Seed Dispersers,Seed Predators,and Browsers Act Synergistically as Biotic Filters in a Mosaic Landscape

In this study, we analize the functional influence of animals on the plants they interact with in a mediterranean mountain. We hypothesise that seed dispersers, seed predators, and browsers can act as biotic filters for plant communities. We analyse the combined effects of mutualistic (seed dispersal) and antagonistic (seed predation, herbivory) animal interactions in a mosaic landscape of Mediterranean mountains, basing our results on observational and experimental field. Most of the dispersed seeds came from tree species, whereas the population of saplings was composed predominantly of zoochorous shrub species. Seed predators preferentially consumed seeds from tree species, whereas seeds from the dominant fleshy-fruited shrubs had a higher probability of escaping these predators. The same pattern was repeated among the different landscape units by browsers, since they browsed selectively and far more intensely on tree-species saplings than on the surrounding shrubs. In synthesis, our work identifies the major biotic processes that appear to be favoring a community dominated by shrubs versus trees because seed dispersers, predators, and herbivores together favored shrub dispersal and establishment versus trees.     

Differential effects of goat browsing on herbaceous plant community in a two-phase mosaic

The impact of herbivores on herbaceous plant communities is usually attributed to direct consumption of plants. We hypothesized that goats affect herbaceous plants both directly (consumption by foraging) and indirectly, by changing environmental conditions through modification of woody plant structure. We assessed the effects of goats browsing on environmental conditions, landscape structure, and herbaceous plants to link the direct and indirect effects of goats on herbaceous communities. Our model system was the Mediterranean woodland in Mt. Carmel, Israel. This is a two-phase mosaic landscape, composed of herbaceous (open) and woody patches. We delineated 10 plots of 1000 m², goats were introduced to five plots and five plots remained without goats. We monitored plant species richness and composition in two adjacent patch types (woody and open) in each plot. For each patch type, in all plots, we collected data on environmental conditions. We analyzed landscape structure using landscape metrics derived from a high-resolution vegetation map. We found that goats modified the structure of woody plants and hence the landscape mosaic. This alteration was associated with changes in environmental conditions, with more light penetration and higher temperatures. The impact of goats on the herbaceous plant community depended on patch type. In open patches, goats affected the herbaceous community mostly by direct consumption, whereas in woody patches they affected the herbaceous community mainly by modification of abiotic conditions. Our results stress the importance of considering landscape and patch structure in analyzing the effect of herbivory on plant communities.     

Composition and diversity of herbaceous patches in woody vegetation: The effects of grazing,soil seed bank,patch spatial properties and scale

Herbaceous plants contribute much to plant diversity in Mediterranean-type ecosystems though mostly occupying relatively small patches within the dense woody vegetation. While studying species diversity in the herbaceous patches, we hypothesized that grazing, soil seed bank, and spatial properties of the patch affect plant diversity and composition at different spatial scales. The study site was in an LTER site located in the Mediterranean region in north Israel. We determined herbaceous species composition in: (1) randomly sampled quadrats in herbaceous patches in grazed and un-grazed plots; (2) soil seed bank samples taken from the same patches and germinated under optimal greenhouse conditions; (3) quadrats in the same patches sown with a homogenous mixture of local soil samples. Using GIS methods, we determined small-scale spatial characteristics of the herbaceous patches. Alpha and beta diversities were calculated at the patch and plot scales using Shannon's entropy H. Grazing increased alpha diversity of local untreated seed bank samples but decreased alpha diversity of the artificial homogenous soil seed bank mixture at both patch and plot scales. Positive relation between alpha diversity and patch area was detected only under grazing. Grazing increased beta diversity in all three treatments at the patch scale. Grazing decreased the similarity in species composition between above-ground vegetation and soil seed bank. The results indicate that moderate cattle-grazing affects species diversity in the herbaceous patches within the dense maquis. This effect is scale-dependent, and interacts with the effects of soil seed bank and patch spatial-properties: without grazing soil seed bank plays a more important role than patch spatial properties, but under grazing the size and the accessibility of the patch are more important in the determination of herbaceous species composition.     

The effects of shrub patch sizes on the colonization of pioneer plants on the volcano Mount Koma,northern Japan

Vegetation recovery on Mount Koma, Hokkaido, Japan, has been slow after the catastrophic eruption in 1929, due to undeveloped soil and limited plant colonization. Nowadays, the seedling establishment is supported mostly by a nurse plant, Salix reinii forming shrub patches, facilitates the plant colonization. Although the effects of shrub patches should differ with patch sizes, the size effects have not been examined well. To examine the size effects, seed-sowing experiments were conducted on two common pioneer herbaceous species, Miscanthus sinensis and Polygonum sachalinense, in the field. The seed germination and seedling survival were monitored by the seeds sown into S. reinii patches (0.97 m²–4.12 m² in area) for 4 months during snow-free periods. Microenvironments altered by the patches were measured. Lab-experiments were performed to characterize the seed germination and seedling growth.Larger patches decreased light intensity and temperature more and increased litter and water content. The large patches promoted the seed germination of the two species. Interspecific interactions, examined by a seed mixture experiment, showed that the interaction increased the seed germination on M. sinensis and decreased that on P. sachalinense. On the lab-experiments at three temperatures (15, 20 and 25 °C), M. sinensis seeds germinated more at higher temperatures and obtained higher seedlings biomass. P. sachalinense germinated the seeds more at 20 °C and grew faster at lower temperatures. The total biomass of the two species was reduced by shade that intercepted 50% of light intensity. The seed germination and seedling growth of these two species became higher on litter with 2 cm in depth than on no litter. Soil water supported seed germination when the seeds of these two species were mixed while the water reduced the growth of P. sachalinense seedlings. Therefore, the dry soils were suitable for their growths. In all the treatments, P. sachalinense seedlings showed higher mortalities than M. sinensis.In conclusion, the large patches facilitated more to the colonization of pioneer plants via seed germination and growth. Large patches acted as a nursery supporting the natural regeneration in the disturbed area by improving litter accumulation, maintaining soil water, reducing strong light and/or protecting from heat.     

Woody structure facilitates invasion of woody plants by providing perches for birds

Woody encroachment threatens prairie ecosystems globally, and thus understanding the mechanisms that facilitate woody encroachment is of critical importance. Coastal tallgrass prairies along the Gulf Coast of the US are currently threatened by the spread of several species of woody plants. We studied a coastal tallgrass prairie in Texas, USA, to determine if existing woody structure increased the supply of seeds from woody plants via dispersal by birds. Specifically, we determined if (i) more seedlings of an invasive tree (Tridacia sebifera) are present surrounding a native woody plant (Myrica cerifera); (ii) wooden perches increase the quantity of seeds dispersed to a grassland; and (iii) perches alter the composition of the seed rain seasonally in prairie habitats with differing amounts of native and invasive woody vegetation, both underneath and away from artificial wooden perches. More T. sebifera seedlings were found within M. cerifera patches than in graminoid‐dominated areas. Although perches did not affect the total number of seeds, perches changed the composition of seed rain to be less dominated by grasses and forbs. Specifically, 20–30 times as many seeds of two invasive species of woody plants were found underneath perches independent of background vegetation, especially during months when seed rain was highest. These results suggest that existing woody structure in a grassland can promote further woody encroachment by enhancing seed dispersal by birds. This finding argues for management to reduce woody plant abundance before exotic plants set seeds and argues against the use of artificial perches as a restoration technique in grasslands threatened by woody species.     

Shrub encroachment in Argentinean savannas

Abstract. In the Wet Chaco region of Argentina, increasing shrub encroachment in savannas over the last few decades has led to important changes in the structure and functioning of the landscape. Some sectors of this territory are characterized by the appearance of circular clusters of woody patches, dispersed throughout the grassland matrix. The increasing size of these patches leads to a gradual change from grassland to dense shrubland. We studied these circular woody patches in the eastern region of the Argentine province of Formosa and characterized the variation in terms of floristic composition, diversity and predominant seed dispersal mode in different size patches. We observed an increase in species richness, diversity and compositional heterogeneity among patches with increasing patch size. Seed dispersal by animals, especially birds, is an important factor in the expansion of these woody vegetation patches within the grassland matrix.     

Seed Dispersal Patterns by Large Frugivorous Mammals in a Degraded Mosaic Landscape

Seed dispersal by Red fox (Vulpes vulpes), Stone marten (Martes foina), and Wild boar (Sus scrofa) was analyzed in an extensively degraded mosaic landscape in Sierra Nevada (SE Spain). The main objective was to determine whether seed dispersal by mammals was related to habitat degradation within a mosaic of adjacent degraded patches mixed with native forest and thereby to determine the potential role of mammals as seed dispersers in degraded landscape units. For three consecutive years, mammal feces were collected in the fruit production period, extracting all seeds of woody species found therein and analyzing their viability. Feces were collected in three different plots for each of five different landscape units: shrubland, native forest, and dense, cleared, and fenced reforestation stands. Seeds from 16 woody species (which represent more than a half of the total fleshy‐fruited woody species available) were recorded, although some agrarian species are also introduced in a low percentage of the scats. Seeds showed a high viability rate for all dispersed species, irrespective of the mammal disperser. No differences in species composition appeared in the overall landscape units or in the seed density between degraded habitats. Due to the small patch size, the high viability of dispersed seeds, and the large home range of the large mammals, these three animal species act as efficient seed dispersers for a diverse assemblage of woody plant species regardless of the habitat type within this degradation framework. This fact has important consequences for the biodiversity recuperation in these degraded habitats, principally in pine plantations.     

Relationships between spatial configuration of tropical forest patches and woody plant diversity in northeastern Puerto Rico

The destruction and fragmentation of tropical forests are major sources of global biodiversity loss. A better understanding of anthropogenically altered landscapes and their relationships with species diversity and composition is needed in order to protect biodiversity in these environments. The spatial patterns of a landscape may control the ecological processes that shape species diversity and composition. However, there is little information about how plant diversity varies with the spatial configuration of forest patches especially in fragmented tropical habitats. The northeastern part of Puerto Rico provides the opportunity to study the relationships between species richness and composition of woody plants (shrubs and trees) and spatial variables [i.e., patch area and shape, patch isolation, connectivity, and distance to the Luquillo Experimental Forest (LEF)] in tropical forest patches that have regenerated from pasturelands. The spatial data were obtained from aerial color photographs from year 2000. Each photo interpretation was digitized into a GIS package, and 12 forest patches (24–34 years old) were selected within a study area of 28 km². The woody plant species composition of the patches was determined by a systematic floristic survey. The species diversity (Shannon index) and species richness of woody plants correlated positively with the area and the shape of the forest patch. Larger patches, and patches with more habitat edge or convolution, provided conditions for a higher diversity of woody plants. Moreover, the distance of the forest patches to the LEF, which is a source of propagules, correlated negatively with species richness. Plant species composition was also related to patch size and shape and distance to the LEF. These results indicate that there is a link between landscape structure and species diversity and composition and that patches that have similar area, shape, and distance to the LEF provide similar conditions for the existence of a particular plant community. In addition, forest patches that were closer together had more similarity in woody plant species composition than patches that were farther apart, suggesting that seed dispersal for some species is limited at the scale of 10 km.     

Landscape heterogeneity as an ecological filter of species traits

Landscape heterogeneity is a major driver of biodiversity in agricultural areas and represents an important parameter in conservation strategies. However, most landscape ecology studies measure gamma diversity of a single habitat type, despite the assessment of multiple habitats at a landscape scale being more appropriate. This study aimed to determine the effects of landscape composition and spatial configuration on life-history trait distribution in carabid beetle and herbaceous plant communities. Here, we assessed the gamma diversity of carabid beetles and plants by sampling three dominant habitats (woody habitats, grasslands and crops) across 20 landscapes in western France. RLQ and Fourth Corner three-table analyses were used to assess the association of dispersal, phenology, reproduction and trophic level traits with landscape characteristics. Landscape composition and configuration were both significant in explaining functional composition. Carabid beetles and plants showed similar response regarding phenology, i.e. open landscapes were associated with earlier breeding species. Carabid beetle dispersal traits exhibited the strongest relationship with landscape structure; for instance, large and apterous species preferentially inhabited woody landscapes, whereas small and macropterous species preferentially inhabited open landscapes. Heavy seeded plant species dominated in intensified agricultural landscapes (high % crops), possibly due to the removal of weeds (which are usually lightweight seeded species). The results of this study emphasise the roles of landscape composition and configuration as ecological filters and the importance of preserving a range of landscape types to maintain functional biodiversity at regional scales.     

Recruitment limitation of forest communities in a degraded Mediterranean landscape

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

鸟类传播种子对几种树篱中侵入植物多样性的影响

2001年4月至2003年11月,在南京中山植物园内调查了冬青卫矛(Euonymus japonicus)、日本珊瑚树(Viburnum awabuki)和溲疏(Deutzia scabra)3种树篱,以及位于不同生境的2个黄杨(Buxus microphyllavar.sinica)树篱中侵入生长的杂草及杂树种类,并选择3 km外南京农业大学实验楼周围的冬青卫茅树篱作为参照。在6个树篱类型中调查到侵入生长的种子植物共计1230株,分隶于42科57属70种植物。其中适应于鸟类传播的植物有55种1047株植物;适应于风力传播的植物有10种161株植物;传播媒介不明的植物有5种22株植物。各树篱类型中,适应于鸟类传播种子的侵入植物种类数目显著多于风力传播的植物种类(t=5.086,df=10,p<0.0001)和种子传播媒介不明的植物种类(t=8.446,df=10,p<0.0001),但各树篱中适应于风力传播的侵入植物与种子传播媒介不明的植物在种类数目上无显著差异(t=1.977,df=10,p=0.076)。南京中山植物园的5个树篱类型中,适应于鸟类传播的侵入植物物种多样性的Shannon-Wiener指数都较高,为2.151至2.917,平均2.671±0.306,并且种类数目也较多,为15~36种,平均25.6±8.1种。而南京农业大学内冬青卫茅树篱的物种多样性指数最低,为1.679,并且种类数目也最少,为12种。Bray-Curtis相似性指数的聚类分析结果显示,不同树篱类型中适应于鸟类传播种子的侵入植物组成的相似性主要受到树篱周围的种子源及其距离、食果鸟类、人为干扰活动等生境因子的影响,而非简单取决于树篱种类本身。鸟类传播种子增加了单一树种构成的树篱中的种子植物多样性,同时也对树篱的景观产生了一定的不利影响,不过人工对树篱的日常修剪等管护措施严格限制了这些侵入植物实生苗及小树的正常生长。研究结果反映出种子源、鸟类传播种子和灌丛在促进植物群落的发展和演替过程中具有重要的生态意义。     

贵阳市区灌木林生态系统生物量及碳储量

采用直接收获法和实测数据,以贵州省贵阳市区天然灌木林内木本和草本植物、凋落物及土壤为研究对象,研究了灌木林生态系统的生物量、碳含量及碳储量。结果表明:灌木林植被层生物量为23.16 t/hm2,其中木本植物层生物量为12.46 t/hm2;草本植物层为3.74 t/hm2;凋落物层为6.96 t/hm2,分别占植被层生物量的53.08%、16.15%、30.05%。木本植物25种的碳含量范围为445.91—603.46 g/kg;草本植物6种的碳含量为408.48—523.04 g/kg;凋落物层碳含量为341.01—392.81 g/kg;土壤层碳含量为5.73—26.68 g/kg。生态系统总碳储量为88.34 t/hm2,其中植被层为8.10 t/hm2;凋落物层为2.56 t/hm2;土壤层为77.68 t/hm2,分别占系统总碳储量的9.17%、2.89%、87.94%。灌木林生态系统碳储量的空间分布格局为:土壤层植被层凋落物层。研究结果,可为喀斯特城市估算森林生态系统碳储量和碳平衡提供科学依据。     

Woody species as landscape modulators: their effect on the herbaceous plants in a Mediterranean maquis

“Landscape modulators” are ecosystem engineers that have an impact on community structure by creating patches in the landscape mosaic. Our aim was to study the effect of evergreen-trees, as landscape modulators, on herbaceous plants in a Mediterranean maquis system in northern Israel. We examined the effects of canopy removal and cattle grazing on species richness, plant functional types, and rare plant species in two patch-types: (1) woody—under tree canopy (or the location of a removed canopy); (2) herbaceous—in open areas with no tree canopy. Patch-type and tree removal affected species richness and plant functional types. The extreme negative effect of the woody patch-type on species richness disappeared soon after the removal of the landscape modulator canopy. We conclude that the dominant effect of the evergreen woody landscape modulators can be regulated by canopy removal and grazing for maintaining patch-type and landscape diversities, and consequent high species richness in Mediterranean ecosystems, which is a main goal of global nature conservation policy.     

生境片段化对千岛湖马尾松林内土壤种子库的影响

土壤种子库是森林群落更新的主要来源之一,对森林的演替和恢复等具有重要意义。生境片段化现象正日益严重地影响着森林群落,并可影响森林土壤种子库。研究了千岛湖地区的大陆及岛屿次生马尾松林内土壤种子库的组成及其影响因素(e.g.,岛屿面积,形状指数,隔离度和距岛屿边缘距离等)。根据大陆和岛屿的面积及边缘梯度,采用大数量小样方法,分别在土壤种子库最大化(初冬,2015年12月)和最小化(晚春,2016年4月)时期对马尾松林内土壤进行了机械取样。对土壤样品进行萌发实验,检测了两个时期的土壤种子库上层(0—2 cm)和下层(2—5 cm)种子组成,并通过广义线性混合效应模型等手段分析其影响因素。结果显示:(1)所有316个土壤样本中,萌发出幼苗1422株,隶属于29科、40属、41种。其中,木本植物幼苗占13种1024株,草本占28种398株。(2)Jaccard指数和相关性分析均显示初冬、晚春时期的土壤种子库组成具有很高的相似性;土壤种子库上、下层组成的相似性也很高。(3)广义线性混合效应模型分析显示,在大陆和岛屿上,土壤种子库下层种子含量低于上层;而大陆样地土壤种子库中的木本植物种子数较岛屿样地高。岛屿上,土壤种子库中的种子数随土层的加深而降低;随边缘梯度升高也下降,尤其是草本植物的种子。对于岛屿上的木本植物,不耐阴种的种子数量远大于耐阴种,尤其是土壤下层。表明千岛湖地区马尾松林内土壤种子库组成受到生境片段化的影响,进而可能作用于该类型森林群落的演替。     

The effect of plant litter on ecosystem properties in a Mediterranean semi‐arid shrubland

Abstract. We studied the distribution of litter in a shrubland of the Negev with a semi‐arid Mediterranean climate of less than 200 mm of rainfall per year. Our focus was on the effects of litter on properties of landscape patches relevant to ecosystem processes (water runoff and soil erosion), annual plant community responses (seedling density, biomass production and species richness), and animal activity (soil disturbance by termites). Three 60‐m transects, extending across a pair of opposing north‐ and south‐facing slopes and their drainage channel, showed that litter accumulates not only under shrubs, but to a lesser extent also on the crusted inter‐shrub open areas. We used 35 experimental units (‘cells’, 0.5m × 1 m), each containing a crust and a shrub patch. Because runoff flows from crusted patches and is intercepted by shrub patches, the latter were in the lower third of the cells. Leaf litter was added in single and double amounts providing ca. 0.5 and 1.0 cm litter depth, to either, both, or none of the patches. Litter addition significantly decreased the amount of runoff, regardless of the location and amount of litter applied. Litter on the crust increased species number and seedling density of species with low abundance. Adding a double litter layer increased annual plant biomass production, while a single amount had no effect. Litter addition to the shrub patch affected neither biomass nor species richness. Litter addition to both patches at both quantities caused a large increase in termite activity. Termites caused disturbance by disrupting the crust, which may contribute to the reduction in runoff amounts. In the open, flat crust patches, annual plant communities are limited in their productivity and species richness, as there are few structures stopping the outflow of water, soil and seeds. Litter adds such structures, but affects the plant communities only when added to litter‐free crust. Litter accumulation and its patchy distribution have large impacts on landscape patch properties affecting resource distribution, plant productivity and diversity, and animal activity. Therefore, understanding litter distribution in relation to the patchy structure of the landscape of semi‐arid shrubland should be viewed as an important component of shrubland management.     

Frugivoría por Aves en un Paisaje Fragmentado: Consecuencias en la Dispersión de Semillas1

Few data exist on seed dispersal by frugivorous birds in fragmented landscapes, originating from tropical dry forests, in contrast to more abundant data from tropical rain forests. In this study, we assessed the effect of frugivorous birds in a fragmented landscape of Veracruz, Mexico, now occupied by remnant fragments of tropical semi‐deciduous forest and dry deciduous forest, grassland, and shrubby patches on sand dunes. We determined four characteristics related to seed dispersal by birds: the interacting species of plants and birds, the characteristics of these species, spatio‐temporal variation in the dispersal system, and the outcome of the process. During one year, we recorded 54 frugivorous bird species and 33 ornithochorous plant species, which engaged in 176 different bird‐plant species interactions. Similarity (Sorensen index) of frugivorous bird communities using different vegetation types was high (>70%), suggesting that many bird species used all of the vegetation types. In contrast, the similarity of ornithochorous plant communities among vegetation types commonly was low (<37%), suggesting that most plant species were restricted to particular sites in this landscape. At the landscape level, as well as for tropical deciduous forest, we detected a significant positive relationship (Spearman's correlation of rank coefficient >0.65, P <0.05) among richness per month of frugivorous birds and plant species bearing fleshy fruits. Seeds of many plant species previously detected in studies of seed rain at the site were eaten by birds during this study. Most seeds of zoochorous species, which are deposited in the dry and decidous tropical forests patches, are produced within these vegetation types (i.e., they are autochthonous species), whereas bird‐dispersed seeds arriving in grassland and shrubby patches are produced outside (i.e., allochthonous) and are mostly woody species. Birds are important seed dispersers among vegetation types in this landscape but they have different effects in each one. The four characteristics studied, as well as the landscape approach of this research, allowed us to detect spatial and temporal patterns that otherwise would have remained undetected.     

Does seed mass drive the differences in relative growth rate between growth forms?

The idea that herbaceous plants have higher relative growth rates (RGRs) compared with woody plants is fundamental to many of the most influential theories in plant ecology. This difference in growth rate is thought to reflect systematic variation in physiology, allocation and leaf construction. Previous studies documenting this effect have, however, ignored differences in seed mass. As woody species often have larger seeds and RGR is negatively correlated with seed mass, it is entirely possible the lower RGRs observed in woody species is a consequence of having larger seeds rather than different growth strategies. Using a synthesis of the published literature, we explored the relationship between RGR and growth form, accounting for the effects of seed mass and study-specific effects (e.g. duration of study and pot volume), using a mixed-effects model. The model showed that herbaceous species do indeed have higher RGRs than woody species, and that the difference was independent of seed mass, thus at all seed masses, herbaceous species on average grow faster than woody ones.     

Non-native and native shrubs have differing impacts on species diversity and composition of associated plant communities

The spread of non-native plants has been depicted as a serious threat to biodiversity. However, it remains unclear whether the indigenousness of the invading plant plays a marked role for the ecological consequences of an invasion as few studies have compared the ecological impacts of non-native shrubs with structurally or functionally comparable native shrubs. We studied patches of introduced and native shrubs to assess whether there are general differences in plant species composition or biomass between patches formed by non-native versus native shrubs. The indigenousness of the shrub (non-native vs. native) did not explain the variation in soil nutrients, neither the production of shoot biomass or allocation of growth to different parts of the shoot. The amount of light reaching ground level did not differ between patches of a non-native and a native shrub. However, species richness and biomass of herbaceous plants were lower in patches of non-native than native shrubs and the amount of litter was higher below non-native than native shrubs. Our results suggest that the indigenousness of the patch-forming plant may be an important factor for the diversity and composition of associated herbaceous vegetation. Based on our results, resource availability (light and nutrients) is not a sufficient explanation for the negative effects of non-native shrubs on plant communities. Further research is needed to investigate whether alternative explanations, such as the novelty of the toxic compounds produced by non-native plants, can explain the differences we observed.     

Herbaceous plant richness and vegetation cover in Mediterranean grasslands and shrublands

Different types of relationship between herbaceous species richness and several parameters indicating abundance of plant material (herbaceous, woody plants, litter and bare ground cover) are presented. The data were obtained from 50 sites along a 300 km strip running from E to W within Spain and Portugal. Each site was representative of the silvo-pastoral landscape of the Mediterranean type ecosystems of the Iberian peninsula, and contained two neighboring patches, one of grassland and the other of shrubland. 3,600 20 × 20 cm subplots were randomly located (72 per site, 36 per patch) crossing the boundary grassland/shrubland. This approach allowed us to analyze the richness-occupation relationship of the space from different points of view: among and within the sites, and among and within the grassland and shrubland plant communities. We found a unimodal relationship between richness-cover similar to the one generally accepted between richness and biomass. Our results show that the dependence of this relationship varies depending on the spatial scale of the analysis and on the type of data used. When the whole region is taken into account, significant unimodal relationships are found between richness and herbaceous cover, litter and bare ground, and a negative linear relationship with woody plant cover. Within the sites there are mainly linear or non-significant relationships. But the results also depend on the type of communities analyzed. In pastures, the unimodal relationship represents the combination of positive and negative linear responses for low and high cover values, respectively. The value for herbaceous cover in which maximum richness occurs is around 60%. In shrublands, this value for cover also corresponds to maximum species richness, although the possibilities of reaching it are limited by other variables, such as woody plant cover. This implies that, on not considering variability at local scale, the relationship is linear and positive. This paper shows the existence of a common model related to herbaceous cover, but this model has multiple controlling factors that act differently in each type of community.     

The use of branch piles to assist in the restoration of degraded semiarid steppes

Desertification is a major environmental problem in arid and semiarid regions. Tree plantation has been commonly employed to foster the recovery of degraded areas. However, this technique is costly, and their outcomes are often uncertain. Therefore, we evaluated an alternative method for the restoration of degraded semiarid steppes that involved the construction of branch piles to attract frugivores as potential seed‐dispersing birds, promoting seed rain, and fostering the formation of woody patches. We measured the success of branch piles in terms of the number of bird visits and seed input compared to naturally occurring shrub patches. Generally, frugivorous birds visited branch piles less frequently than shrub patches. Yet, branch piles accumulated seeds of patch‐forming shrub species. Seed rain was higher under patches of the dominant shrub Rhamnus lycioides than under branch piles. In contrast, woody patches and branch piles did not differ in seed input of the less abundant Pistacia lentiscus shrub. Our study demonstrates that branch piles are used by frugivorous birds and accumulate seeds of patch‐forming shrubs. Branch piles may be a suitable method to promote the expansion of bird‐dispersed plant species and restore semiarid wooded steppes. However, their efficiency largely depends on pile persistence and economic cost.