Seed Dispersal in Páramo Plants: Epizoochorous and Hydrochorous Taxa

Abstract: On the basis of a recent checklist of plant diversity in páramos, diaspores collected from herbaria were studied for adaptations to dispersal on animals and by water. This study shows that the páramo flora has a relatively high percentage of genera with morphological adaptations to epizoochorous and to hydrochorous diaspore dispersal. Genera with hooked and straight appendages are present throughout the páramo belt, while their number decreases in the higher páramo zones. About half of the hydrochorous genera and one-third of the epizoochorous ones can be found throughout all páramo zones. The contribution of holarctic epizoochorous genera to the páramo flora seems to be greater than that of austral-antarctic genera, whereas in hydrochorous genera it is the reverse.     

The growth form composition of plant communities in the ecuadorian páramos

A growth form classification for the plants of the Ecuadorian páramos is proposed, consisting of ten forms: stem rosettes, basal rosettes, tussocks, acaulescent rosettes, cushions and mats, upright shrubs, prostrate shrubs, erect herbs, prostrate herbs, and trailing herbs.The growth form composition of 192 samples of páramo vegetation from twelve different regions in Ecuador was analysed using multivariate techniques to determine physiognomic types. The distribution of growth forms was significantly related to altitude, rock cover, bare ground and to measures of disturbance and exposure.The dominant growth forms in all the samples belonged to one of Hedberg's (1964) five types, but other forms were also significant components of the vegetation. The majority of Ecuadorian páramo vegetation showed a relatively consistent growth form composition, dominated by tussocks. The accompanying growth forms were mostly acaulescent rosettes, cushions, upright shrubs, prostrate shrubs, erect herbs and prostrate herbs, sometimes with stem rosettes, basal rosettes or trailing herbs.At higher altitudes, the dominance of tussocks was reduced. At first, acaulescent rosettes became dominant, but higher still their dominance was shared with cushions. At the highest altitudes of all, where plant cover was thin, no single growth form was dominant. In other locations where plant cover was sparse, once again no single growth form was dominant. In humid páramos, stem rosettes were co-dominant with tussocks or erect herbs. Basal rosettes, erect herbs and prostrate herbs were locally co-dominant at higher altitudes.The growth form composition of the Ecuadorian páramos showed similarities with other tropical alpine regions, though no comparable quantitative data for these regions are available yet. The quantitative determination of growth form composition may also lead to a better understanding of community structure and the mechanisms which govern it.     

Sensitivity of Soil Restoration Indicators during Páramo Succession in the High Tropical Andes: Chronosequence and Permanent Plot Approaches

The aim of this study was to analyze indicators of soil restoration during secondary succession in a heterogeneous valley in the high tropical Andes. A combination of chronosequence and permanent plot methods was used to detect changes in this heterogeneous matrix. Thirty-six plots with different fallow times (1–9 years) and four noncultivated plots with natural vegetation (páramo) were sampled twice in a 3-year interval (1996 and 1999). The following soil properties were determined: total C and N, pH, exchangeable bases, cation exchange capacity, and microbial biomass N (MB-N). Using the chronosequence approach, successional increases in soil pH and Mg were detected, pointing to these variables as indicators of soil restoration during the fallow period. Comparing the noncultivated páramo with the fallow plots, a significant decrease in MB-N was found, suggesting that this is a sensitive agricultural disturbance indicator. The permanent plot analysis failed to detect successional trends in any of the study variables, probably as a result of a lack of sensitivity of the indicators used within the 3-year interval. Nevertheless, a strong acidification was detected by the permanent plot method when fallow plots were cultivated. We conclude that the size of important soil components such as total soil organic matter or microbial biomass is not a sensitive soil restoration indicator in these heterogeneous mountain systems but that other integrative variables such as pH could be more sensitive to successional changes in key soil processes (e.g., nitrification or nutrient losses).     

Analysis of Andes frogs (Phrynopus, Leptodactylidae, Anura) phylogeny based on 12S and 16S mitochondrial rDNA sequences

South American leptodactylid frogs of the genus Phrynopus occur in cloud-forest, páramo, subpáramo and puna habitats (1000–4400 m elevation) from Colombia to Bolivia. Currently, there are 34 described species; however, many additional species new to science have been reported from Colombia, Peru, and Bolivia. The phylogeny of the species-diverse Phrynopus is unknown and the position of the genus within Leptodactylidae is poorly understood. We present the results of a phylogenetic study based on 12S and 16S mitochondrial rDNA. Fifteen species of Phrynopus from Bolivia to Ecuador are included, along with several other genera of Leptodactylidae and representatives of other frog families. Our results indicate that Phrynopus is phylogenetically nested within Eleutherodactylus , whereas Phyllonastes is phylogenetically nested within Phrynopus . Based on the recovered phylogeny, we transfer Phrynopus simonsii to Eleutherodactylus , and show that Phrynopus carpish needs to be removed from Phrynopus . Eleven of the 13 Phrynopus species occurring in Central Peru share the absence of an external and internal ear.     

Presence of cushion plants increases community diversity in the high equatorial Andes

Cushion plants are a common growth form in the equatorial páramo vegetation and their surfaces are often colonized by other plants. This paper analyzes the effect of the cushion plants on the community diversity at 4650 m on the eastern slope of the Iliniza volcano in Ecuador. Ninety sample plots of 1 m² size were located in the study area and were divided into 25 subplots in which presence and abundance of plant species was recorded. The community diversity was expressed as species richness, Simpson's diversity index, and evenness. Correlation between the cushion species and the composition of the colonists was measured with the CCA ordination analysis, correlation between the cushion cover and community diversity was measured by means of correlation analysis. Randomized species–area curves were used to compare richness of plant communities with and without the cushions. A total of 32 species were found including five cushion plants. Most species preferred to grow on the cushion surface whereas only a few species were able to colonize open ground. Species richness and Simpson's index were significantly correlated to the cushion area but no correlation was found for evenness. The cushions were usually composed of more than one species which hampered the examination of the cushion–colonist specific relationships. Nevertheless, cushions of Azorella and Arenaria seemed to provide more favorable habitat for colonization than the other cushion species. Comparison with an earlier study made on Iliniza indicates that the presence of the cushions significantly increases the richness of the plant community.     

The impact of roads on the avifauna of páramo grasslands in Cajas National Park,Ecuador

National parks are an important tool for conserving biodiversity, particularly in areas of high biodiversity and endemism such as the tropical Andes. However, national parks often face a variety of stressors related to recreation, road construction and illegal extraction of natural resources. Unfortunately, the influence of these stressors for biodiversity is rarely well documented. Cajas National Park in Ecuador is no exception. Despite being traversed by the Cuenca-Molleturo-Naranjal road, effects of the road construction on biodiversity have not been determined. We therefore assessed the influence of road proximity on bird species richness and abundance as well as composition of bird habitat groups in Cajas National Park using transect walks at 25 m and 250 m distance to the road (overall 18 transects, each 1 km length). In total, we recorded 1110 individuals of 28 páramo bird species. Overall species richness did not differ between transects near and far from the road. Nevertheless, the average abundance of shrubby páramo species was significantly higher far from the road than near the road (Far = 36, Near = 25). Moreover, we found a tendency towards differences in the composition of bird habitat groups between transects near and far from the road. One aspect potentially driving the observed patterns was the increasing proportion of planted non-native woody tree species within páramo grassland near the road, which may have caused reduced abundances of shrubby páramo bird species there. While roads represented a clear impact on the composition of bird species in the páramo, the major effect seems to be driven by the introduction of non-native plant species along the roadside. In order to reduce the impact of roads to a minimum, we suggest that park managers should control the introduction of such plant species.     

Late Holocene vegetation, fire, climate and upper forest line dynamics in the Podocarpus National Park, southeastern Ecuador

Late Holocene vegetation, fire, climate and upper forest line dynamics were studied based on detailed pollen and charcoal analyses. Two sediment cores, from the Rabadilla de Vaca mire (RVM) and the Valle Peque?o bog (VP), with an age of about 2100 and 1630 cal yrs b.p., respectively, were taken at the modern upper forest line in the Parque Nacional Podocarpus (Podocarpus National Park) in southeastern Ecuador. The two pollen records reflect relatively stable vegetation with slight changes in floral composition during the recorded period. Changes of the proportion between subpáramo and páramo vegetation are related to lower and higher frequency of fires. The RVM records show that the upper forest line moved to a higher elevation between 1630 and 880 cal yrs b.p., stabilising after 310 cal yrs b.p. Human impact is suggested by a high fire frequency, mainly between 1800–1600 and 880–310 cal yrs b.p. The VP records indicate no marked changes in the upper forest line. The charcoal records suggest an increased human impact from 230 cal yrs b.p. to the present. The results indicate that high fire frequency is an important factor in reducing the expansion of subpáramo vegetation and upper montane rainforest and in favouring the distribution of grass páramo. Since there is a clear correlation between fire and vegetation dynamics, it is difficult to detect how far climate change also played a significant role in upper forest line changes during the late Holocene.     

Hidden biodiversity in the Arctic – a study of soil seed banks at Disko Island,Qeqertarsuaq, West Greenland

Seed dispersal is a key process in plant community dynamics, and soil seed banks represent seed dispersal in time rather than in space. Despite their potential importance, seed bank dynamics in the Arctic are poorly understood. We investigated soil seed banks and corresponding plant community composition in three contrasting vegetation types in West Greenland, viz. dwarf shrub heaths, herb slopes and fell‐fields. Through germination testing, 31 species were documented in soil seed banks. All of these were herbaceous, while no dwarf‐shrub species, which represents the dominating growth form in the above‐ground vegetation, were emerging from the seed bank. Consequently, across vegetation types, the lowest similarity between seed bank and above‐ground vegetation was found in dwarf shrub heath. Nine plant species were exclusively found in seed bank, all of which were non‐clonal forbs. Seed bank size (total number of seeds) and species richness seemed to increase with the level of natural disturbance. Additionally, we examined the effect of different experimental light and temperature conditions on the quantity and diversity of germinating seeds. The difference in diversity in vegetation and seed bank at the species level will impact population dynamics, regeneration of vegetation after disturbances and its potential to respond to climate change.     

Structure and diversity of stream invertebrate assemblages: the influence of temperature with altitude and latitude

1. Structure and diversity of the macroinvertebrate fauna were studied in relation to altitude and latitude among three groups of streams from Ecuador (lowland: 100–600 m, Central Valley: 2600–3100 m, páramo: 3500–4000 m), and one group from the temperate lowland region of Denmark. The streams in the four regions were comparable with regard to physical characteristics such as size, current and substratum.
2. In terms of faunal composition the Ecuadorian highland streams bore more resemblance to the Danish lowland streams than the Ecuadorian lowland streams. The greater similarity between the Ecuadorian highland and the Danish streams, however, was due to the large number of insect families in the Ecuadorian lowlands, many of which were not found in the other regions. Of ten physico-chemical parameters measured, maximum stream temperature explained by far the most variability in faunal composition.
3. The number of insect orders and families increased linearly with maximum stream temperature and therefore decreased with altitude and latitude. A compilation of literature data on insect richness and maximum water temperature from streams around the world confirmed this pattern, yielding a common linear relation for both temperate and tropical streams. This pattern may arise due to a direct temperature effect on speciation but is probably also related to geological history and the influence of climatic changes on stream ecosystems. We estimate that small, tropical, lowland streams have, on average, a two- to fourfold higher species richness than temperate lowland streams.     

Seed dispersal in heterogeneous environments: bridging the gap between mechanistic dispersal and forest dynamics models

Seed dispersal is an important determinant of vegetation composition. We present a mechanistic model of seed dispersal by wind that incorporates heterogeneous vegetation structure. Vegetation affects wind speeds, a primary determinant of dispersal distance. Existing models combine wind speed and fall velocity of seeds. We expand on them by allowing vegetation, and thus wind profiles, to vary along seed trajectories, making the model applicable to any wind-dispersed plant in any community. Using seed trap data on seeds dispersing from forests into adjacent sites of two distinct vegetation structures, we show that our model was unbiased and accurate, even though dispersal patterns differed greatly between the two structures. Our spatially heterogeneous model performed better than models that assumed homogeneous vegetation for the same system. Its sensitivity to vegetation structure and ability to predict seed arrival when vegetation structure was incorporated demonstrates the model's utility for providing realistic estimates of seed arrival in realistic landscapes. Thus, we begin to bridge mechanistic seed dispersal and forest dynamics models. We discuss the merits of our model for incorporation into forest simulators, applications where such incorporation has been or is likely to be especially fruitful, and future model refinements to increase understanding of seed dispersal by wind.     

Superpáramo plant species diversity and phytogeography in Ecuador

In 18 superpáramo sites in Ecuador we found 388 species of vascular plants belonging to 146 genera and 52 families, making the Ecuadorian superpáramo flora richer in species than that of Venezuela and that of Colombia which appears to have fewer species although the number remains uncertain. The most species rich families were Asteraceae (83 species) and Poaceae (49) which also dominate the grasspáramo that surrounds the superpáramo. Otherwise the superpáramo is dominated by families that are mostly herbaceous such as Cyperaceae, Brassicaceae, Caryophyllaceae and Valerianaceae, whereas shrubby families that dominate the subpáramo and grasspáramo, such as Melastomataceae, Ericaceae, and Solanaceae, have only few species or are absent in the superpáramo. The generic spectrum was dominated by a suite of species-rich genera (Lachemilla, Gentianella, Valeriana, Calamagrostis, Draba) with many páramo-endemic species reflecting a high level of autochthonous speciation in the (super)páramo. Species richness varied from 71 to 149 in the individual superpáramos surveyed, but species richness was only weakly correlated to their area and log-transformed area, and negatively correlated to the vertical range of each superpáramo. β-diversity was significantly correlated to the vertical range (i.e., the number of surveyed altitudinal levels) but it was not correlated to the area or log-transformed area of each superpáramo. Most of the species were narrowly distributed and 112 (29%) of them were found in a single superpáramo, while eight (2%) occurred in all 18 superpáramo sites. Floristic similarity was not correlated to the geographical distance between the sites. Redundance Analysis suggested that geological origin of the substrate (metamorphic versus volcanic bedrock) was important for the floristic composition. Occurrence of mountains built from metamorphic rocks, is however, correlated to areas with high rainfall and the amount of rainfall may be a stronger determinant for species distribution than the presence of volcanic versus metamorphic bedrocks. TWINSPAN cluster analysis divided the 18 sites into three groups, which corresponded to dry, humid and very humid superpáramos. The groups were also separated along the first ordination axis in Correspondence Analysis, while the second axis may correlate to the volcanic history of the area.     

Latitudinal and altitudinal patterns of plant community diversity on mountain summits across the tropical Andes

The high tropical Andes host one of the richest alpine floras of the world, with exceptionally high levels of endemism and turnover rates. Yet, little is known about the patterns and processes that structure altitudinal and latitudinal variation in plant community diversity. Herein we present the first continental‐scale comparative study of plant community diversity on summits of the tropical Andes. Data were obtained from 792 permanent vegetation plots (1 m²) within 50 summits, distributed along a 4200 km transect; summit elevations ranged between 3220 and 5498 m a.s.l. We analyzed the plant community data to assess: 1) differences in species abundance patterns in summits across the region, 2) the role of geographic distance in explaining floristic similarity and 3) the importance of altitudinal and latitudinal environmental gradients in explaining plant community composition and richness. On the basis of species abundance patterns, our summit communities were separated into two major groups: Puna and Páramo. Floristic similarity declined with increasing geographic distance between study‐sites, the correlation being stronger in the more insular Páramo than in the Puna (corresponding to higher species turnover rates within the Páramo). Ordination analysis (CCA) showed that precipitation, maximum temperature and rock cover were the strongest predictors of community similarity across all summits. Generalized linear model (GLM) quasi‐Poisson regression indicated that across all summits species richness increased with maximum air temperature and above‐ground necromass and decreased on summits where scree was the dominant substrate. Our results point to different environmental variables as key factors for explaining vertical and latitudinal species turnover and species richness patterns on high Andean summits, offering a powerful tool to detect contrasting latitudinal and altitudinal effects of climate change across the tropical Andes.     

Temporal changes of local and regional processes in the assembly of herbivorous insect communities

Herbivorous insect communities are structured by multiple processes operating locally (e.g. bottom–up effects of plants) and regionally (e.g. dispersal limitation). Although the relative strength of these processes has been well documented, how it varies in time is less understood, especially in relation with the temporal dynamics of plant communities. If temporal turnover of local plant species composition is too rapid for insect community assembly to keep up with, bottom–up effects are expected to be weak. Here, in plant and herbivorous insect communities in Japanese grasslands, we studied how the relative importance of local (bottom–up effects of plants, structures of plant communities and top–down effects of predators) and regional (dispersal limitation) processes varies over the growing season. In addition, we tested the hypothesis that larger temporal turnover of plant species composition is related to the weaker bottom–up effects, that is, the lower explanation power of plant communities for insect communities. We found that, throughout the growing season, the insect species composition was mainly explained by local variables (plant species composition, vegetation height and predator abundance), and their explanation power was higher during later phases of the season (late summer). Furthermore, the variation not explained by plant species composition was correlated with the degree of temporal turnover of plants, suggesting that insect communities failed to track the temporal turnover of plant species. These results were pronounced when we focused on leaf sucker insects, whose host plant range is presumably more limited. We conclude that herbivorous insect communities are mainly regulated by local processes, especially bottom–up effects from plants, while stochasticity may have played a role in early phases of the season. Furthermore, we underscore the importance of considering relative time scale of community assembly and environmental shifts, especially in systems characterized by dynamic changes.     

湿地土壤种子库与地上植被相似性关系研究评述

土壤种子库与地上植被的关系是土壤种子库研究的重要组成部分。当前,湿地生态系统面临严重威胁,研究湿地土壤种子库和地上植被关系既可以加强对土壤种子库和植物群落特征的认识,又可以为湿地保护与管理提供理论指导。检索了科学引文索引扩展版(SCIE)数据库中收录的1900—2012年间研究湿地土壤种子库与地上植被关系的文献,通过分析土壤种子库与地上植被的Srensen相似性系数,结果发现:不同湿地类型的土壤种子库和地上植被的相似性存在显著差异,河流湿地中两者的相似性最小;不同植被类型中土壤种子库与地上植被的相似性差异显著:草本群落的相似性大于乔木群落;不同气候带的湿地中两者的相似性也存在显著差异,其中亚热带地区相似性最小。总结了湿地种子库与地上植被相似性关系的时空变化特征。二者的相似性通常随着植物群落的演替而减小,在空间上也随着环境梯度而变化。分析了两者关系的影响因素,如种子传播、环境条件和繁殖策略等。对研究中存在的问题及发展方向提出建议。     

The relationship between soil seed bank,above‐ground vegetation and disturbance intensity on old‐field successional permanent plots

Questions: How does disturbance and successional age influence richness, size and composition of the soil seed bank? What is the potential contribution of the soil seed bank to the plant community composition on sites differing in their successional age or disturbance intensity? Location: Experimental Botanical Garden of Göttingen University, central Germany. Methods: Above‐ground vegetation and soil seed bank were studied on formerly arable fields in a 36‐year‐old permanent plot study with five disturbance intensities, ranging from yearly ploughing via mowing to long‐term uninterrupted succession. We compared species compositions, seed densities and functional features of the seed bank and above‐ground vegetation by using several methods in parallel. Results: The seed bank was mainly composed of early successional species typical of strongly disturbed habitats. The difference between seed bank composition and above‐ground vegetation decreased with increasing disturbance intensity. The species of greatest quantitative importance in the seed bank was the non‐native forb Solidago canadensis. Conclusions: The ability of a plant community to regenerate from the soil seed bank dramatically decreases with increasing time since abandonment (successional age) and with decreasing disturbance intensity. The present study underlines that plant species typical of grasslands and woodlands are limited by dispersal capacity, owing to low capacity for accumulation of seeds in the soil and the fact that most species do not build up persistent seed banks. Rare and target species were almost absent from the seed bank and will, after local elimination, depend on reintroduction for continuation of their presence.     

Identifying drivers of species compositional change in a semi‐natural upland grassland over a 40‐year period

Question: Few long‐term studies exist with integrated vegetation and soil composition data, coupled with detailed environmental driver records. Can changes in community composition in an upland grassland be identified by revisitation after a 40‐year period and allow the main environmental drivers of change to be identified? Location: Snowdon, Wales, UK. Methods: Changes in plant community and soil composition were assessed by resurveying an upland Agrostis–Festuca grassland in 2008, 40 years after the original survey. PCA and ecological indicators were used to determine changes in plant community composition. Redundancy analysis (RDA) allowed the impact of soil chemical composition on the vegetation community to be assessed. Results: A significant shift in community composition was found between years. A 35% reduction in species richness and an increase in the grass:forb ratio, suggest significant ecosystem degradation. Indicator values suggest acidification of the community with an increased acidity preference of species recorded in 2008. However, soil pH measurements showed that soil pH had increased. RDA suggested that the main shifts in species composition were correlated with an increase in pH and a reduction in soil exchangeable base cation concentration. Clear ecosystem responses to climate, land‐use change or nitrogen enrichment were not observed. Conclusions: Shifts in vegetation and soil composition are clearly identifiable after 40 years. The shifts in community composition are consistent with ecosystem degradation due to acidification during the period between surveys. Ecological indicator values and soil chemical composition displayed differing degrees of change. Whilst soils appear to be recovering from historic effects of sulphur deposition, vegetation community composition changes appear to lag behind those in soil chemistry.     

Factors Controlling Compositional Changes in a Northern Andean Páramo (La Rusia,Colombia)

In this study, we aimed to assess the processes controlling compositional change in a Northern Andean páramo highly affected by human‐induced disturbances over the last few decades (La Rusia, Colombia). Along the 3000–3800 m asl altitudinal range, we randomly sampled fifty 10 × 10 m plots. Therein, we measured altitude and variables related to soil conditions (i.e., moisture, nutrient contents, bulk density, and texture), occurrence of human‐induced disturbances (i.e., fire, vegetation clearing, potato cultivation, and cattle grazing), and land‐use history. We also recorded richness and abundance of plant species, identifying them as exotic or native. We differentiated four groups of plots according to their species composition. The groups had significant differences in altitude, soil conditions, land‐use history, and particularly, in richness of exotic species and exotic/native cover ratio. They could be ascribed to shrub‐ and grass‐páramo vegetation types based on their relative dominance of woody and herbaceous species; however, these groups were not arranged according to the hypothetical composition of altitudinal belts, but rather formed a mosaic of patches. This mosaic was determined not only by altitude but also by soil conditions and disturbance history of sites. Our results corroborate recent findings which highlight shrub‐ and grass‐páramo vegetation types as patches of contrasting species composition and structure that depend on local environmental variables, as well as human‐induced disturbances as a major determinant of compositional discontinuities in these ‘high mountain’ tropical ecosystems.     

Environmental Controls on Fungal Community Composition and Abundance Over 3 Years in Native and Degraded Shrublands

Soil fungal communities have high local diversity and turnover, but the relative contribution of environmental and regional drivers to those patterns remains poorly understood. Local factors that contribute to fungal diversity include soil properties and the plant community, but there is also evidence for regional dispersal limitation in some fungal communities. We used different plant communities with different soil conditions and experimental manipulations of both vegetation and dispersal to distinguish among these factors. Specifically, we compared native shrublands with former native shrublands that had been disturbed or converted to pasture, resulting in soils progressively more enriched in carbon and nutrients. We tested the role of vegetation via active removal, and we manipulated dispersal by adding living soil inoculum from undisturbed native sites. Soil fungi were tracked for 3 years, with samples taken at ten time points from June 2006 to June 2009. We found that soil fungal abundance, richness, and community composition responded primarily to soil properties, which in this case were a legacy of plant community degradation. In contrast, dispersal had no effect on soil fungi. Temporal variation in soil fungi was partly related to drought status, yet it was much broader in native sites compared to pastures, suggesting some buffering due to the increased soil resources in the pasture sites. The persistence of soil fungal communities over 3 years in this study suggests that soil properties can act as a strong local environmental filter. Largely persistent soil fungal communities also indicate the potential for strong biotic resistance and soil legacies, which presents a challenge for both the prediction of how fungi respond to environmental change and our ability to manipulate fungi in efforts such as ecosystem restoration.     

Distinguishing signal from noise: Long-term studies of vegetation in Makhtesh Ramon erosion cirque,Negev desert,Israel

We consider three case studies of long-term plant population dynamics in Makhtesh Ramon erosion cirque in the central Negev desert of Israel. We show that rainfall is the major driving variable in this system, and that it creates large temporal and spatial variation in plant species diversity and vegetation community composition. This variability makes it extremely difficult to distinguish `signal' (= pattern in vegetation) from `noise' (random spatial and temporal variance). Our long-term vegetation studies in permanent plots arranged along the length of the cirque, initiated in 1990 and continuing, show that there is high spatial and temporal variance in plant species' incidences and abundances. This is particularly true of annual plant species. However, using pairs of fenced and unfenced plots arranged along the major abiotic gradient, altitude (which mirrors changes in rainfall), we were able to tease apart the effects of variance in rainfall and herbivory. We found significant negative effects of herbivory by the re-introduced Asiatic wild ass (Equus hemionus) on plant cover and on vegetation community composition. In a study of the effects of herbivory by dorcas gazelles (Gazella dorcas) on the lily Pancratium sickenbergeri, we found that there was little inter-annual fluctuation in lily population size in two sand dunes in Makhtesh Ramon in spite of the high percentage of lilies that is removed by the gazelles and the almost complete herbivory of flowers by these herbivores. This result indicates that the dune lily populations may be maintained by dispersal of seed from other lily populations elsewhere in Makhtesh Ramon where gazelles are rare or absent.     

Seed Dispersal Modes of the Sandstone Plateau Vegetation of the Middle Caquetá River Region,Colombian Amazonia1

We characterized the dispersal spectra and phenology of 298 vascular plant species of the sandstone plateaus of Colombian Amazonia. Dispersal modes were determined by the morphology of dispersion units, personal observations on fruit consumption, and an extensive literature review. We obtained the number of species per dispersal mode for the sandstone plateaus and for two recognized vegetation types: open‐herbaceous vegetation and low forest‐shrub vegetation. Dispersal modes were assigned to 295 plant species. Animals dispersed the highest percentage of species (46.6%), while the percentage of autochorous and anemochorous species was 29.4 and 23 percent, respectively. The dispersal spectrum of the low forest‐shrub vegetation type, based on the coverage of every species, showed that percentages of anemochorous (40.2%) and zoochorous species (37.8%) were similar. Autochory was the most important seed dispersal mode of the open‐herbaceous vegetation (60%). Birds were the principal group of potential dispersers (58.9%) of zoochorous species and reptiles the least important. We found two marked fruiting peaks, one from the end of the dry season to the beginning of the wet season and the second one from the beginning to the middle of the dry season. Our results showed that besides the differences in the vegetation structure and floristic composition between the sandstone plateaus and the adjacent tall forest, there also exist differences in the dispersal spectra and the fruiting rhythms.