration and natural regeneration of abandoned pastures can recreate the forest structure and microclimatic conditions favourable to seed germination and seedling establishment. Remnant individuals of Cavanillesia arborea in agricultural landscapes may have a negligible contribution to current recruitment, but they can attract potential dispersers of pioneer species, with positive feedbacks to future recruitment during secondary succession.     

High Acorn Predation Prevents the Regeneration of Quercus liaotungensis in the Dongling Mountain Region of North China

The mature oak (Quercus liaotungensis Koidz) forests in the Dongling mountains of northern China have become degraded in recent years because regeneration has been limited. To determine whether or not seedling establishment of the oak is seed limited, microsite limited, or predator limited and to determine whether seedling establishment is affected by ground cover, we conducted field experiments during a mast year and investigated the fate of seeds and the soil seed bank dynamics of the oak. A large acorn crop (128.8 acorns/m²) was observed in the study period, and the peak density of acorns on the forest floor reached 46.5 acorns/m², suggesting that tree recruitment was not seed limited. Acorns in the soil seed bank were mainly lost through decay (principally after fungal attack), consumption in situ, and removal by animals. Predation (including consumption in situ and removal) accounted for 86.4% of acorn loss and was therefore likely to have been the most important factor influencing seed dynamics. More than 70% of acorns were found to have germinated, but no established seedling was observed on the forest floor. Using cages to exclude predators, it was estimated that 87% of acorns germinated and 49% became established as seedlings, indicating that the acorns on the forest floor could emerge and grow in the absence of predators. We conclude that the regeneration of the tree population is limited by predators rather than by the availability of microsites. The presence of ground cover increased the germination rate and increased the chance of seed survival in the early stage of the experiment, but at the end of the investigation, no established seedling was found in the quadrats both with and without ground cover, possibly because of high density of animal predators. On the basis of these results, we suggest that selective tree felling will increase the coverage of the herbaceous layer, which can further decrease the population density of the rodents, and thereby improve the regeneration of oak trees.     

Patch to landscape patterns in post fire recruitment of a serotinous conifer

Obligate seeding species are highly specialized to fire disturbance and many conifers such as cypress, which are adapted to high intensity stand-replacing fires, have canopy seed banks stored in serotinous cones. Resilience of these trees to fire disturbance is a function of disturbance frequency and one focus of this study was to determine the effect of patch age on postfire recruitment. A second focus was to determine the extent to which fire induced a landscape level change in the location of the forest boundary. Prior to a fire in 1994, a large Cupressus sargentii forest was a mosaic landscape of different aged patches of nearly pure cypress bordered by chaparral. Patches less than 60 years of age were relatively dense with roughly one tree every 1–2 m² but older patches had thinned to one tree every 3–15 m². Older trees had substantially greater canopy cone crops but the stand level seed bank size was not significantly correlated with stand age. Fire-dependent obligate seeding species are sensitive to fire return interval because of potential changes in the size of seed banks – facing both a potential `immaturity risk' and a `senescence risk'. At our site, C. sargentii regeneration was substantial in stands as young as 20 years, suggesting that fire return interval would need to be shorter than this to pose any significant risk. Reduced seedling recruitment in stands nearly 100 years of age may indicate risk from senescence is greater, however, even the lowest density seedling recruitment was many times greater than the density of mature forests – thus this cypress would appear to be resilient to a wide range of fire return intervals. Changes in landscape patterning of forest and chaparral are unlikely except after fire. Factors that inhibit tree establishment within the shrubland, as well as factors that affect shrub establishment within the forest border likely affect the `permeability' of this ecotone. After the 1994 fire this boundary appeared to be stable in that cypress recruited best within the shadow of burned canopies and cypress were weak invaders of adjacent shrublands.     

Contrasting demographics of tropical savanna and temperate forest eucalypts provide insight into how savannas and forests function. A case study using Corymbia clarksoniana from north‐eastern Australia

Eucalypts (Eucalyptus spp. and Corymbia spp.) dominate many communities across Australia, including frequently burnt tropical savannas and temperate forests, which receive less frequent but more intense fires. Understanding the demographic characteristics that allow related trees to persist in tropical savannas and temperate forest ecosystems can provide insight into how savannas and forests function, including grass–tree coexistence. This study reviews differences in critical stages in the life cycle of savanna and temperate forest eucalypts, especially in relation to fire. It adds to the limited data on tropical eucalypts, by evaluating the effect of fire regimes on the population biology of Corymbia clarksoniana, a tree that dominates some tropical savannas of north‐eastern Australia. Corymbia clarksoniana displays similar demographic characteristics to other tropical savanna species, except that seedling emergence is enhanced when seed falls onto recently burnt ground during a high rainfall period. In contrast to many temperate forest eucalypts, tropical savanna eucalypts lack canopy‐stored seed banks; time annual seed fall to coincide with the onset of predictable wet season rain; have very rare seedling emergence events, including a lack of mass germination after each fire; possess an abundant sapling bank; and every tropical eucalypt species has the ability to maintain canopy structure by epicormically resprouting after all but the most intense fires. The combination of poor seedling recruitment strategies, coupled with characteristics allowing long‐term persistence of established plants, indicate tropical savanna eucalypts function through the persistence niche rather than the regeneration niche. The high rainfall‐promoted seedling emergence of C. clarksoniana and the reduction of seedling survival and sapling growth by fire, support the predictions that grass–tree coexistence in savannas is governed by rainfall limiting tree seedling recruitment and regular fires limiting the growth of juvenile trees to the canopy.     

Multiple stages of tree seedling recruitment are altered in tropical forests degraded by selective logging

Tropical forest degradation is a global environmental issue. In degraded forests, seedling recruitment of canopy trees is vital for forest regeneration and recovery. We investigated how selective logging, a pervasive driver of tropical forest degradation, impacts canopy tree seedling recruitment, focusing on an endemic dipterocarp Dryobalanops lanceolata in Sabah, Borneo. During a mast‐fruiting event in intensively logged and nearby unlogged forest, we examined four stages of the seedling recruitment process: seed production, seed predation, and negative density‐dependent germination and seedling survival. Our results suggest that each stage of the seedling recruitment process is altered in logged forest. The seed crop of D. lanceolata trees in logged forest was one‐third smaller than that produced by trees in unlogged forest. The functional role of vertebrates in seed predation increased in logged forest while that of non‐vertebrates declined. Seeds in logged forest were less likely to germinate than those in unlogged forest. Germination increased with local‐scale conspecific seed density in unlogged forest, but seedling survival tended to decline. However, both germination and seedling survival increased with local‐scale conspecific seed density in logged forest. Notably, seed crop size, germination, and seedling survival tended to increase for larger trees in both unlogged and logged forests, suggesting that sustainable timber extraction and silvicultural practices designed to minimize damage to the residual stand are important to prevent seedling recruitment failure. Overall, these impacts sustained by several aspects of seedling recruitment in a mast‐fruiting year suggest that intensive selective logging may affect long‐term population dynamics of D. lanceolata. It is necessary to establish if other dipterocarp species, many of which are threatened by the timber trade, are similarly affected in tropical forests degraded by intensive selective logging.     

Gnawing damage by rodents to the seedlings ofFagus crenata andQuercus mongolica var.grosseserrata in a temperateSasa grassland-deciduous forest series in southwestern Japan

The effects of dwarf bamboo,Sasa, cover on the initial morrality of hardwood seedlings were investigated by transplanting 1-year-old beech (Fagus crenata) and current-year oak (Quercus mongolica var.grosseserrata) seedling to three different stands; old-growth beech and secondary oak forests withSasa undergrowth, and aSasa grassland in a grassland-forest series near the top of Mt Jippo, southwestern Japan. The most frequent cause of seedling morrality was gnawing of the stems by rodents. In the beech forest, the gnawing was more likely to occur underSasa cover, suggesting that it provides a good habitat for rodents on the beech forest floor. TheSasa under growth may thus play an imporrant role in regeneration of beech forest. In the oak floor, mortality of both species was low and only a little gnawing occurred during a year. However, no natural oak seedling were found in the forest even after a mast year. This may be because most of the acorns disappeated before establishment. The early-stage demography of hardwood seedling as oak may thus play an imporrant role in regeneration of oak forest. In theSasa grassland where the seed supply is small, almost all of the seedlings died fromo gnawing regardless of the presence ofSasa cover. These factors prevent the recruitment of a sizable seedling bank. Rodents may thus play an imporrant role in maintenance of theSasa grassland.     

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.     

Effects of altitude and livestock on the regeneration of two tree line forming <Emphasis Type="Italic">Polylepis</Emphasis> species in Ecuador

Regeneration is known to be limited at many temperate tree lines, but very little data is available on the impacts of altitude and anthropogenic disturbance on regeneration patterns along tropical tree lines. The study focused on the reproductive traits of two Polylepis species in the Páramo de Papallacta in Ecuador along an altitudinal gradient, and involved different intensities of cattle trampling within subsequent altitudinal ranges. We analyzed flowering, fruit set, seed viability, germination, and seedling establishment as well as stand structure of Polylepis incana and P. pauta. The numbers of P. incana inflorescences and seedlings per m2 showed a marginally significant decrease with increasing altitude. Mean tree height was significantly lower at higher altitudes, while stem number increased. The number of P. pauta inflorescences also decreased significantly upslope. In both forest types, trampling was found to have a positive impact on seedling abundance, presumably due to the removal of the litter layer. Thus, there was no evidence of negative effects of moderate cattle grazing on both tree line species. However, sapling establishment was minimal inside the forest stands at all altitudes and grazing levels, and we consequently observed a low proportion of narrow stems within all investigated forests. Our results show that, along with vegetative growth limitations of adult trees, important regeneration traits such as seedling and inflorescence numbers are also influenced by altitude, which might contribute to the formation of the upper tree line. Nevertheless, recruitment in the forest interior was low overall indicating that further factors, such as light conditions, affect regeneration of the studied species.     

Causes of arrested succession in coastal dune forest

Dispersal or recruitment limitation may arrest succession after disturbance. In north-eastern South Africa the Acacia karroo successional pathway is used to facilitate coastal forest recovery after strip-mining. However, although A. karroo establishes naturally, it forms monospecific stands, arresting forest succession for decades. This casts doubt on the efficacy of this restoration pathway. We investigated the causes of arrested succession. The seed and seedling banks of A. karroo stands and of forest at Cape Vidal, and three A. karroo stands (7–27 years old) on rehabilitated strip-mined dunes at nearby Richards Bay were examined. The establishment and growth of seedlings at Cape Vidal were also considered. The seed bank was larger and more diverse in forest, but the seedling bank was larger in Acacia stands. At Richards Bay, the size of the seed bank increased and the seedling bank decreased with Acacia stand age. Excluding mammalian herbivores in Acacia stands at Cape Vidal resulted in greater species richness and survival of naturally established seedlings, as well as two experimentally planted species. Neither seed dispersal nor seedling establishment limited recruitment of tree species in Acacia stands. Herbivory arrested forest succession by causing the differential mortality of seedlings. In contrast, at Richards Bay where there were few mammalian herbivores, the advanced regeneration in A. karroo stands converged on the diversity of nearby forests 29 years after restoration. Controlling herbivore access and seeding Acacia stands with forest species are site-specific options for preventing arrested succession when using the A. karroo successional pathway.     

Contrasting patterns of seed dispersal between alien mammals and native lizards in a declining plant species

The introduction of carnivorous mammals has led many native island species to extinction. In the Balearic Islands, the introduction of carnivorous mammals in the main islands has contributed to the extinction of the endemic lizard Podarcis lilfordi, which is an important seed disperser of many plant species. One of the introduced mammals, Martes martes (pine marten), is also known to disperse seeds and may replace the native lizard in this role in islands. However, little is known on the patterns of seed dispersal by these two different species and their possible implications for plant regeneration ability and population structure. We have compared the quality of seed deposition provided by both to the vulnerable Mediterranean shrub Cneorum tricoccon, by studying the pattern of seed distribution among sites generated by both dispersers as well as the suitability of those sites for seedling emergence and establishment. The study was carried out in two types of habitats (coastal shrublands and pine forests) and two islands (Mallorca and Dragonera). Lizards and mammals showed contrasting patterns of seed deposition and, where lizards were absent, mammals played their role as seed dispersers in pine forests but not in coastal shrublands. The lack of seed dispersal in the coastal shrubland seriously limits recruitment, by concentrating seeds under conspecifics and hindering colonisation opportunities, marking a long-term trend towards decline. The introduced predator replaces the native seed disperser in its service where it has gone extinct, although with important differences in the dispersal service provided.     

兴安落叶松(Larix gmelinii)火后种子萌发影响因子

林火干扰是影响兴安落叶松林结构和功能的主要因子之一。兴安落叶松种群火后更新受多种因子的影响,并决定着该群落演替轨迹。通过在大兴安岭呼中自然保护区的火烧迹地内设置原位控制实验,利用增强回归树分析方法,量化研究了落叶松火后恢复初期不同影响因子（温度、有机质层厚度、覆盖度等）对种子萌发的相对重要性。研究结果表明：落叶松火后种子萌发的最主要影响因子为温度,第二影响因子为种源,第三影响因子为草本覆盖度,分别解释了幼苗数量变异的28.51%、22.40%、20.66%;各影响因子的相对重要性在不同地形条件下有明显差异：温度在山坡顶部和阳坡底部占有重要影响,种源在阳坡中部和阴坡中部最为重要,土壤含水量在阴坡底部占有重要地位。同时,去除土壤表面有机质可以显著提高种子萌发数量,去除地面杂草则会使种子萌发数量降低。从研究结果可知,落叶松火后种子萌发在不同地形（环境条件）下的限制因子不同,人工辅助需因地制宜采取相应措施,才能更有效地促进种子萌发与森林恢复。     

Effects of Soil Burn Severity on Post-Fire Tree Recruitment in Boreal Forest

Fire, which is the dominant disturbance in the boreal forest, creates substantial heterogeneity in soil burn severity at patch and landscape scales. We present results from five field experiments in Yukon Territory, Canada, and Alaska, USA that document the effects of soil burn severity on the germination and establishment of four common boreal trees: Picea glauca, Picea mariana, Pinus contorta subsp. latifolia, and Populus tremuloides. Burn severity had strong positive effects on seed germination and net seedling establishment after 3 years. Growth of transplanted seedlings was also significantly higher on severely burned soils. Our data and a synthesis of the literature indicated a consistent, steep decline in conifer establishment on organic soils at depths greater than 2.5 cm. A meta-analysis of seedling responses found no difference in the magnitude of severity effects on germination versus net establishment. There were, however, significant differences in establishment but not germination responses among deciduous trees, spruce, and pine, suggesting that small-seeded species experience greater mortality on lightly burned, organic soils than large-seeded species. Together, our analyses indicate that variations in burn severity can influence multiple aspects of forest stand structure, by affecting the density and composition of tree seedlings that establish after fire. These effects are predicted to be most important in moderately-drained forest stands, where a high potential variability in soil burn severity is coupled with strong severity effects on tree recruitment.     

Tree Regeneration in Church Forests of Ethiopia: Effects of Microsites and Management

Tree regeneration is severely hampered in the fragmented afromontane forests of northern Ethiopia. We explored how trees regenerate in remnant forests along the gradient from open field, forest edge to closed sites and canopy gaps inside the forest. We investigated the effects of seed sowing, litter removal, and weeding on the regeneration success along this gradient. Regeneration success was investigated for four indigenous tree species, and measured in terms of seedling establishment, growth, and survival. Species performed differently according to site conditions. Within the forest, local canopy openings facilitated seed germination ( Ekebergia ), seedling growth (all species except Olea ), or survival ( Ekebergia and Olea ), suggesting that all species benefited from local high light conditions in the forest. Outside the forest, germination (all species) and growth rates ( Juniperus and Olea ) were lower in the open field, most probably due to water stress in the dry season. Outer edge conditions favored growth for three of the four species. Natural seed germination was, however, zero at any site for Juniperus and Olea and low for Ekebergia and Prunus in the open field. Soil scarification influenced germination positively, while weeding did not have a positive effect. These results suggest that simple measures may improve seedling establishment, and that, for some species, forest edges are particularly useful for growth and survival after succesful establishment. Together with erecting fences, needed to protect seedlings against grazing, seed sowing, planting seedling, and soil scarification may contribute to maintain and restore church forests in the fragmented landscapes of northern Ethiopia.     

PEG模拟干旱胁迫对栓皮栎种子萌发及生长生理的影响

为了阐明栓皮栎种子萌发期对水分胁迫的耐受性,以当年采集的栓皮栎种子为材料,采用培养皿滤纸萌发的方法研究了不同浓度（0%、5%、10%、20%、30%）PEG-6000溶液对其种子萌发、生长、保护酶活性和有机渗透调节物质的影响,并对种子萌发率、胚根生长速率等与PEG浓度进行了回归分析。结果表明:（1）随着干旱胁迫程度的加剧,栓皮栎种子的发芽率、发芽指数、胚根长度、胚根生长速率和活力指数均呈先升高后下降趋势,说明低浓度（5%、10%）PEG胁迫可以促进栓皮栎种子萌发及生长,而高浓度（20%、30%）则具有抑制作用。（2）建立水分胁迫条件下种子萌发率、胚根生长速率等与PEG浓度的回归方程发现,栓皮栎种子在PEG胁迫下的萌发临界值和极限值分别为32.01%和36.99%,胚根生长速率的临界值和极限值分别为30.1%和33.35%。（3）随着PEG浓度的升高,栓皮栎种子中3种保护酶（SOD、POD、CAT）活性和3种有机渗透调节物质（可溶性蛋白、脯氨酸、可溶性糖）含量呈现不同的变化,但均有不同程度升高趋势。由此可见,栓皮栎种子可通过增加保护酶活性和体内渗透调节物质来适应水分胁迫环境,使自身萌发和生长受抑制程度降到最低。     

Recruitment limitation in dry sclerophyll forests: Regeneration requirements and potential density‐dependent effects in Eucalyptus tricarpa (L.A.S. Johnson) L.A.S. Johnson & K.D. Hill (Myrtaceae)

Recruitment limitation in canopy trees is receiving increasing attention as restoration of tree cover for connectivity, biodiversity offsets, carbon‐trading and improved catchment health becomes more prominent. Recruitment limitation is often addressed by examining seed traits and germination requirements. Distance between trees is also often explored as a factor, particularly in agricultural landscapes where forest structure has been altered, with large distances between fragments generating Allee effects linked to pollen limitation. Fewer studies have examined how short distances between trees (high stem densities), which can characterize regenerated forests, might affect recruitment. This study examined recruitment limitation by exploring germination requirements of Eucalyptus tricarpa (red ironbark) and the effects of stem density on seed characters and germination. Eucalyptus tricarpa is a canopy species in the box‐ironbark forests of central Victoria, Australia. During the early European settlement phase in the 1800s these forests were heavily impacted by human disturbances including mining, logging and livestock grazing. In many regenerated stands, now more than 100 years old, current stem density exceeds those documented in 19th century reports and there is little regeneration of key canopy species such as E. tricarpa. In laboratory trials seed germinated at 18°C in darkness, and while viability varied among populations, overall it remained high (91–96%) and did not differ among populations with different stem densities. An in situ germination trial addressing sowing season, ground‐cover, soil‐ripping, canopy‐cover and stem density recorded no germination. High stem density in coppicing box‐ironbark forest was hypothesized to limit resources for reproduction; however, no relationship was found between nearest neighbour distance, allocation to selected reproductive structures or seed viability. The relative insensitivity of reproductive allocation to intraspecific competition (high stem density) may be interpreted as a trait linked to unpredictable environments and consistent with long‐lived species. In terms of management, recruitment limitation in E. tricarpa appears not primarily attributable to seed viability, tested seedbed treatments or stem density so further factors, including fire, now require exploration.