Testing for Janzen–Connell Effects in a West African Montane Forest

The Janzen–Connell hypothesis proposes that density dependent seed and seedling mortality, combined with increasing seed and seedling survival away from the parent tree, together promote regular spacing of species and thus α diversity. This hypothesis has rarely been tested in tropical Africa, and rarely in montane forests anywhere. We tested this hypothesis using a combination of field experiments and observations in the most floristically diverse dry submontane forest in Nigeria. We investigated distance effects on seedling herbivory, seedling survival and seedling height growth. We found a significant decrease in herbivory with distance from conspecific adult trees for all three species of experimentally planted seedlings (Entandrophragma angolense, Deinbollia pinnata and Sterculia setigera), and also for naturally occurring seedlings of Pouteria altissima but not of Newtonia buchananii or Isolona pleurocarpa. The relative density of large seedlings/saplings of P. altissima, N. buchananii and I. pleurocarpa increased significantly at greater distance from conspecific adult trees; however, we found no significant distance effect on survival or height growth over 3 mo for all three experimentally planted species. Taken together, our results are some of the first to show that Janzen–Connell effects occur on the African continent and in particular montane tropical forest and suggest that such effects may be pantropical.     

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.     

Effect of the grass Leersia hexandra on the dispersal,seed germination,and establishment of Pachira aquatica seedlings

1. In wetlands, hydrochory is one of the main mechanisms of seed dispersal and there is often synchrony between propagule production and the flood season. Different sources of disturbance can prevent seed dispersal to suitable sites, and if environmental conditions are not adequate for germination and seedling establishment, recruitment will be limited, affecting succession.
2. We worked in a disturbed tropical freshwater swamp where the native grass Leersia hexandra has dominated open areas, creating a grass matrix that surrounds patches of swamp forest. Leersia grows vigorously, forming cushions of dry matter that cover the soil, forming a potential obstacle to seed dispersal. We asked whether the vegetative growth of this grass prevents the entry and dispersal of seeds of the tropical swamp tree Pachira aquatica, thwarting seed germination and seedling establishment, and arresting succession. We set up transects in the grass matrix in two zones: close to the river (R) and bordering the tree patches or fragments (F). We quantified tree seed and seedling presence, survival and growth in situ and experimentally introduced seeds and seedlings in the field and monitored seed germination and the survival and growth of their seedlings, as well as that of transplanted seedlings.
3. There was a negative relationship between the number of seeds and established seedlings, and the distance to river or fragment (r = −0.86, p < 0.001 for zone R; and r = −0.77, p < 0.001 for zone F) and with the grass cushion (r = −0.68, p = 0.005 for zone R; and r = −0.66, p = 0.007 for zone F); the grass creating a barrier to dispersal. When seeds were sown after clearing the grass cushion, germination success was high, so this stage is not limited. The transplanted seedlings had better survival and a greater final height than the seedlings of the sown seeds. Grass cover had a negative effect on both types of seedlings. Seedling survival rates were inversely related to grass cover, showing that seedlings overgrown by grass had low survival rates. Flooding is a stress factor for seedlings and produced mortality, in addition to the effects of the grass.
4. Together, the field survey and the experiment show that succession is being arrested in two ways: (1) by limiting seed dispersal because the grass cushion slows the dispersal and penetration of seeds into the vegetation; and (2) by limiting seedling establishment because the grass competes for space and light. Our results show that even where the grass is native, slower growing, seed-dependent species may struggle to compete and establish. If grass cover is increasing, these swamps are very vulnerable to a decrease in area because it is very difficult for them to regenerate naturally.

     

Variability in survival and mortality of Acacia drepanolobium Sjøstedt following prescribed burning at Olpejeta Conservancy,Kenya

Mortalities to Acacia drepanolobium, a main item in the diet of the eastern black rhino (Diceros bicornis Michaeli) in Olpejeta conservancy, Kenya, are caused by three main factors: drought, browsers and fire. The effect of controlled fires on A. drepanolobium was examined by monitoring survival and growth in experimental plots before and after fire episodes between 2003 and 2007. Density, survival and growth in A. drepanolobium trees were compared eighteen months before and after burning. Tall trees were more likely to die from fire compared to short trees (R² = 0.575; df = 6; P = 0.048), while seedling densities increased after fire (χ² = 36.57; df = 1; P = 0.001). Even with increased seedling densities, burned areas attracted large numbers of seedling predators, lowering the possibility of seedling recruitment into adult, as mean seedling heights reduced significantly (ANOVA, F = 204.42; df = 1; P = 0.036). Fires also significantly lowered flowering (F = 346; df = 1; P < 0.05) in A. drepanolobium, thereby affecting fruit production. Although fires caused mortalities to adult A. drepanolobium, the most significant effect was tree reversals into seedling height class as trees resprouted. Although fire may increase browse biomass of A. drepanolobium available for black rhino, it is not an appropriate black rhino habitat management tool because burnt areas attract many seedling predators that lower seedling recruitment into adult trees.     

Offspring spatial patterns in Picconia excelsa (Oleaceae) in the Canarian laurel forest

We studied the spatial patterns of seedlings and seeds in isolated Picconia excelsa (Oleaceae) trees in the laurel forest of Anaga, Tenerife (Canary Islands). By finding isolated trees we assessed the correlation of seed and seedling bank traits and parent trees by removing the confounding effects of proximity (<100 m radius) of conspecific fruiting trees. We counted all the seedlings per age (height) class within its parental range, and sampled the seed number along transects departing from beneath the parent canopy at regular intervals. We mapped all seedlings per age class and plotted seed and seedling profiles in relation to distance to parent trees. Older Picconia seedlings tended to clump significantly further from parent trees than younger seedlings, which clumped just beneath the parents. We found significant differences among distances to parent tree in numbers of seedlings per age class. The seedling bank area was significantly correlated with maximum distance of seedlings to parent trees. The majority of seeds were deposited within the first 4 m below the parent crown. Seedlings amount at further distances from the trees is larger than seeds/fruits as counted on the ground. Our results suggest that disseminated, older seedlings have occupied germination sites far from the parent tree because there is probably lower seedling–seedling and parent–seedling competition for resources, and perhaps no intraspecific allelopathy and predation/disease.     

A comparison of sites suitable for the seedling establishment of two co-occurring species, Swintonia glauca and Stemonurus scorpioides, in a tropical peat swamp forest

Spatial and temporal ground-surface dynamics are major factors that affect regeneration and species coexistence in tropical peat swamp forests. We studied the seedling survivorship and morphological features of two tree species that play important roles in maintaining the ground-surface dynamics of a peat swamp forest in Sumatra. Large Swintonia glauca trees form mounds, whereas large Stemonurus scorpioides trees occupy non-mounds. We monitored the demography of naturally dispersed Swintonia and Stemonurus seedlings that germinated in 2000. Survivorship of Swintonia seedlings was high under conditions of late germination, high-light environment, and elevated ground surface, and was negatively affected by distance to the nearest conspecific adult. Survivorship of Stemonurus was high under conditions of early germination and high conspecific seedling density, and was also negatively affected by distance to the nearest conspecific adult. The allometric features of Stemonurus seedlings indicated characteristics of stress tolerance, that is, low growth rate and thick, porous roots. Stemonurus, which has large wingless seeds, regenerated in non-mounds around the parental trees, while winged Swintonia seeds dispersed farther from the parent and established in patchily distributed gaps and mounds. Thus, Swintonia seedlings can survive on non-mound sites within gaps and possibly create mounds, while Stemonurus seedlings tend to maintain non-mounds around the parental trees.     

Distance- and density-dependent leaf dynamics of seedlings of a tropical rainforest tree

Parental distance and plant density dependence of seedling leaf turnover and survival was examined to investigate predictions of the Janzen–Connell hypothesis. The focal study species, Shorea macroptera is a canopy tree species in a lowland rain forest in peninsular Malaysia. We found that the peak of the distribution of plants shifted from 3–6 m to 6–9 m during the course of the change from seedling to sapling stage. The leaf demography of the seedlings was influenced by their distance from the adult tree and also by the seedling density. Although significant density- and distance dependence in leaf production was not detected, seedling leaf loss decreased with distance from the parent tree and with seedling density. Similarly, leaf damage was not found to be distance- or density-dependent, but net leaf gain of seedlings increased with distance from the parent tree. Although no significant distance- or density-dependence was evident in terms of leaf damage, significant distance dependence of the net leaf gain was found. Thus, we concluded that positive distance dependence in the leaf turnover of seedlings may gradually contribute to a shift in the distribution pattern of the progeny through reductions in growth and survivorship.     

Distribution ofAcer palmatum seedlings under the crown of the maternal tree

To investigate the relationship between crown width of a parent plant and seedling distribution, seedling ages and sizes, and their distances from the maternal tree were surveyed under a maple (Acer palmatum) crown at Chungdam Park, Chungdam-dong, Kangnam-gu, Seoul, in 1998. Maple crown width increased 33.0 cm per year during the eight-year study. Seedlings ranged from 1 to 10 years old; their density was highest the third year, and decreased with age. Seedling-distribution ranges from the parent plant were 43 cm to 487 cm, and seedling density was highest at the border of the parent plant canopy. One-year-old seedlings were distributed broadly, but the main distribution areas became more distant from the maternal plant with seedling age. Seedling heights increased with age; differences between two consecutive ages were significant at the 0.1% level for ages less than 5 years, but not for over 6 years. For seedlings of the same age, heights were similar, but short under the tree crown, but were varied and taller outside the crown. The variation in seedling height increased with age and distance from the parent plant In particular, seedlings older than 5 years that were distributed outside the crown showed conspicuous variations among distance classes. Therefore, maple seedlings were distributed to an appropriate distance at which their growth could be supported by the understory-light environment. This study demonstrated that newA. palmatum trees are not recruited from the seedlings growing beneath the crown but from those that grow vigorously outside the crown.     

Ecology of Aglaia mackiana (Meliaceae) Seedlings in a New Guinea Rain Forest1

The large seeds of Aglaia mackiana (Meliaceae) germinate and produce vigorous seedlings under closed canopies or in large gaps. To assess seedling ecology after germination, we measured growth, herbivore damage, and survivorship of seedlings over one year. The sample included shaded seedlings from dispersed seeds, undispersed seeds under parent trees, and seedlings transplanted to gaps. We quantified the light environment using hemispherical canopy photographs taken above seedlings at the beginning and end of the one–year study. Seedlings transplanted to gaps grew faster and had more leaves, larger total leaf surface area, longer secondary roots, and greater root mass than shaded seedlings. Seedlings in gaps did not differ from shaded seedlings in survivorship or amount of herbivore– and pathogen–caused leaf damage. The canopy photographs taken one year apart suggest there is a rough equilibrium in closed canopies with slight changes occurring around an average light level. Sites with < 0.06 ISF (a unitless, relative measure of canopy openness or reflected sunlight) tended to remain the same with minor fluctuations toward brighter or darker. Sites with canopy openness > 0.06 ISF tended to close; few gaps grew larger. Seedlings under parenr trees and seedlings away from parent trees had similar amounts of leaf damage and virtually identical survivorship after 18 months, but seedlings under parent trees had slower growth rates and smaller total leaf surface areas. Dispersal did not strongly benefit seeds via escaping high levels of mortality or comperition around the parent.     

Effects of Dispersal Pattern and Mammalian Herbivores on Seedling Recruitment for Virola michelii (Myristicaceae) in French Guiana1

The effects of dispersal pattern (seeds in small clumps vs. seeds scattered in pairs) and distance to the nearest Carapa procera (Meliaceae; a tree that produces seeds preferred by terrestrial vertebrates) on survival of seeds and seedlings were examined for the animal–dispersed tree species Virola michelii (Myristicaceae) in a mature forest‘at Paracou, French Guiana, in 1992 and 1993. We assessed the putative role of ground–dwelling mammalian herbivores, rodents, and ungulates that filter the seed shadow, acting either as dispersers or predators and thus modifying the original pattern of seed dispersal made by frugivores. We measured the effects of simulated seed burial by rodents using marked seeds and quantified the effect of protecting seeds and seedlings from ground–dwelling vertebrates on seedling germination and survival with fence exclosures in 1992. Dispersal pattern had short–term but no long–term effects on the proportion of V. michelii seeds that survived one year later as seedlings. In the short term, within six weeks, clumped seeds survived better than scattered seeds in both years. Marked seeds that were removed from their site of dispersal were eaten; rodents only rarely buried seeds of V. michelii, and seed burial reduced seed and seedling survivorship. The combined effect of the factors year and Carapa proximity significantly affected seed survival within six weeks. Although six–week seed survival was greater in 1993 than in 1992, seedling establishment was lower in 1993 than in 1992 following a lower rainfall regime during the key period of seed germination (February). One–year seed and seedling survivorship was similar between treatments and years. Seed survival and seedling establishment in V. michelii was dependent on vertebrates in the short term and on climate in the long term. Overall, seed and seedling survivorship depended on a combination of these factors.     

Ectomycorrhizal Networks of Pseudotsuga menziesii var. glauca Trees Facilitate Establishment of Conspecific Seedlings Under Drought

Ectomycorrhizal (EM) networks are hypothesized to facilitate regeneration under abiotic stress. We tested the role of networks in interactions between P. menziesii var. glauca trees and conspecific seedlings along a climatic moisture gradient to: (1) determine the effects of climatic factors on network facilitation of Pseudotsuga menziesii (Mirb.) Franco var. glauca (Mayr) seedling establishment, (2) infer the changing importance of P. menziesii var. glauca parent trees in conspecific regeneration with climate, and (3) parse the competitive from facilitative effects of P. menziesii var. glauca trees on seedlings. When drought conditions were greatest, seedling growth increased when seedlings could form a network with trees in the absence of root competition, but was reduced when unable to form a network. Survival was maximized when seedlings were able to form a network in the absence of root competition. Seedling stem natural abundance δ¹³C increased with drought due to increasing water use efficiency, but was unaffected by distance from tree or network potential. We conclude that P. menziesii seedlings may benefit from the presence of established P. menziesii trees when growing under climatic drought, but that this benefit is contingent upon the establishment of an EM network prior to the onset of summer drought. These results suggest that networks are an important mechanism for EM plants establishing in a pattern consistent with the stress-gradient hypothesis, and therefore the importance of EM networks to facilitation in regeneration of EM trees is expected to increase with drought.     

Limitations on tree seedling establishment across ecotones between abandoned fields and adjacent broad-leaved forests in eastern Japan

Field abandonment has increased over several decades under a long-term recession in Japanese agriculture. To support effective farmland management or reforestation, the patterns of tree recruitment in abandoned fields, which influence management costs, need to be clarified. We investigated tree seedling establishment and microsite variables along forest–field transects in 11 abandoned fields adjacent to secondary broad-leaved forests in the eastern Kanto region of Japan. Generalized linear mixed model analysis indicated that tree seedling establishment in abandoned fields was not correlated with the period of abandonment. Instead, it was positively correlated with vegetation openness. The dominance of a woody vine (Pueraria lobata), an annual liana (Humulus japonicus), and a dwarf bamboo (Pleioblastus chino) mainly explained low vegetation openness within the fields. In addition, the establishment of evergreen tree seedlings and tree seedlings dispersed by hoarding was negatively correlated with the distance from the forest edge. In the forest interior, taller understory vegetation at the edge correlated with lower seedling establishment. These results suggest that seedling establishment across forest–field ecotones is restricted by the dominance of competitive natives within abandoned fields and distance-dependent limitations in both abandoned fields and adjacent forest.     

Association with mature plants protects seedlings from predation in an arid grassland shrub,Gutierrezia microcephala

Summary Survivorship of Gutierrezia microcephala (Compositae) seedlings was studied in an undisturbed arid grassland and in experimental plots where various components of the natural vegetation were removed following seed germination. The major causes of seedling mortality were herbivore damage from the specialist grasshopper, Hesperotettix viridis, and drought stress associated with competition from established plants. The relative intensity of these mortality factors varied strongly with seedling size. Large seedlings had higher overall survivorship but were most likely to be killed by defoliation; most small seedlings died of drought stress.In plots where all perennial grasses were removed (leaving established G. microcephala plants), seedling survivorship was 5 times greater than in undisturbed vegetation. Surviorship in plots where both grasses and mature G. microcephala were removed was slightly poorer than in undisturbed vegetation due to a large increase in mortality from defoliation. From May–August, when the herbivore H. viridis was abundant, seedling survivorship was better in the immediate vicinity of mature conspecific plants than in plots lacking mature G. microcephala, both in the presence and absence of perennial grasses. These results provide a counter example to theories predicting that the impact of specialist herbivores on seedling recruitment is greatest in the vicinity of parent plants.     

Changes in spatial distribution during the life history of a tropical tree,Scaphium macropodum (Sterculiaceae) in borneo

We studied the spatial distribution of fruits and plants, mortality and growth rates ofScaphium macropodum (Sterculiaceae) in four 1-ha plots in a tropical rain forest in West Kalimantan, Indonesia. The species is a large deciduous tree and produces wind-dispersed fruits on defoliated twigs. The density of dispersed fruits on the ground decreased with increasing distance from a parent tree. The area under the parent's crown had the highest density of the fruits and the highest mortality of the seedlings immediately after germination. Consequently, the density of the established seedlings peaked 14 m from the tree which is outside its crown. Thick litter mainly from the parent tree seemed to physically prevent the seedlings' root from reaching the soil surface and caused the high mortality. Juvenile and mature trees distributed exclusively, suggesting that regeneration is the most successful outside of the crown of mature trees. Saplings under canopy shade did not grow well.Scaphium macropodum is hypothesized to require a gap for seedling growth and successful regeneration, whereas it can germinate and last under closed canopies as suppressed seedlings or saplings.     

Dispersion pattern and mortality of seeds and seedlings ofFagus crenata Blume in a cool temperate forest in western Japan

The seed and seedling mortality ofFagus crenata Blume after a mast year (1993) was examined in relation to density and distance from the nearest conspecific adult tree in a mixed conifer-hardwood forest in Ohdaigahara, western Japan. The mortality of fallen seeds during winter amounted to 93.7%, and 79.2% of the current-year seedlings died in the first growing season. The most important factor of death for both seeds and seedlings was predation by vertebrates. The mortality of seeds during winter was positively correlated with sound seed density. The mortality of seedlings was positively correlated with density but not significantly related to the distance from the nearest crown edge of a conspecific adult tree. Mortality patterns varied with stages and spatial scales due to the behavior of predators; it is thus important to investigate the spatial pattern of seeds and seedling mortality at various temporal and spatial scales. After the first growing season, the difference in seedling density between distance classes was not significant at <4m from the nearest adult trees due to density-dependent mortality. However, seedling density was significantly lower in the ≥4 m class than in the <4 m classes.     

Effects of Tree Guards and Weed Mats on the Establishment of Native Tree Seedlings: Implications for Forest Restoration in Hong Kong, China

The effects of tree guards and weed mats on establishment and growth of native tree seedlings, Thick‐leaved oak (Cyclobalanopsis edithiae (Skan) Schott., Fagaceae), planted in an exposed hillside grassland in Hong Kong, were investigated. The natural regeneration of C. edithiae is poor due to a lack of seed dispersal agents and high seed predation, and therefore, this species is often targeted for forest restoration. The experiment lasted for 3.5 years during which the height, basal diameter, and crown diameter of individual seedlings were measured and survivorship recorded. The use of weed mats alone did not have a significant effect, but a combination of tree guards and weed mats led to a significant improvement in establishment, survivorship, and growth of the seedlings during the experimental period. Initially, the guards promoted rapid height growth of the seedlings, although lateral growth and secondary stem thickening were compromised. After the seedlings grew over the tree guards, the basal diameter and crown diameter increased at a notably faster rate. The combined effect of the tree guard and weed mat on the seedling growth pattern was found to be beneficial and contributed to the high survivorship of the seedlings. Comparing the survivorship data and the costs of various treatments, the use of tree guards in combination with the weed mats was demonstrated to be more cost‐effective than planting the seedlings without tree guards or weed mats. The potential for applying the technique in afforestation programs with native tree species for forest restoration in Hong Kong and other tropical regions is discussed.     

Northern Bobwhites and Postfire Succession

ABSTRACT Our study evaluated the effects of prescribed fire on northern bobwhites (Colinus virginianus) occupying native rangelands in Rolling Plains of Texas, USA, during 2002 and 2003. Prescribed fires were conducted during February of 1996, 1998, and 2000; pastures with no recent treatment history served as controls. We quantified bobwhite densities from line transects using distance sampling. We used a repeated-measures analysis of variance to test for treatment-year differences in bobwhite densities. We measured postfire herbaceous and woody vegetation attributes and evaluated vegetation relationships to bobwhite density using simple linear regression. We found significant between-year differences in fall bobwhite densities (F = 13.05, df = 3, P = 0.036) but no differences among treatments or controls. Fall bobwhite densities were inversely related to visual obstruction (r² = 0.179, df = 15, P = 0.058) and positively associated with increasing heterogeneity of grass cover (r² = 0.416, df = 15, P = 0.004). Our results suggest prescribed fire at large spatial scales may be a neutral practice for managing bobwhite habitat on semiarid rangelands.     

Understorey gaps influence regeneration dynamics in subtropical coastal dune forest

Dominant understorey species influence forest dynamics by preventing tree regeneration at the seedling stage. We examined factors driving the spatial distribution of the monocarpic species Isoglossa woodii, a dominant understorey herb in coastal dune forests, and the effect that its cover has on forest regeneration. We used line transects to quantify the area of the forest understorey with I. woodii cover and with gaps in the cover. Paired experimental plots were established in semi-permanent understorey gaps with I. woodii naturally absent and in adjacent areas with I. woodii present to compare plant community composition, soil, and light availability between the two habitats. Isoglossa woodii was widespread, covering 65–95% of the understorey, while gaps covered the remaining 5–35% of the area. The spatial distribution of this species was strongly related to tree canopy structure, with I.␣woodii excluded from sites with dense tree cover. Seedling establishment was inhibited by low light availability (<1% of PAR) beneath I.␣woodii. When present, I. woodii reduced the density and species richness of tree seedlings. The tree seedling community beneath I. woodii represented a subset of the seedling community in gaps. Some species that were found in gaps did not occur beneath I. woodii at all. There were no significant differences between the sapling and canopy tree communities in areas with I. woodii gaps and cover. In the coastal dune forest system, seedling survival under I. woodii is dependent on a species’ shade tolerance, its ability to grow quickly during I. woodii dieback, and/or the capacity to regenerate by re-sprouting and multi-stemming. We propose a general conceptual model of forest regeneration dynamics in which the abundant understorey species, I. woodii, limits local tree seedling establishment and survival but gaps in the understorey maintain tree species diversity on a landscape scale.     

Reproductive characteristics in an understory bamboo and gradual environmental changes after its dieback provide an extended opportunity for overstory tree regeneration in a mixed cool-temperate forest in central Japan

• The reproductive characteristics of understory bamboo and the effects of dieback on overstory tree seedlings through temporal changes in the environment at the forest floor have only been examined in a few bamboo species, due to the unpredictable occurrence of flowering events and long intervals between them but provide valuable information on tree regeneration and succession in a forest with dense dwarf bamboo cover.
• We investigated environmental conditions and assessed seedlings (< 30-cm tall) of the dwarf bamboo Sasa borealis and overstory tree species at 44–50 measurement points during 2016–2021, which included a S. borealis mass flowering event in 2017. We also conducted seed germination tests to determine germination rates and patterns in S. borealis. Environmental factors affecting seedling recruitment of S. borealis and of overstory trees were analysed using spatiotemporal generalized linear mixed models in the Bayesian framework.
• We observed gradual temporal changes in the environment, including increasing canopy openness and decreasing maximum height of dead S. borealis culms. The seeds germinated slowly and the emergence of current-year S. borealis seedlings peaked in spring–summer in 2019. The tree seedling density after 2019 increased significantly compared to that before the dieback. The model results suggest that tree seedling establishment was enhanced by increased light availability.
• Continuous field observation beginning before S. borealis dieback revealed gradually enhanced tree recruitment in response to slow decay of the remaining dead culms and slow recovery of S. borealis. The seedling regeneration pattern of understory bamboo partly contributes to a prolonged opportunity for overstory tree regeneration.

     

Differential seedling regeneration patterns across forest–grassland ecotones in two tropical treeline species (Polylepis spp.)

Successful forest expansion into grassland can be limited by seed dispersal and adverse conditions for tree seedlings in the grassland environment. In the high‐elevation Andes, human‐induced fragmentation has exacerbated the patchy distribution of Polylepis forests, threatening their unique biological communities and spurring restoration interest. Studies of Polylepis forest extent in Peru suggest that forest borders have remained stable over the past century despite decreasing anthropogenic disturbance, suggesting that tree seedling recruitment is being limited in the open grassland habitat. We studied natural seedling dispersion patterns of Polylepis sericea and Polylepis weberbaueri (Rosaceae) at forest–grassland edges across a range of environmental conditions to examine seedling recruitment and colonization of grasslands in Huascaran National Park (Peru). Using data from 2367 seedlings found in 48 forest–grassland edge plots (15 m × 15 m) at forest patches between 3900–4500 masl, we employed generalized mixed modelling to identify the significant associations of seedling densities with environmental covariates. In addition, we compared these associations to patterns of adult presence on the landscape. Seedling densities were associated with a combination of variables varying within (distance to forest edge) and among (elevation and dry season solar irradiation) plots across the landscape. For both species, seedling densities decreased with increasing distance away from the forest in a manner consistent with short‐distance seed dispersal by wind. Our results suggest that such short‐distance dispersal may slow forest expansion, but that there also appear to be substantial post‐dispersal limitations to seedling establishment in the grassland. Polylepis sericea densities decreased with elevation, while P. weberbaueri increased with elevation and decreased with solar irradiation. Associations of adult presence with elevation and solar irradiation mirrored those of seedling densities. Management of areas with forest patches dominated by these species should consider these differences in their environmental tolerances, particularly during species selection and zonation for reforestation.