Experiments on the mechanism of tree and shrub establishment in temperate grassy woodlands: Seedling emergence

Field experiments were designed to examine tree and shrub seedling emergence in temperate grassy woodlands on the New England Tablelands. The effects of study sites, intensity of previous grazing, removal of ground cover by fire or clearing, burial of seeds and ant seed theft on seedling emergence were tested in two field experiments. Six tree and seven shrub species were used in the experiments and their cumulative emergence was compared with laboratory germination studies. All species used in field experiments had lower cumulative emergence than those in laboratory germination studies despite prolonged periods of above average rainfall before and after seeds were sown. Eucalypt species emerged faster in the field than the shrub species and generally attained higher cumulative emergence than the shrubs. Spatial effects of sites and patches within sites, and of previous grazing history did not strongly influence patterns of seedling emergence in most species. Ground and litter cover generally did not enhance or suppress the emergence of seedlings, although the removal of cover in recently grazed areas enhanced the emergence of some species. Burning enhanced the emergence of some tree and shrub species where plots had more fuel and intense fires, but this effect was not strong. Compared with other treatments, seedbed manipulations produced the strongest effects. In the absence of both invertebrate and vertebrate predators, seedling emergence was lower for surface‐sown seed, compared with seed sown on scarified soil surfaces. Higher seedling emergence of buried seeds in the presence of invertebrate predators probably resulted from the combined effects of predator escape and enhanced moisture status of the germination environment. Some promotion of emergence was achieved for all species in most sown treatments probably as a result of a prolonged above average rainfall. In contrast, the natural recruitment of trees and shrubs was negligible in experimental plots, highlighting the importance of seed supply and dispersal as ultimate determinants of recruitment.     

Emergence and survival of herbaceous seedlings in temperate grassy woodlands: Recruitment limitations and regeneration niche

Abstract Field experiments examined herbaceous seedling emergence and survival in temperate grassy woodlands on the New England Tablelands of New South Wales. Effects of intensity of previous grazing, removal of ground cover by fire or clearing, burial of seeds, grazing and seed theft by ants on seedling emergence and survival were studied in two field experiments. Thirteen species with a range of traits were used in the experiments and their cumulative emergence was compared with laboratory germination studies. Field emergence correlated to laboratory germination but all species had lower emergence in the field. Little natural emergence of native species was observed in the field in unsown treatments. Short‐lived forbs had the highest emergence, followed by perennial grasses; rhizomatous graminoids and perennial forbs had the lowest emergence. Soil surface and cover treatments did not markedly enhance emergence suggesting that intertussock spaces were not prerequisites for forb emergence. No consistent pattern of enhanced emergence was found for any treatment combination across all species. Seedling survival varied among species, with perennial grasses and short‐lived forbs having the highest seedling mortality. Low mortality rates in the graminoids and rhizomatous forbs appeared partially to compensate for lower seedling emergence. All perennial grasses and some short‐lived forbs showed increased risk of mortality with grazing. Differences in emergence and survival of species were related to ground cover heterogeneity, soil surfaces and, to some extent, herbivory. The complexity of these patterns when superimposed on temporal variability suggests that no generalizations can be made about the regeneration niche of herbaceous species groups. Strong recruitment limitation and partitioning of resources in the regeneration niche may reduce competition among native species and explain the high species richness of the herbaceous layer in the temperate grassy communities of eastern Australia.     

Fire frequency effects in a grassy woodland: Trees and grasses

Our study investigated the effects of fire frequency on the tree and grass components of Cumberland Plain Woodland, an endangered grassy eucalypt woodland in eastern Australia. We located three sites within each of three fire frequency classes (high, moderate, low) based on fire history and a similar time since last fire. For trees, we asked whether fire frequency affected density, eucalypt population structure, basal area and spatial patterning. For grasses, we tested for fire frequency effects on total abundance and the abundance of two common species. Density of trees was not significantly affected by fire frequency for juveniles, saplings or adult trees and neither was basal area per hectare. Features of a recruitment bottleneck model were present for eucalypts. There was a large pool of suppressed juveniles making up more than half the population at each fire frequency. Saplings were the smallest group and were susceptible to fire‐induced stem mortality, with particularly low numbers relative to juveniles and small trees where fire was frequent or intense. Despite this difference in sapling mortality, numbers of small trees did not differ with fire frequency. Saplings could transition to small trees in equal numbers if they did so more quickly at high than at lower fire frequencies, and if recruitment of saplings into the tree layer was controlled independently of fire frequency. The size hierarchy of small to medium eucalypt trees was homogeneous over eight of the nine sites, spatial patterning of adult trees was random tending to regular, and mean tree size decreased with density at all sites. These features of eucalypt population structure are indicative of possible resource competition which could regulate tree recruitment. Total grass cover index was high across all fire frequencies, with Themeda triandra dominating at high and moderate fire frequencies and Microlaena stipoides at low fire frequency.     

Differences in tree and shrub establishment due to tree guard type in a temperate upland pasture

Success of establishing native trees in cool temperate environments depends on the ability of seedlings to withstand subzero temperatures and recurrent frosts. This study compared the survival and growth of five tree and shrub species with two guard types at three landscape positions in an upland pasture. Seedlings were planted between December 2013 and March 2014. Half of the seedlings were planted in tall Corflute^® guards (60 cm high), and the remaining seedlings were interplanted in milk cartons (30 cm). Seedling survival and height were measured in November 2014. Hourly temperature readings were recorded between March and November 2014. Seedling height for all species was greater in tall guards than milk cartons at all landscape positions, possibly at least partly due to etiolation. However, seedlings in tall guards survived better than seedlings in milk cartons at mid‐ and upper‐slope positions. Higher temperatures may have benefited seedling performance by prolonging the growing season as average maximum temperature was significantly higher inside tall guards than milk cartons and ambient conditions at all landscape positions. Average daily temperature was significantly higher in tall guards compared to milk cartons and ambient conditions at the upper‐slope site only. There were no significant differences in average minimum temperature between guard types at all landscape positions.     

Fire season and intensity affect shrub recruitment in temperate sclerophyllous woodlands

The season in which a fire occurs may regulate plant seedling recruitment because of: (1) the interaction of season and intensity of fire and the temperature requirements for seed release, germination and growth; (2) post-fire rainfall and temperature patterns affecting germination; (3) the interaction of post-fire germination conditions and competition from surrounding vegetation; and (4) the interaction of post-fire germination conditions and seed predators and/or seedling herbivores. This study examined the effects of different fire intensities and fire seasons on the emergence and survival of shrubs representing a range of fire response syndromes from a summer rainfall cool climate region. Replicated experimental burns were conducted in two seasons (spring and autumn) in 2 consecutive years and fuel loads were increased to examine the effects of fire intensity (low intensity and moderate intensity). Post-fire watering treatments partitioned the effects of seasonal temperature from soil moisture. Higher intensity fires resulted in enhanced seedling emergence for hard-seeded species but rarely influenced survival. Spring fires enhanced seedling emergence across all functional groups. Reduced autumn recruitment was related to seasonal temperature inhibiting germination rather than a lack of soil moisture or competition. In Mediterranean-type climate regions, seedling emergence has been related to post-fire rainfall and exposure of seeds to seed predators. We think a similar model may operate in temperate summer rainfall regions where cold-induced dormancy over winter exposes seeds to predators for a longer time and subsequently results in recruitment failure. Our results support the theory that the effect of fire season is more predictable where there are strong seasonal patterns in climate. In this study seasonal temperature rather than rainfall appears to be more influential.     

Fire frequency influences composition and structure of the shrub layer in an Australian subcoastal temperate grassy woodland

Little is known about the relationship between fire regimes and plant diversity in Australia's temperate grassy woodlands. The effect of fire frequency on shrubs in grassy woodland remnants across Western Sydney's Cumberland Plain was examined. Shrub species richness and composition were compared in sites that had experienced a high, moderate or low frequency of fire over the previous 20 years. Nine sites were surveyed, three in each fire frequency category; most sites, including all low‐fire‐frequency sites, had burnt 9–36 months prior to sampling. Fire frequency had a profound effect on the composition and structure of the shrub layer. Per cent frequency and density of the prickly shrub Bursaria spinosa (Pittosporaceae) was considerably higher in low‐fire‐frequency sites than where fires had occurred at least once a decade. In sites where fire had been absent for decades prior to a recent fire, this species dominated the landscape, while elsewhere it occurred as clumps in a grassy matrix. Per cent frequency of other native shrubs, particularly obligate seeders, was greatest at moderate fire frequencies. Exotic shrubs were recorded most often where fire had been rare. While ordination clearly separated out the low‐fire‐frequency sites, complete separation between high‐ and moderate‐fire‐frequency blocks was not achieved. The increase in Bursaria in the absence of fire mirrors the encroachment of woody plants into a range of grassy ecosystems around the world. The sensitivity of obligate seeder species, many of them short‐lived legumes with fire‐cued seeds, to both very frequent and very infrequent fire shows the vulnerability of these species to extreme fire regimes, despite the safeguards conferred by hard‐seededness. Competition from Bursaria, as well as loss of viable seed in the soil, may have contributed to the low frequency of these species after a long inter‐fire interval.     

Effects of herbivory on growth and survival of seedlings of a rainforest tree,Alphitonia whitei (Rhamnaceae)

This study examined the effects of insect herbivory on growth and mortality of seedlings of a mid- successional rainforest tree, Alphitonia whitei Braid. Two experiments were conducted in which seedlings were exposed to 0% and 50% natural defoliation by insect herbivores and placed in light gaps in simple notophyll vine forest at Paluma, near Townsville, North Queensland. In the first experiment, insect herbivory significantly increased mortality of 2-month-old seedlings. Smaller seedlings had significantly greater mortality rates than larger seedlings, irrespective of herbivory. A significantly greater proportion of smaller seedlings died from being smothered by fallen leaves and soil as a result of digging by vertebrates than for larger seedlings. In the second experiment, the effects of seedling age were examined by comparing 2-month and 4-month-old seedlings. Mortality rates were significantly influenced by seedling age, with eight times greater survival of older seedlings than of younger seedlings. Although insect herbivory was correlated with a significant decrease in shoot mass and a significant increase in root:shoot ratio, there was no effect of insect herbivory on seedling survival in the second experiment. Thus, mortality rates were greater for seedlings if they were young or small (which indirectly results from insect herbivory), because small, young seedlings were more susceptibile to other mortality factors, such as burial by fallen debris and digging by vertebrates.     

Early emergence increases survival of tree seedlings in Central European temperate forests despite severe late frost

Global warming is expected to result in earlier emergence of tree seedlings that may experience higher damages and mortality due to late frost in spring. We monitored emergence, characteristics, and survival of seedlings across ten tree species in temperate mixed deciduous forests of Central Europe over one and a half year. We tested whether the timing of emergence represents a trade‐off for seedling survival between minimizing frost risk and maximizing the length of the growing period. Almost two‐thirds of the seedlings died during the first growing period. The timing of emergence was decisive for seedling survival. Although seedlings that emerged early faced a severe late frost event, they benefited from a longer growing period resulting in increased overall survival. Larger seedling height and higher number of leaves positively influenced survival. Seedlings growing on moss had higher survival compared to mineral soil, litter, or herbaceous vegetation. Synthesis. Our findings demonstrate the importance of emergence time for survival of tree seedlings, with early‐emerging seedlings more likely surviving the first growing period.     

Seedling establishment of Acacia tortilis and Hyphaene compressa in the Turkwel riverine forest, Kenya

This study reports on the spatial and temporal patterns of seedling establishment in the Turkwel riverine forest, Kenya. Seedlings of the dominant tree species Acacia tortilis and Hyphaene compressa were mapped and monitored to assess the underlying causes of seedling recruitment and mortality. The broad‐scale distribution of A. tortilis was not correlated with any environmental variables, while H. compressa seedlings were confined to flood‐exposed sites in the arid downstream section of the floodplain. One year of monitoring showed that seedling recruitment of A. tortilis was evoked by prolonged rainfall, while seedling mortality was caused by desiccation, browsing and trampling. In contrast, seedling recruitment and mortality of H. compressa was largely unaffected by rainfall and livestock, probably due to the high moisture requirements of seeds and the tolerance of seedlings to disturbance. There were no effects of soils, light, or seedling density on the establishment of A. tortilis and H. compressa seedlings. This study demonstrates the importance of parallel mapping and monitoring of riverine seedlings in order to understand patterns and processes of forest regeneration in arid and semi‐arid floodplains.     

Effects of sika deer on tree seedlings in a warm temperate forest on Yakushima Island,Japan

High-density herbivore species often play an important role in forest regeneration. Native sika deer (Cervus nippon yakushimae) inhabit a high density (51.5–63.8head/km², estimated by a pellet count method) area in the western part of a lowland natural forest on Yakushima Island, Japan. To test experimentally the impact of sika deer on the mortality and the survivability of current-year seedlings, which are at a more vulnerable stage than the later stages, we constructed fenced exclosures, planted seeds of nine sapfruit tree species and compared the mortality and the survivability of current-year seedlings between fenced and unfenced quadrats. Large seeded species had significantly greater survivability in fenced quadrats than in unfenced quadrats. However, the survivability disagreed with feeding preferences. Sika deer activity increased seedling mortality of large-seeded species more than that of small-seeded species, and did not decrease much seedling survivability of not-preferred species. We found that the physical disturbance by the high density of sika deer resulted in mortality for both preferred and not-preferred species, and that deer herbivory was important for preferred species.     

鼠类对山杏种子存活和萌发的影响

山杏（Prunus armeniaca)是广布于北京山区阳坡的矮林或灌丛,易生存在土壤贫脊、干旱、或严重退化的山坡、裸地,对于保护生态环境,减少水土流失及促进山区经济发展均具有重要作用。有关鼠类在山杏更新中作用的研究较少,本研究主要是确定影响山杏更新的鼠类种类、鼠类对山杏种子的取食压力、人工埋藏种子及植被因素对种子存活和萌发的影响。于1997年10月3－4日,采用夹捕法同时调查取食花生和山杏种子的鼠类种类及种群密度。于1997年9月24－25日,通过在地表放置山桔种子,调查在鼠类取食下,地表种子消失的速度。于1997年10月20日,作种子埋藏实验,调查种子存活及萌发情况。于1997年10月26日,进行扣网实验,研究网内、外山杏种子存活及萌发情况。结果表明：大林姬鼠（Apodemus speciosus)、社鼠（Rattus confucianus),黑线姬鼠（Apodemus agrarius)为取食山杏种子的主要鼠种种类;与取食花生相比,社鼠更偏爱山杏种子,大林姬鼠对山杏种子喜食性略低,黑线姬鼠对二者喜食程度相近;山杏种子放置地表后1、10、20、30d的消失率分别为13.06%,64.46%,90.70%,96.69%;扣网实验说明,山杏在地表的萌发率极低,生长也慢;而埋入土层内可明显提高萌发率和生长率（若除去鼠类的取食,萌发率还要高）;人工埋藏实验表明,将山杏种子埋入土层5cm后,能够有效地减少鼠类对种子的取食;通过分析山杏种子萌发与植被的关系,发现山杏易在开阔、阳光充分的草丛中存活和生长,而不易在阴闭的灌丛下存活和生长。     

Interspecific and intraspecific variation in tree seedling survival: effects of allocation to roots versus carbohydrate reserves

We examined interspecific and intraspecific variation in tree seedling survival as a function of allocation to carbohydrate reserves and structural root biomass. We predicted that allocation to carbohydrate reserves would vary as a function of the phenology of shoot growth, because of a hypothesized tradeoff between aboveground growth and carbohydrate storage. Intraspecific variation in levels of carbohydrate reserves was induced through experimental defoliation of naturally occurring, 2-year-old seedlings of four northeastern tree species –Acer rubrum, A. saccharum, Quercus rubra, and Prunus serotina– with shoot growth strategies that ranged from highly determinate to indeterminate. Allocation to root structural biomass varied among species and as a function of light, but did not respond to the defoliation treatments. Allocation to carbohydrate reserves varied among species, and the two species with the most determinate shoot growth patterns had the highest total mass of carbohydrate reserves, but not the highest concentrations. Both the total mass and concentrations of carbohydrate reserves were significantly reduced by defoliation. Seedling survival during the year following the defoliation treatments did not vary among species, but did vary dramatically in response to defoliation. In general, there was an approximately linear relationship between carbohydrate reserves and subsequent survival, but no clear relationship between allocation to root structural biomass and subsequent survival. Because of the disproportionate amounts of reserves stored in roots, we would have erroneously concluded that allocation to roots was significantly and positively related to seedling survival if we had failed to distinguish between reserves and structural biomass in roots. Received: 14 December 1999 / Accepted: 2 June 1999     

Composition of grazed and cleared temperate grassy woodlands in eastern Australia: patterns in space and inferences in time

Abstract. A regional vegetation survey of the temperate grassy woodlands (temperate savanna) in Australia was designed to assess the effects of clearing and grazing on the composition of vegetation remnants and the adjacent pasture matrix. Vegetation was sampled across a range of habitats using 77 0.1024‐ha quadrats; the relative abundance of species was recorded. Classification analysis clustered the sites into three main groups that corresponded to intensity of grazing/clearing followed by groups based on underlying lithology (basalt, metasediment, granites). Using Canonical Correspondence Analysis, exogenous disturbance and environmental variables were related to the relative abundance of species; grazing intensity had the highest eigenvalue (0.27) followed by tree canopy cover (0.25), lithology (0.18), altitude (0.17) and slope (0.10). Based on two‐dimensional ordination scores, six species response groups were defined relating to intensity of pastoralism and nutrient status of the landscape. Abundance and dominance of native shrubs, sub‐shrubs, twiners and geophytes were strongly associated with areas of less‐intense pastoralism on low‐nutrient soils. The strongest effects on species richness were grazing followed by canopy cover. Continuously grazed sites had lower native species richness across all growth forms except native grasses. There was no indication that intermediate grazing intensities enhanced forb richness as a result of competitive release. Species richness for all native plants was lowest where trees were absent especially under grazed conditions. Canopy cover in ungrazed sites appeared to promote the co‐existence of shrubs with the herbaceous layer. Predicted declines in forb richness in treeless, ungrazed, sites were not detected. The lack of a disturbance‐mediated enhancement of the herbaceous layer was attributed to habitat heterogeneity at 0.1 ha sampling scale.     

Effects of local biotic neighbors and habitat heterogeneity on tree and shrub seedling survival in an old-growth temperate forest

Seedling dynamics play a crucial role in determining species distributions and coexistence. Exploring causes of variation in seedling dynamics can therefore provide key insights into the factors affecting these phenomena. We examined the relative importance of biotic neighborhood processes and habitat heterogeneity using survival data for 5,827 seedlings in 39 tree and shrub species over 2?years from an old-growth temperate forest in northeastern China. We found significant negative density-dependence effects on survival of tree seedlings, and limited effects of habitat heterogeneity (edaphic and topographic variables) on survival of shrub seedlings. The importance of negative density dependence on young tree seedling survival was replaced by habitat in tree seedlings ??4?years old. As expected, negative density dependence was more apparent in gravity-dispersed species compared to wind-dispersed and animal-dispersed species. Moreover, we found that a community compensatory trend existed for trees. Therefore, although negative density dependence was not as pervasive as in other forest communities, it is an important mechanism for the maintenance of community diversity in this temperate forest. We conclude that both negative density dependence and habitat heterogeneity drive seedling survival, but their relative importance varies with seedling age classes and species traits.