Experiments on tree and shrub establishment in temperate grassy woodlands: Seedling survival

Abstract Experimental studies of the emergence of shrubs and trees in grassy woodlands on the New England Tablelands, New South Wales, Australia, showed that emergence of seedlings was determined by seed supply, seed predators and seed burial. The survival of these seedlings was then observed in an experiment to test the effects of previous land use, grazing by stock and grazing by other vertebrates. The fate of four eucalypts and six shrub species was followed over 5 years. Across all species more than 50% mortality occurred in the first 6 months prior to the imposition of grazing treatments. These deaths were attributed to the combined effects of insect defoliation, cold, and low soil moisture. Average mortality over all treatments showed two distinct trends: eucalypts and one unpalatable shrub (Leptospermum) had greater than 1% survival over 5 years, whereas Acacia, Cassinia, Indigophera, Lomatia and Xanthorrhoea either had very low or no survival after 5 years. The effect of livestock grazing on seedling numbers was rarely detected because of patchy emergence and mortality due to other causes. However, proportional hazard regression models showed that there was often an increased hazard associated with grazing or grazed landscapes. Overall, those species with high hazard coefficients associated with stock are rare in the landscape, whereas those with lesser risk are more common. Recruitment is likely to be an extremely rare event because the highest proportion of germinable seed sown that survived to a juvenile stage was 0.42% and the mean across all species was 0.12%. No natural recruitment of shrub species was observed over 5 years of observation, suggesting that recruitment is episodic and disturbance driven. Enhancing natural ‘regeneration’ of woody plants under these circumstances may be more challenging than simply fencing off remnants.     

Biomass dynamics of herbaceous vegetation in Zambezi riverine woodlands

Seasonal changes in biomass, productivity and large mammal utilization of annual grasslands on Zambezi alluvium in Mana Pools National Park, Zimbabwe, were recorded over four years and compared with perennial grasslands. Annual grasses grew only in the wet season, but forb productivity peaked at the start of the dry season. Large mammals and termites ate dead annual grasses in the dry season. Production was usually greater in perennial grasslands, than in annual ones, and perennial grasses often grew throughout the year. Large mammal utilization of grasslands varied between 53 and 99% of annual production.

Résumé

L'on rapporte les changements saisonniers de biomasse, productivité et utilisation par les grands mammifères des herbages annuels dans les plaines alluviales du Zambèze au Parc National de Mana Pools, Zimbabwe, pendant 4 ans et on les compare avec ceux des herbages pérennes. Les herbes annuelles ne croissent que pendant la saison des pluies mais la productivité du fourrage était au maximum en début de pérennes que dans les annuels et les herbes pérennes poussaient souvent tout au long de I'année. L'utilisation par les grands mammifères variait de 53 à 99% de la production annuelle.     

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.     

林窗对格氏栲天然林更新层物种生态位的影响

采用改进更新生态位宽度和更新生态位重叠模型分析林窗干扰对福建三明市格氏栲天然林更新层物种生态位的影响。结果表明:林窗中格氏栲更新生态宽度值大于林下,林窗在促进格氏栲更新过程中具有重要作用。林窗和林下更新生态位宽度最大的为桂北木姜子,其与格氏栲更新生态位重叠值也较大。林窗中的格氏栲与其它物种更新生态位重叠值低于0.6,林窗微生境的异质性导致格氏栲与其它树种间对资源的利用存在明显的共享趋势,促进了物种间共存。林下的格氏栲与木荷、木荚红豆和短尾越桔的更新生态位重叠值均高于0.6,格氏栲与这些物种相互争夺资源与空间,种间竞争较强。格氏栲天然林未来树种组成中,主要由桂北木姜子、木荷、矩圆叶鼠刺与格氏栲等组成的混交群落,整个群落正向物种组成多样化的方向演变。     

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.     

Taxonomic scale-dependence of habitat niche partitioning and biotic neighbourhood on survival of tropical tree seedlings

In order to differentiate between mechanisms of species coexistence, we examined the relative importance of local biotic neighbourhood, abiotic habitat factors and species differences as factors influencing the survival of 2330 spatially mapped tropical tree seedlings of 15 species of Myristicaceae in two separate analyses in which individuals were identified first to species and then to genus. Using likelihood methods, we selected the most parsimonious candidate models as predictors of 3 year seedling survival in both sets of analyses. We found evidence for differential effects of abiotic niche and neighbourhood processes on individual survival between analyses at the genus and species levels. Niche partitioning (defined as an interaction of taxonomic identity and abiotic neighbourhood) was significant in analyses at the genus level, but did not differentiate among species in models of individual seedling survival. By contrast, conspecific and congeneric seedling and adult density were retained in the minimum adequate models of seedling survival at species and genus levels, respectively. We conclude that abiotic niche effects express differences in seedling survival among genera but not among species, and that, within genera, community and/or local variation in adult and seedling abundance drives variation in seedling survival. These data suggest that different mechanisms of coexistence among tropical tree taxa may function at different taxonomic or phylogenetic scales. This perspective helps to reconcile perceived differences of importance in the various non-mutually exclusive mechanisms of species coexistence in hyper-diverse tropical forests.     

The regeneration niche of the grey mangrove (Avicennia marina): effects of salinity,light and sediment factors on establishment,growth and survival in the field

Field observations of seedlings and saplings of Avicennia marina showed patterns that correlated with salinity, light and sediment. Models that account for these observations were subsequently tested in a series of field experiments. Establishment varied within an estuary under controlled conditions but was not related to salinity or sediment type. Seedling survival was uniform over 3 years regardless of position in estuary and sediment type. Seedling densities and survival under canopies or in canopy gaps were not significantly different. However, seedling growth and density of saplings were greater in canopy gaps. Experiments involving manipulations of canopies showed no differences in seedling survival under canopies or in light gaps, but addition of slow-release fertilizer enhanced growth and survival in canopy gaps and under canopies. Long-term comparison of areas denuded of a canopy and with sediment disturbance showed enhanced establishment and survival when compared with areas with canopy gaps but with undisturbed sediments. Overall there appears to be no restriction to establishment of propagules within mangrove stands other than the supply of propagules and tidal or wave action. In contrast, recruitment to the sapling stage appears to be restricted by light and sediment resources. We suggest that propagules need to establish in a regeneration niche for seedling recruitment to the sapling stage. This differs from the view that seedlings in the under-storey are analogous to a seed pool in the soil.     

Performance of seedlings of Mediterranean woody species under experimental gradients of irradiance and water availability: trade-offs and evidence for niche differentiation

The aim of the study was to assess the potential importance for Mediterranean plants of trade-offs in the response to irradiance and water availability at the regeneration stage. Survival and growth patterns across an experimentally imposed irradiance gradient (1, 6, 20 and 100% sunlight) were studied in seedlings of eight Mediterranean woody species, together with the impact of a simulated summer drought. We found evidence of some of the trade-offs previously reported for non-Mediterranean plant communities, such as between survival in the shade and relative growth rate (RGR) at high light, but no evidence for others, such as between shade and drought tolerances. The impact of drought on survival and RGR was stronger in high light than in deep shade. The observed species-specific differences in performance provide a mechanistic basis for niche differentiation at the regeneration stage, contributing to possible explanations of species coexistence in Mediterranean ecosystems.     

Distribution and regeneration strategies of major canopy dominants in species-rich subtropical/warm temperate rainforests in south-western Japan

The distribution and regeneration strategy of the major canopy dominants in species-rich subtropical/warm temperate rainforests in south-western Japan was studied in a hilly zone below 1100 m a.s.l. Using the patch sampling method, four dominance-community types were numerically identified and they corresponded to four habitats which represented a combination of topography and altitude (i.e. ridgesvs slopes, and belowvs above 500 m a.s.l.) Seven major canopy dominants had their respective distributional core in one of these topo-altitudinal habitats. The seven canopy dominants could be classified into three species groups according to their size structure and growth habits. Group A (Tsuga sieboldii andQuercus gilva) and group B (Quercus acuta, Quercus salicina, Machilus thunbergii) were restricted to the emergent and canopy layer, respectively. These two groups had only a few subcanopy trees and saplings. Saplings of group B showed a rapid growth rate in canopy gaps. Group C (Distylium racemosum) was characterized by many subcanopy trees and saplings that grew steadily under the closed canopy.Castanopsis sieboldii showed intermediate characteristics between group B and C in the size structure and growth habit of its saplings. The density ofD. racemosum canopy trees was markedly reduced on ridges and slopes above 500 m and on slopes below 500 m. In these marginal habitats, the three species groups coexisted by sharing different strata within a community. This situation was possible due to the differences in regeneration strategies among the canopy species.     

Evolutionary history and distance dependence control survival of dipterocarp seedlings

One important hypothesis to explain tree-species coexistence in tropical forests suggests that increased attack by natural enemies near conspecific trees gives locally rare species a competitive advantage. Host ranges of natural enemies generally encompass several closely related plant taxa suggesting that seedlings should also do poorly around adults of closely related species. We investigated the effects of adult Parashorea malaanonan on seedling survival in a Bornean rain forest. Survival of P. malaanonan seedlings was highest at intermediate distances from parent trees while heterospecific seedlings were unaffected by distance. Leaf herbivores did not drive this relationship. Survival of seedlings was lowest for P. malaanonan , and increased with phylogenetic dissimilarity from this species, suggesting that survival of close relatives of common species is reduced. This study suggests that distance dependence contributes to species coexistence and highlights the need for further investigation into the role of shared plant enemies in community dynamics.
Ecology Letters (2010) 13: 51–59     

广西大石围天坑群草本植物多样性及其生态位变化规律

研究不同发育时期天坑草本植物的多样性,能更好的理解草本层对天坑环境变化的响应机制。以广西大石围天坑群中不同发育时期天坑(初期、中期和后期)为研究对象,采用空间代替时间的方法,分析不同发育时期天坑草本植物多样性和生态位变化规律。结果表明：(1)广西大石围天坑群调查样地共记录到草本植物34科71属98种,以鳞毛蕨科、荨麻科、凤尾蕨科、菊科和禾本科为主。(2)后期天坑草本植物的Pielou′s指数和Shannon-Wiener指数均显著高于初期天坑(P<0.05),表明随着天坑发育到后期,草本层物种数逐渐增多,物种分布更为均匀。(3)随着天坑的发育,草本层喜阴植物重要值下降,喜阳、半阴植物重要值上升,喜阳植物逐渐取代喜阴植物在群落中占据优势地位,草本层将经历喜阴植物—喜阴、半阴植物—喜阳植物的三个生态型转变过程。(4)各发育时期优势草本生态位相似性比例和生态位重叠平均值均<0.5,表明天坑不同发育时期对草本植物的生态位影响较小,各时期草本层总体呈现生态分化明显、种间竞争较弱的特点。(5)各发育时期优势草本生态位重叠值大、相对高度差小的种对数中,不同生态习性的种对数占比不同,中期天...     

Biophysical and human influences on plant species richness in grasslands: Comparing variegated landscapes in subtropical and temperate regions

A survey of grassy woodlands in the Queensland subtropics was conducted, recording herbaceous species richness at 212 sites on three properties (2756 ha). A range of habitats typical of cattle grazing enterprises was sampled and site variables included lithology, slope position, tree density, soil disturbance, soil enrichment and grazing. Results were compared with a previously published survey of temperate grasslands. Lithology, slope position and tree density had relatively minor effects on plant species richness, although in both surveys there was some evidence of lower species richness on the more fertile substrates. Soil disturbance and soil enrichment significantly reduced the richness of native species in both surveys, while exotic species were insensitive (subtropics) or increased (temperate) with disturbance. Rare native species were highly sensitive to disturbances, including grazing, in the temperate study. Although some trends were similar for rare species in the subtropics, the results were not significant and there were complex interactions between grazing, lithology and slope position. Grazing did not have a negative effect on native species richness, except in the closely grazed patches within pastures, and then only on the most intensively developed property. At the scale recorded (30 m²), the native pastures, roadsides and stock routes sampled in the subtropics appear to be among the most species‐rich grasslands ever reported, both nationally and globally. Native species richness was approximately 50% higher than the temperate survey figures across all the comparable habitats. While there are no clear reasons for this result, potential explanations are proposed.     

三峡库区消落带现存草本植物组成与生态位

为揭示经历长期水位变动后,三峡水库消落带优势植物对可利用资源的分享状况、不同资源环境下植物种群生态位特征以及植物种间竞争和共存机制,对三峡库区秭归段典型消落带现存优势草本物种的种群空间格局及生态位特征进行研究.结果表明: 该区共有草本植物39种,隶属18科32属,以禾本科、菊科、蓼科和大戟科植物为主;狗牙根、狗尾草、狼把草和毛马唐具有较大的重要值和生态位宽度,为主要优势种;海拔145～155 m段植物生态位宽度前3位依次为狗牙根＞酸模叶蓼＞狗尾草;海拔155～165 m段植物生态位宽度前3位依次为狗尾草＞毛马唐＞狗牙根;海拔165～175 m段植物生态位宽度前3位依次为狗尾草＞狼把草＞红蓼;不同海拔物种间生态位重叠均保持较高状态,生态位宽度较小物种也可与生态位宽度较大物种有较大的生态位重叠值,仅凭生态位宽度不能判定生态位重叠值.消落带经历7次水位涨落周年后,大多数优势物种为一年生草本植物,生态位分化程度较低,在资源匮乏且不稳定的生境中种间竞争强烈,消落带植被仍处于群落演替的初级阶段.     

Recruitment and survival of Argyrodendron actinophyllum seedlings in an Australian rainforest

The capacity of seedlings to survive for extended periods beneath intact forest increases the likelihood of regeneration of many species of canopy trees in rainforests. I studied the demographics of Argyrodendron actinophyllum (F.M.Bail.) H.L.Edlin seedlings in a subtropical rainforest in northern New South Wales. A mast seeding of A. actinophyllum was observed and subsequent survival of seedlings monitored over a four year period. Densities of seedlings that emerged correlated with seedfall, while seedfall depended on the size and distance to the surrounding trees. Mortality of seedlings showed density-dependence at higher seedling densities (above about 100 seedlings m^?2), apparently in response to browsing pressure that varied with the density of seedlings. Seedlings that were protected from vertebrates by exclosure cages had lower mortality rates than unprotected seedlings and showed no density response. Glasshouse experiments showed seedling growth was reduced by defoliation, light intensity and initial seed weight, and that seedlings could not persist at light intensities below about 1% ambient, which occur in darker patches on the forest floor. Possible mechanisms whereby the observed spatial and temporal patterns of seedling recruitment could reduce the likelihood of the species becoming more common relative to other tree species in the forest are discussed.     

Contrasting effects of insect and molluscan herbivores on plant diversity in a long-term field experiment

The importance of invertebrate herbivores in regulating plant communities remains unclear, due to the absence of long-term exclusion experiments. An experiment in an English grassland involving long-term exclusions of insect and mollusc herbivores, along with rabbit fencing, showed strong, but opposing, effects of the invertebrate herbivores. Plant species richness declined and biomass increased following insect exclusion, due to increased dominance by a grass species, whereas mollusc exclusion led to increased herbs abundance. The two herbivores had a compensatory interaction: molluscs had no effects in the absence of insects and large insect effects depended on the absence of molluscs. The effects of invertebrate exclusion became apparent only after 8 years, and would have been seriously underestimated in shorter studies. Our results suggest that theorists and conservation managers need to shift from their historic focus on vertebrate herbivory, to a recognition that invertebrates can be equally important drivers of plant community structure.     

Ecological niche breadth and microhabitat guild structure in temperate Australian reptiles: Implications for natural resource management in endangered grassy woodland ecosystems

Ecological theory predicts that species with narrow niche requirements (habitat specialists) are more vulnerable to anthropocentric disturbances than those with broad niche requirements (habitat generalists). Hence, understanding a species ecological niche and guild membership would serve as a valuable management tool for providing a priori assessments of a species extinction risk. It also would help to forecast a species capacity to respond to land use change, as what might be expected to occur under financial incentive schemes to improve threatened ecological vegetation communities. However, basic natural history information is lacking for many terrestrial species, particularly reptiles in temperate regions of the world. To overcome this limitation, we collated 3527 reptile observations from 52 species across an endangered woodland ecoregion in south‐eastern Australia and examined ecological niche breadth and microhabitat guild structure. We found 30% of species had low ecological niche values and were classified as habitat specialists associated with large eucalypt trees, woody debris, surface rock or rocky outcrops. Cluster analysis separated species into six broad guilds based on microhabitat similarity. Approximately 80% of species belonged to guilds associated with old growth vegetation attributes or non‐renewable litho‐resources such as surface rock or rocky outcrops. Our results suggest that agri‐environment schemes that focus purely on grazing management are unlikely to provide immediate benefits to broad suites of reptiles associated with old growth vegetation and litho‐resources. Our classification scheme will be useful for identifying reptile species that are potentially vulnerable to anthropocentric disturbances and may require alternative strategies for improving habitat suitability and reptile conservation outcomes in grassy woodland ecosystems.