Can Establishment Characteristics Explain the Poor Colonization Success of Late Successional Grassland Species on Ex‐Arable Land?

Abstract Many grassland restoration projects on former arable land face problems because early successional grassland species establish vigorously and persistently but late successional grassland species fail to establish. Differences in establishment characteristics of early and late successional species might provide an explanation for the failure of many late successional species to colonize grasslands on ex‐arable land. I examined whether early and late successional species had different establishment rates in the initial years of a grassland succession, whether a specific establishment stage (seedling emergence, mortality or growth) could be identified as the key process controlling establishment, and what management would enhance the establishment of late successional grassland species. Seeds of three early and three late successional species were sown separately in ex‐arable plots with bare soil, 1‐year‐old vegetation, and 2‐year‐old vegetation. Emergence, mortality, and seedling growth were monitored for 1 year. Early successional species established successfully in the bare soil plots but failed to establish in plots with 1‐ and 2‐year‐old vegetation. Late successional species showed either lower establishment rates in the younger succession stages or decreased establishment with succession that nevertheless resulted in significant establishment in the oldest plots. Seedling emergence proved to be the key factor determining the establishment pattern of early and late successional species. In absolute numbers, emergence of late successional species was, however, similar or higher than that of early successional species, even in the earliest succession stage. The poor establishment of late successional species on former arable land could therefore not be explained solely by differences in establishment characteristics between early and late successional grassland species. Competitive processes between early and late successional species later in the life cycle probably play an important role. The results do point out that establishment of late successional species can be promoted by creating vegetative cover from the start of the restoration effort.     

Patterns of phytophagous insect herbivory and abundance on juvenile regrowth of Eucalyptus subgenera coexisting in southeastern Tasmania

Abstract Eucalypts from the subgenus Monocalyptus tend to be more abundant and dominate species from the subgenus Symphyomyrtus where they occur together in the mixed species stands of southeastern Australia. The differential impacts of herbivory by phytophagous insects has been postulated as a causal mechanism in the creation and maintenance of such stands. This research aimed to quantify phytophagous insect abundance and herbivory in mixed species juvenile regrowth of Eucalyptus globulus, E. viminalis (Symphyomyrtus), E. obliqua and E. pulchella (Monocalyptus} southwest of Hobart, Tasmania. Monocalyptus experienced a higher level of herbivory than Symphyomyrtus. However, mean damage levels were relatively low at less than 11% throughout. Furthermore, due to the positively skewed nature of herbivory data the mean was an inappropriate measure of central tendency; median damage levels ranged from 4.9% to 8.4%. Patterns of herbivory tended to be different for each eucalypt species: E. obliqua was particularly prone to chewing damage, E. pulchella and E. globulus suffered higher levels of distortion while E. viminalis was least affected by insect attack. Even though some trends in insect community structure seemed apparent at the level of eucalypt subgenus, closer examination revealed patterns of abundance were characteristic of each Eucalyptus species. The composition of foliar damage corresponded with the prominence of particular insect groups. Sucking insects tended to dominate the fauna except on E. obliqua where chewing insects in general, and chrysomelids in particular, were most prevalent. Both the distribution and magnitude of herbivory suggested that phytophagous insects had a negligible effect on competition between coexisting juvenile eucalypts and were unlikely to be responsible for the dominance of Monocalyptus or the maintenance of mixed species stands.     

Effects of leaf litter on woody seedlings in xeric successional communities

We investigated the effect of leaf litter on the establishment of Eucalyptus incrassata, a mallee eucalypt. It has been suggested that litter accumulation may hinder seedling establishment, and that the removal of litter may be one of the mechanisms through which fire enhances recruitment. We conducted factorial experiments testing the effects of three kinds of leaf litter on E. incrassata seeds and seedlings at three contiguous sites with different land use histories. One site was an uncleared E. incrassata open mallee woodland (Mallee site), one a cleared area that had been ungrazed for about five years (Pasture site) and the third an area of mallee rolled some 40 years ago and permitted to regenerate (Regrowth site). Litter had no effect on emergence of planted E. incrassata seeds, but emergence differed between sites. Overall, the percentage of seeds that germinated and emerged was substantial (mean 35.2% ± 25.9%). Seedling shoot biomass did not differ between sites or litter treatments. Although seedlings grown in Pasture litter suffered higher mortality rates, overall mortality rates were low (mean 13.2% ± 15.5%), suggesting that leaf litter has little effect on recruitment rates during winter and spring. We conclude that leaf litter does not affect emergence or growth in young E. incrassata seedlings during winter and spring, when most establishment occurs. Our results emphasize the difficulty in predicting litter effects on recruitment.     

Leaf litter of <Emphasis Type="Italic">Kageneckia angustifolia</Emphasis> D. Don (Rosaceae) inhibits seed germination in sclerophyllous montane woodlands of central Chile

Leaf litter accumulation can have either positive, negative or neutral effects on seed germination and seedling recruitment. In montane woodlands of the Mediterranean zone of central Chile, large amounts of leaf litter accumulate beneath the crowns of the summer semi-deciduous tree Kageneckia angustifolia and no regeneration of this or other plant species has been observed beneath this tree throughout the year. In a sample plot of 5000 m² we selected ten K. angustifolia trees and measured (1) leaf litter accumulation beneath and outside canopy; (2) the effects of time elapsed since burial on viability of K. angustifolia seeds with and without a leaf litter cover; (3) field seed germination with presence or absence of leaf litter and (4) the possible chemical effects of K. angustifolia leaf litter leachates on seed germination of its own seeds and of other two co-occurring native shrubs species (Guindilia trinervis and Solanum ligustrinum). Our results show that a considerable accumulation of leaf litter occurred beneath K. angustifolia, and litter negatively affected seed viability and germination of this species in the field. Under laboratory conditions, K. angustifolia leaf litter leachates inhibited seed germination of its own seeds and of the two native shrub species. Chemical effects are likely involved in the negative effects of leaf litter on the recruitment of K. angustifolia in the montane sclerophyllous woodland of central Chile.     

Using experimental manipulation to assess the roles of leaf litter in the functioning of forest ecosystems

The widespread use of forest litter as animal bedding in central Europe for many centuries gave rise to the first litter manipulation studies, and their results demonstrated that litter and its decomposition are a vital part of ecosystem function. Litter plays two major roles in forest ecosystems: firstly, litterfall is an inherent part of nutrient and carbon cycling, and secondly, litter forms a protective layer on the soil surface that also regulates microclimatic conditions. By reviewing 152 years of litter manipulation experiments, I show that the effects of manipulating litter stem from changes in one, or both, of these two functions, and interactions between the variables influenced by the accumulation of litter can result in feedback mechanisms that may intensify treatment effects or mask responses, making the interpretation of results difficult.Long-term litter removal increased soil bulk density, overland flow, erosion, and temperature fluctuations and upset the soil water balance, causing lower soil water content during dry periods. Soil pH increased or decreased in response to manipulation treatments depending on forest type and initial soil pH, but it is unclear why there was no uniform response. Long-term litter harvesting severely depleted the forests of nutrients. Decreases in the concentrations of available P, Ca, Mg, and K in the soil occurred after only three to five years. The decline in soil N occurred over longer periods of time, and the relative loss was greater in soils with high initial nitrogen concentration. Tree growth declined with long-term litter removal, probably due to lower nutrient availability. Litter manipulation also added or removed large amounts of carbon thereby affecting microbial communities and altering soil respiration rates.Litter manipulation experiments have shown that litter cover acts as a physical barrier to the shoot emergence of small-seeded species; further, the microclimate maintained by the litter layer may be favourable to herbivores and pathogens and is important in determining later seedling survival and performance. Litter manipulation altered the competitive outcomes between tree seedlings and forbs, thereby influencing species composition and diversity; changes in the species composition of understorey vegetation following treatments occurred fairly rapidly. By decreasing substrate availability and altering the microclimate, litter removal changed fungal species composition and diversity and led to a decline in populations of soil fauna. However, litter addition did not provoke a corresponding increase in the abundance or diversity of fungi or soil fauna.Large-scale long-term studies are still needed in order to investigate the interactions between the many variables affected by litter, especially in tropical and boreal forests, which have received little attention. Litter manipulation treatments present an opportunity to assess the effects of increasing primary production in forest ecosystems; specific research aims include assessing the effects of changes in litter inputs on the carbon and nutrient cycles, decomposition processes, and the turnover of organic matter.     

生物土壤结皮对种子萌发和幼苗建植的影响研究进展

生物土壤结皮（BSC）在陆地生态系统中具有重要的生态地位,尤其是旱地生态系统中,BSC占据了种子植物之间的广阔地面。因此,BSC的发展必然影响种子植物更新过程与植被空间格局;但其作用方式、影响程度等因相关研究涉及多气候要素、土壤类型、BSC组成物种和种子植物物种的差异及其不同组合,导致目前的研究结论存在广泛争议。研究综合论述了BSC改变地表微形态对种子传播过程的影响;BSC改变土壤特性（物理、化学、生物学）对种子萌发和幼苗存活与建植等关键环节的影响;并结合种子形态特征及种子萌发、幼苗建植的性状等,综合分析了BSC对种子传播、种子萌发与幼苗建植等更新过程的潜在影响机理;探讨了目前研究矛盾性结论产生的原因。总体来说,深入研究并全面揭示BSC对种子植物更新过程的影响,应加强学科交叉,将分子生物学、植物生理学、生物化学等微观研究,与遥感、野外生态因子过程监测、控制实验等宏观、中观研究结合,从机理到过程方面动态研究BSC对种子植物更新过程的影响,并引入水文模型、气候模型、种群动态模型等模型预测方法,研究气候变化、各类干扰频发的情景下,BSC对种子传播、萌发及幼苗建植过程的潜在影响,以期促进对BSC与种子植物间相互关系的研究,加深对干旱脆弱生态系统植被发展规律的认识。     

桉-桤混合凋落物分解及其土壤动物群落动态

采用放置不同孔径凋落袋(6目、30目和260目)的方法,研究了四川省乐山市苏稽镇不同比例巨桉(Eucalyptus grandis)与台湾桤木(Alnus formosana)混合凋落物的质量损失率及土壤动物群落结构的变化.结果表明:不同比例桉-桤混合凋落物均表现出前期分解迅速,后期分解较慢的规律.不同孔径凋落物袋中凋落物的分解率表现为6目最大,30目次之,260目最小.同孔径凋落物袋中不同比例桉-桤混合凋落物的分解速率也有不同,6目中各种凋落物分解时长相差较小,而30目和260目中纯巨桉、纯台湾桤木凋落物分解率达95%的时间之差分别为1175 d和908 d.凋落物分解过程中大型土壤动物类群结构发生了明显变化,分解初期主要为啮虫目,中期为后孔寡毛目,后期为鞘翅目,末期为双翅目.这些结果为进一步研究桉桤混交林物质循环提供了重要数据.     

Germination requirements and responses to leaf litter of four species of eucalypt

We studied how the responses of four species of eucalypt to leaf litter related to their germination responses to light and water availability. Two of the species (Eucalyptus obliqua and E. baxteri) have a mesic distribution, while the other two (E. oleosa, and E. incrassata) are more xerophytic. We studied the effect of litter on emergence of the four species in a glasshouse experiment. Litter did not affect the emergence of E. incrassata and E. oleosa, but enhanced the emergence of E. obliqua and E. baxteri. Litter increased the seedling mortality of all four species. Germination responses to light and water availability were studied in growth cabinets under controlled conditions. The germination of E. obliqua and E. baxteri was substantially lower under fluorescent light than in darkness, but that of E. oleosa and E. incrassata was not affected by the light environment. The germination of E. obliqua and E. baxteri was significantly reduced by reduced water potential (a). Substantial germination of E. oleosa and E. incrassata occurred even at very low a (less than –1.05 MPa). We conclude that both the shade and the humid micro-environment provided by litter may have contributed to the emergence responses of the four species to litter, and these responses may correspond to ecological adaptations to the different environments in which they live.     

巨桉凋落叶分解初期对菊苣生长和光合特性的影响

采用盆栽试验,于2010年3-5月在四川农业大学教学科研园区内研究了巨桉凋落叶分解初期对受体植物菊苣幼苗生长和光合特性的影响.试验设置CK (0 g·pot-1)、A1(30 g·pot-1)、A2(60 g·pot-1)和A3 (90 g·pot-1)4个凋落叶施用水平,将各处理的凋落叶分别与12 kg土壤混合后装盆,播种菊苣,分别在播种30、45、60和75 d测定菊苣生长指标,待凋落叶量的最高处理组A3植株第3片真叶完全展开后,测定菊苣叶片的光合生理指标.结果表明:在各测定时间下,不同巨桉凋落叶施用水平的菊苣生物量积累和叶面积增长受到显著抑制;在凋落叶分解初期,菊苣幼苗叶片光合色素合成受到明显抑制,且随着凋落叶施用量增加抑制作用加大,各处理幼苗的光合速率日变化均呈午休双峰型曲线,气孔导度和水分利用效率的变化趋势与净光合速率相同,日光合总量表现为CK＞A1 ＞A2 ＞A3.经GC-MS定期检测,凋落叶中有33种小分子化合物随着凋落叶的分解而逐步释放,以具有化感作用的萜类物质为主.     

Similar breakdown rates and benthic macroinvertebrate assemblages on native and Eucalyptus globulus leaf litter in Californian streams

1. Eucalyptus globulus, a tree species planted worldwide in many riparian zones, has been reported to affect benthic macroinvertebrates negatively. Although there is no consensus about the effects of Eucalyptus on aquatic macrobenthos, its removal is sometimes proposed as a means of ecological restoration. 2. We combined the sampling of macroinvertebrates with measurement of the colonisation of leaf packs in mesh bags, to examine the effects of riparian Eucalyptus and its litter on benthic macroinvertebrates in three small streams in California, U.S.A. Each stream included one reach bordered by Eucalyptus (E‐site) and a second bordered by native vegetation (N‐site). 3. The macrobenthos was sampled and two sets of litter bags were deployed at each site: one set with Eucalyptus litter (Euc‐bags) and one with mixed native tree litter (Nat‐bags) containing Quercus, Umbellularia, Acer and Alnus. Bags were exposed for 28, 56 and 90 days and this experiment was repeated in the autumn, winter and spring to account for effects of changing stream flow and insect phenology. 4. Litter input (average dry mass: 950 g m^?2 year^?1 in E‐sites versus 669 g m^?2 year^?1 in N‐sites) was similar, although in‐stream litter composition differed between E‐ and N‐sites. Litter broke down at similar rates in Euc‐bags and Nat‐bags (0.0193 day^?1 versus 0.0134 day^?1), perhaps reflecting the refractory nature of some of the leaves of the native trees (Quercus agrifolia). 5. Summary metrics for macroinvertebrates (taxon richness, Shannon diversity, pollution tolerance index) did not differ significantly between the E and N sites, or between Euc‐bags and Nat‐bags. No effect of exposure time or site was detected by ordination of the taxa sampled. However, distinct seasonal ordination clusters were observed in winter, spring and autumn, and one of the three streams formed a separate cluster. 6. The presence of Eucalyptus was less important in explaining the taxonomic composition of the macrobenthos than either ‘season’ or ‘stream’. Similarly, these same two factors (but not litter species) also helped explain the variation in leaf breakdown. We conclude that patches of riparian Eucalyptus and its litter have little effect on stream macrobenthos in this region.     

内蒙古典型草原群落实生苗动态对刈割和放牧干扰的响应

该文通过刈割和放牧样地的实生苗动态观测,试图揭示刈割和放牧干扰对冰草(Agropyron cristatum)、克氏针茅(Stipa krylovii)和糙隐子 草(Cleistogenes squarrosa )等 3种禾草实生苗出生、存活和死亡季节动态的影响,以及这种效应是如何受气温和降水影响的。利用有重复测 量的方差分析,比较了不同刈割留茬高度(0、5和15 cm)与放牧处理之间在实生苗密度、死亡率和出现率的时间动态方面的差异。结果表明：3 种禾草的实生苗密度自5月下旬至8月中旬一直呈增加态势。刈割在一定程度上抑制了克氏针茅的实生苗出现率,原因可能是相对高大的克氏针 茅在前一年刈割时会被去掉相当多的生殖部分,其种子数量必然受到影响,导致实生苗出现率的降低。同时,刈割对相对低矮的糙隐子草实生 苗有正向效应 ,即促进其实生苗的出现,可能的原因是刈割使植被高度降低、植被稀疏,植被冠层以下至地面的光照较好、地温较高,同时位 于下层的糙隐子草受到的高大植株的竞争压力减小。生长季早期和晚期实生苗死亡率较高,这与这两个时期的干旱和低温有关,而自6月中旬至 7月初之间的死亡率小高峰也是先前的干旱少雨导致的。3种禾草实生苗的大规模出现几乎都在5～7月,这与该时期相对适宜的气温、地温和水 分条件有关。对实生苗更新相关过程 (如种子萌发、休眠、种子库存量等)进行更加深入细致的研究,将进一步揭示种子生产和有性繁殖在群落 过程中的作用。     

内蒙古典型草原群落实生苗动态对刈割和放牧干扰的响应

 该文通过刈割和放牧样地的实生苗动态观测,试图揭示刈割和放牧干扰对冰草(Agropyron cristatum)、克氏针茅(Stipa krylovii)和糙隐子 草(Cleistogenes squarrosa )等 3种禾草实生苗出生、存活和死亡季节动态的影响,以及这种效应是如何受气温和降水影响的。利用有重复测 量的方差分析,比较了不同刈割留茬高度(0、5和15 cm)与放牧处理之间在实生苗密度、死亡率和出现率的时间动态方面的差异。结果表明：3 种禾草的实生苗密度自5月下旬至8月中旬一直呈增加态势。刈割在一定程度上抑制了克氏针茅的实生苗出现率,原因可能是相对高大的克氏针 茅在前一年刈割时会被去掉相当多的生殖部分,其种子数量必然受到影响,导致实生苗出现率的降低。同时,刈割对相对低矮的糙隐子草实生 苗有正向效应 ,即促进其实生苗的出现,可能的原因是刈割使植被高度降低、植被稀疏,植被冠层以下至地面的光照较好、地温较高,同时位 于下层的糙隐子草受到的高大植株的竞争压力减小。生长季早期和晚期实生苗死亡率较高,这与这两个时期的干旱和低温有关,而自6月中旬至 7月初之间的死亡率小高峰也是先前的干旱少雨导致的。3种禾草实生苗的大规模出现几乎都在5～7月,这与该时期相对适宜的气温、地温和水 分条件有关。对实生苗更新相关过程 (如种子萌发、休眠、种子库存量等)进行更加深入细致的研究,将进一步揭示种子生产和有性繁殖在群落 过程中的作用。