Grazing as a measure against grass encroachment in Dutch dry dune grassland: effects on vegetation and soil

A grazing experiment was started in 1984 and 1989 respectively, in two parts of a dune grassland in the nature reserve ‘Zwanenwater’, North Holland; a third part with similar geology and topography was used as a control area and not grazed. An evaluation of the effects of grazing on vegetation patterns, species composition, vegetation structure and humus form was made with the help of vegetation maps from 1986 and 1992 as well as field surveys. Dense tall-grass communities dominated byAmmophila arenaria increased over the period 1986–1992 in the grazed areas, and especially in the non-grazed area (increase in area to 20%, 22% and 50%, respectively). Open communities decreased in the grazed areas, but are still prevalent, while in the ungrazed area they virtually disappeared, with the result that the present percentage areas are 53%, 38% and 17%. Field survey data were classified by TWINSPAN producing four vegetation types. These occur more or less equally in grazed and ungrazed areas, albeit with different percentage areas: (1) open vegetation dominated byCorynephorus canescens; (2) open vegetation characteized, byKoeleria macrantha; (3) heathland dominated byEmpetrum nigrum; and (4) tall-grass communities dominated byAmmophila arenaria. Within a vegetation type, species composition was only marginally affected by grazing regime. Within the open communities the number of species, vegetation height, vegetation cover and soil organic horizons were not affected by grazing. In the tall-grass communities the number of species was significantly larger and the height of the vegetation significantly lower in the area grazed since 1984. In the heathland community the number of species and cover of the moss layer were significantly higher in the 1984 area and ectorganic and endorganic horizons significantly thicker in the ungrazed area. It is suggested that these effects are the result of an increased availability of light, but possibly also of a decreased stock of organic matter and nutrients, due to a decreased input of litter and accelerated rates of decomposition.     

Grazing as a measure against grass encroachment in Dutch dry dune grassland: effects on vegetation and soil

A grazing experiment was started in 1984 and 1989 respectively, in two parts of a dune grassland in the nature reserve ‘Zwanenwater’, North Holland; a third part with similar geology and topography was used as a control area and not grazed. An evaluation of the effects of grazing on vegetation patterns, species composition, vegetation structure and humus form was made with the help of vegetation maps from 1986 and 1992 as well as field surveys. Dense tall-grass communities dominated byAmmophila arenaria increased over the period 1986–1992 in the grazed areas, and especially in the non-grazed area (increase in area to 20 %, 22 % and 50 %, respectively). Open communities decreased in the grazed areas, but are still prevalent, while in the ungrazed area they virtually disappeared, with the result that the present percentage areas are 53 %, 38 % and 17 %. Field survey data were classified by TWINSPAN producing four vegetation types. These occur more or less equally in grazed and ungrazed areas, albeit with different percentage areas: (1) open vegetation dominated byCorynephorus canescens; (2) open vegetation characterized byKoeleria macrantha; (3) heathland dominated byEmpetrum nigrum; and (4) tall-grass communities dominated byAmmophila arenaria. Within a vegetation type, species composition was only marginally affected by grazing regime. Within the open communities the number of species, vegetation height, vegetation cover and soil organic horizons were not affected by grazing. In the tall-grass communities the number of species was significantly larger and the height of the vegetation significantly lower in the area grazed since 1984. In the heathland community the number of species and cover of the moss layer were significantly higher in the 1984 area and ectorganic and endorganic horizons significantly thicker in the ungrazed area. It is suggested that these effects are the result of an increased availability of light, but possibly also of a decreased stock of organic matter and nutrients, due to a decreased input of litter and accelerated rates of decomposition. Nomenclature: Van der Meijden et al. (1992) for phanerogams; Corley et al. (1981) for mosses; Grolle (1983) for hepatics.     

Differences in gypsum plant communities associated with habitat fragmentation and livestock grazing.

The negative consequences of habitat fragmentation for plant communities have been documented in many regions of the world. In some fragmented habitats, livestock grazing has been proposed to be a dispersal mechanism reducing isolation between fragments. In others, grazing acts together with fragmentation in a way that increases habitat degradation. Iberian gypsum plant communities have been grazed and fragmented by agricultural practices for centuries. Although their conservation is considered a priority by the European Community, the effects of fragmentation on gypsum plant communities and the possible role of livestock grazing remain unknown. In addition, a substantial proportion of plant species growing in gypsum environments are gypsum specialists. They could be particularly affected by fragmentation, as was found for other habitat specialists (i.e., serpentine and calcareous specialists). In this study (1) we investigated the effect of fragmentation and grazing on gypsum plant community composition (species and life-forms), and (2) we tested to see if gypsum specialists were differently affected by fragmentation and grazing than habitat generalists. A vegetation survey was conducted in the largest gypsum outcrop of Europe (Middle Ebro Valley, northeast Spain). Fragmented and continuous sites in grazed and ungrazed areas were compared. Measurements related to species and composition of life-forms were contrasted first for the whole gypsum plant community and then specifically for the gypsum specialists. In the whole community, our results showed lower plant species diversity in fragmented sites, mainly due to the larger dominance of species more tolerant to fragmented habitat conditions. With livestock grazing, the plant species richness and the similarity in plant species composition between remnants was larger, suggesting that animals were acting as dispersal agents between fragments. As expected, gypsum specialists were less abundant in fragmented areas, and grazing led to the disappearance of the rare gypsum specialist Campanula fastigiata. According to our results, conservation strategies for gypsum plant communities in human-dominated landscapes should consider that fragmentation and grazing modify plant community composition affecting gypsum specialists in particular.     

The impact of grazing on spider communities in a mesophytic calcareous dune grassland

During 1994–1995 and 1997–1998 spiders were sampled with pitfall traps in a botanically rich, mesophytic, calcareous dune grassland in Belgium. As a consequence of intensive cattle grazing, vegetation variation in a large part of the area had diminished. The study area was also patchily grazed by rabbits. Community analysis with TWINSPAN revealed five distinct spider communities. Ecological differentiation was best explained by combination of the habitat variables: distance from grazed or non-grazed vegetation,Rosa pimpinellifolia cover and grass cover in both summer and winter. Species diversity was highest in the border zone between the cattle-grazed and non cattle-grazed sites. Correlation of the most abundant spider species with the vegetation determinants explains the ecological differentiation between the spider communities. Species were classified into seven major groups that reflect the species’ habitat preferences. The group showing clear association with non cattle-grazed, tall vegetation consists of common species. Characteristic species for the intensively cattle-grazed sites are common aeronauts and rare species such asWalckenaeria stylifrons, Mastigusa arietina, Ceratinopsis romana andPardosa monticola. The latter are shown to be dependent on ungrazed vegetation for juvenile development and overwintering. Intensive grazing results in homogeneous short vegetation, which can only be colonized by ‘open ground’ species with a well-developed dispersal capacity, or by species which are not dependent on litter-rich situations for juvenile development. An extensive cattle grazing regime results in a patchy mosaic grassland where, in addition to the above mentioned groups of species, other species survive by migrating between the buffered litter rich ungrazed vegetation and the short vegetation. Additionally, some typical and rare species prefer the transition zone between the grazed and the ungrazed vegetation because they are associated with specific habitat structures or inhabiting ant-species.     

The impact of grazing on spider communities in a mesophytic calcareous dune grassland

During 1994–1995 and 1997–1998 spiders were sampled with pitfall traps in a botanically rich, mesophytic, calcareous dune grassland in Belgium. As a consequence of intensive cattle grazing, vegetation variation in a large part of the area had diminished. The study area was also patchily grazed by rabbits. Community analysis with TWINSPAN revealed five distinct spider communities. Ecological differentiation was best explained by combination of the habitat variables: distance from grazed or non-grazed vegetation,Rosa pimpinellifolia cover and grass cover in both summer and winter. Species diversity was highest in the border zone between the cattle-grazed and non cattle-grazed sites. Correlation of the most abundant spider species with the vegetation determinants explains the ecological differentiation between the spider communities. Species were classified into seven major groups that reflect the species’ habitat preferences. The group showing clear association with non cattle-grazed, tall vegetation consists of common species. Characteristic species for the intensively cattle-grazed sites are common aeronauts and rare species such asWalckenaeria stylifrons, Mastigusa arietina, Ceratinopsis romana andPardosa monticola. The latter are shown to be dependent on ungrazed vegetation for juvenile development and overwintering. Intensive grazing results in homogeneous short vegetation, which can only be colonized by ‘open ground’ species with a well-developed dispersal capacity, or by species which are not dependent on litter-rich situations for juvenile development. An extensive cattle grazing regime results in a patchy mosaic grassland where, in addition to the above mentioned groups of species, other species survive by migrating between the buffered litter rich ungrazed vegetation and the short vegetation. Additionally, some typical and rare species prefer the transition zone between the grazed and the ungrazed vegetation because they are associated with specific habitat structures or inhabiting ant-species. Nomenclature: Roberts (1987, 1995) forAraneae; van der Meijden et al. (1990) for vascular plants; Corly et al. (1981) for bryophytes; Schaminée et al. (1996) for vegetation associations.     

Synergisms among Habitat Fragmentation, Livestock Grazing, and Biotic Invasions in Southwestern Australia

Abstract: The agricultural development of southern Australia over the past 200 years has resulted in extensively fragmented systems, often with only small, isolated remnants of native vegetation remaining. Grazing by sheep and cattle has affected both the remaining fragments and the surrounding matrix, and non-native plant and animal species have had dramatic effects on the native biota. Invasive plant species have the potential to significantly alter ecosystem composition and functioning, and invasive animals, particularly rabbits ( Oryctolagus cuniculatus ), foxes (    Vulpes vulpes ) and cats (    Felis catus ) effectively alter habitat and drive native fauna to local extinction. These different influences often interact. For instance, smaller fragments are often more prone to plant invasion and are more likely to have been grazed in the past. Invasion of plant species is often linked with livestock grazing or rabbit invasion, and other higher-order interactions are also apparent. Classical fragmentation studies that concentrate on parameters such as habitat area and isolation but ignore changes in habitat condition brought about by livestock and invasive species are unlikely to yield meaningful results. Similarly, management of fragmented ecosystems must account for not only the spatial characteristics of the remaining habitat but also the importance of other influences, particularly those that impinge on fragments from the surrounding matrix.     

Cattle Grazing Impacts on Annual Forbs and Vegetation Composition of Mesic Grasslands in California

Abstract:   Livestock grazing represents a major human alteration of natural disturbance regimes in grasslands throughout the world, and its impacts on plant communities have been highly debated. We investigated the impact of cattle grazing on the California coastal prairie plant community with a focus on native annual forbs, a number of which are of conservation concern. In spring 2000 and 2001, we surveyed the vegetation community composition, vegetation structure, and soil chemical parameters at 25 paired grazed and ungrazed sites over a 670-km range of the ecosystem. Native annual forb species richness and cover were higher in grazed sites, and this effect was concomitant with decreased vegetation height and litter depth. Soil properties explained less of the variation. Exotic annual grass and forb cover were higher in grazed sites. Native grass cover and species richness did not differ in grazed and ungrazed sites, but cover and species richness of native perennial forbs were higher in ungrazed sites. Our results suggest that cattle grazing may be a valuable management tool with which to conserve native annual forbs in the ecosystem we studied but that grazing differentially affects the various life-history guilds. Therefore, land managers must focus on creating a matrix of disturbance regimes to maintain the suite of species native to these mesic grasslands. The results of this and other studies highlight the importance of considering the adaptation of vegetation communities to disturbance in making recommendations for grazing management.     

Effects of Grazing on the Demography and Growth of the Texas Tortoise

Abstract: Considerable effort has been exerted in attempts to understand the complex ecological effects of grazing. North American tortoises, by virtue of their distribution, provide a good model taxon through which to study how grazing effects vary with grazing regime, habitat, and climate. We studied the Texas tortoise (  Gopherus berlandieri ) , which is restricted primarily to privately owned rangelands of southern Texas and northeastern Mexico. Management of this species is hampered by a lack of information on the effects of common land-use practices. We evaluated the effects of moderate grazing by cattle (short-duration, winter-spring rotational grazing regime; 6–28 animal-unit days/ha/year) on this tortoise by comparing two grazed and two ungrazed sites in the Western Rio Grande Plains, Texas ( U.S.A.), from April 1994 to October 1997. We made 132 captures of 106 individuals in the ungrazed pastures and 324 captures of 237 individuals in the grazed pastures. We also radiotracked 22 tortoises in the ungrazed pastures and 25 tortoises in the grazed pastures. Comparisons of relative abundance, body-size distribution, age distribution, body mass, sex ratio, adult survival, proportion of juveniles, and growth rates revealed no differences (  p > 0.05 for all parameters) between tortoises on grazed and ungrazed areas. Based on these results, we suggest that moderate grazing by cattle is not incompatible with maintenance of Texas tortoise populations. Our data were consistent with a general model of tortoise biogeography and tolerance of disturbance which suggests that Texas tortoises are tolerant to intermediate levels of disturbance. Generalities about the effect of cattle grazing on the four North American tortoises should be avoided unless they can be placed in the context of grazing regime, precipitation, habitat quality, and tortoise requirements.     

Rodent Communities in an Exurbanizing Southwestern Landscape (U.S.A.)

Abstract:  Ranches are being converted to exurban housing developments in the southwestern United States, with potentially significant but little-studied impacts on biological diversity. We captured rodents on 48 traplines in grasslands, mesquite savannas, and oak savannas in southeastern Arizona that were grazed by livestock, embedded in exurban housing developments, grazed and embedded in development, or neither grazed nor embedded in development. Independent of habitat or development, rodent species richness, mean rank abundance, and capture rates of all rodents combined were negatively related to presence of livestock grazing or to its effects on vegetative ground cover. Exurban development had no obvious effects on rodent variety or abundance. Results suggest southwestern exurban developments can sustain a rich assemblage of grassland and savanna rodents if housing densities are low and houses are embedded in a matrix of natural vegetation with little grazing.     

Impacts of Livestock Grazing and Tree Clearing on Birds of Woodland and Riparian Habitats

Abstract:  We investigated the impact of pastoral management on birds in subtropical grassy eucalypt woodland in southeastern Queensland, Australia, where the patterns of land management have made it possible to disentangle the effects of livestock grazing from those of tree clearing. We recorded changes in bird species composition, density, and relative abundance across two woodland habitat types (riparian and nonriparian) and two levels of clearing (wooded and nonwooded) and three levels of livestock grazing (low, moderate, and high) replicated over space (1000 km²) and time (2001–2002). We predicted that species that depend on understory vegetation would be most negatively affected by livestock grazing. A Bayesian generalized linear model showed that the level of grazing had the greatest effect when trees were present. When trees were absent, the impact of grazing was overshadowed by the effects of a lack of trees. Over 65% of species responded to different levels of grazing, and the abundance of 42% of species varied markedly with habitat and grazing. The most common response to grazing was high species relative abundance under low levels of grazing (28% of species), species absence at high levels of grazing (20%), and an increase in abundance with increasing grazing (18%). Despite having similar bird assemblages, the effect of grazing was stronger in riparian habitat than in adjacent woodland habitat. Our results suggest that any level of commercial livestock grazing is detrimental to some woodland birds, particularly the understory-dependant species, as predicted. Nevertheless, provided trees are not cleared, a rich and abundant bird fauna can coexist with moderate levels of grazing. Habitats with high levels of grazing, on the other hand, resulted in a species-poor bird assemblage dominated by birds that are increasing in abundance nationally .     

The effect of grazing on biodiversity in coastal dune heathlands

The effect of sheep grazing on species richness, higher order diversity measures, inequality, species composition, functional diversity and allometric relationships at a coastal dune heathland site was investigated. After a prescribed fire in 2002, the site was divided into two parts, where one of the parts was unmanaged and the other part was fenced and grazed by sheep. Sheep grazing had a positive effect on species richness as well as a significant positive effect on the functional diversity at the coastal dune heathland site. Generally, the cover of dwarf shrubs was negatively affected by grazing, whereas the cover of sedges and grasses was positively affected by grazing. There is a need for comparative investigations of the effect of different management methods not only on floristic biodiversity, but on all relevant kinds of biodiversity as well as on soil structure, soil chemistry and habitat micro- and macrostructure. Consequently, we advocate the initiation of an international systematic investigation of the effect of different management methods.     

Key Role of European Rabbits in the Conservation of the Western Mediterranean Basin Hotspot

Abstract: The Mediterranean Basin is a global hotspot of biodiversity. Hotspots are said to be experiencing a major loss of habitat, but an added risk could be the decline of some species having a special role in ecological relationships of the system. We reviewed the role of European rabbits (Oryctolagus cuniculus) as a keystone species in the Iberian Peninsula portion of the Mediterranean hotspot. Rabbits conspicuously alter plant species composition and vegetation structure through grazing and seed dispersal, which creates open areas and preserves plant species diversity. Moreover, rabbit latrines have a demonstrable effect on soil fertility and plant growth and provide new feeding resources for many invertebrate species. Rabbit burrows provide nest sites and shelter for vertebrates and invertebrates. In addition, rabbits serve as prey for a number of predators, including the critically endangered Iberian lynx (Lynx pardinus) and Spanish Imperial Eagle (Aquila adalberti). Thus, the Mediterranean ecosystem of the Iberian Peninsula should be termed “the rabbit's ecosystem.” To our knowledge, this is the first empirical support for existence of a multifunctional keystone species in a global hotspot of biodiversity. Rabbit populations have declined drastically on the Iberian Peninsula, with potential cascading effects and serious ecological and economic consequences. From this perspective, rabbit recovery is one of the biggest challenges for conservation of the Mediterranean Basin hotspot.     

Vegetation development influenced by grazing in the coastal dunes near The Hague, The Netherlands

In 1990, grazing was introduced in a section of Meijendel, a coastal sand dune system near The Hague, The Netherlands. After five years an evaluation was made of the effects of grazing on vegetation development. Three transects were established, two in grazed areas and one in an ungrazed area. Field survey data were classified by means of TWIN-SPAN, ordinated with Detrended Correspondence Analysis and the resulting vegetation types interpreted according to Westhoff & den Held (1969). All associations were found in both the grazed and the ungrazed areas, but at the subassociation and variant level some communities appeared to be restricted to the grazed area. These variants were five grassland variants characterized by disturbance indicators such asSenecio sylvaticus andCynoglossum officinale. The total number of plant species in the 19 permanent plots, which had been observed to have been decreasing since 1960, showed a considerable increase after the introduction of horses and cows in 1990. A marked decrease in the cover ofCalamagrostis epigejos andCarex arenaria since 1990 was evident, while in some plots species such asRibes rubrum andViburnum opulus increased considerably. A series of false-colour aerial photographs were used to compare vegetation structure in the three transects between 1990 and 1995. In the grazed area the tall grass vegetation had almost totally disappeared, whereas the areas of open sand. sand with moss and lichens, and low grass vegetation had increased and the pattern had become more fine-grained. In the ungrazed area the area covered by low grass vegetation had increased at the expense of the area of sand with moss and lichens and the pattern had become more coarse-grained.     

Impact of cattle grazing on the occupancy of a cryptic, threatened rail

Impacts of livestock grazing in arid and semiarid environments are often concentrated in and around wetlands where animals congregate for water, cooler temperatures, and green forage. We assessed the impacts of winter-spring (November-May) cattle grazing on marsh vegetation cover and occupancy of a highly secretive marsh bird that relies on dense vegetation cover, the California Black Rail (Laterallus jamaicensis coturniculus), in the northern Sierra Nevada foothills of California, U.S.A. Using detection-nondetection data collected during repeated call playback surveys at grazed vs. ungrazed marshes and a "random changes in occupancy" parameterization of a multi-season occupancy model, we examined relationships between occupancy and habitat covariates, while accounting for imperfect detection. Marsh vegetation cover was significantly lower at grazed marshes than at ungrazed marshes during the grazing season in 2007 but not in 2008. Winter-spring grazing had little effect on Black Rail occupancy at irrigated marshes. However, at nonirrigated marshes fed by natural springs and streams, grazed sites had lower occupancy than ungrazed sites. Black Rail occupancy was positively associated with marsh area, irrigation as a water source, and summer vegetation cover, and negatively associated with marsh isolation. Residual dry matter (RDM), a commonly used metric of grazing intensity, was significantly associated with summer marsh vegetation cover at grazed sites but not spring cover. Direct monitoring of marsh vegetation cover, particularly at natural spring- or stream-fed marshes, is recommended to prevent negative impacts to rails from overgrazing.     

Effect of Vertebrate Grazing on Plant and Insect Community Structure

Abstract: We compared species diversity of plants and insects among grazed and ungrazed areas of Ponderosa pine–grassland communities in Arizona. Plant species richness was higher in two of three grassland communities that were grazed by native elk and deer and domestic cattle than in ungrazed areas inside a series of three large (approximately 40-ha) grazing exclosures. Similarly, plant species richness was higher in grazed areas relative to ungrazed areas at one of two series of smaller (approximately 25-m²) and short-term exclosure sites. Evenness of plant distribution, however, was greater inside ungrazed long-term exclosures but was reduced inside ungrazed short-term exclosures relative to grazed areas. Relative abundances of forbs, grasses, trees, and shrubs, and native and introduced plants did not differ between the long- and short-term grazing exclosures and their grazed counterparts. Relative abundances of some plant species changed when grazers were excluded, however. In contrast, insect species richness was not different between grazed and ungrazed habitats, although insect abundance increased 4- to 10-fold in ungrazed vegetation. Our results suggest that vertebrate grazing may increase plant richness, even in nutrient-poor, semi-arid grasslands, but may decrease insect abundances.     

The effects of shifting vegetation mosaics on habitat suitability for coastal dune fauna—a case study on sand lizards (Lacerta agilis)

Encroachment of tall grasses and shrubs in coastal dunes has resulted in loss of vegetation heterogeneity. This is expected to have negative effects on animal diversity. To counteract encroachment and develop structural heterogeneity grazing is a widely used management practice. Here, we aim to functionally interpret changes in vegetation composition and configuration following grazing management on habitat suitability for sand lizards. Aerial photographs taken over a period of 16 years were used to quantify changes in vegetation composition. A GIS-based method was developed to calculate habitat suitability for sand lizards in a spatially explicit manner, encompassing differences in vegetation structure and patch size. From 1987 to 2003 dune vegetation shifted from small patches of moss and sand to larger patches covered by shrubs and grasses. Grazing management did not have any significant effect on the overall level of heterogeneity, measured as habitat suitability for sand lizards. However, on a more local scale highly suitable patches in 1987 were deteriorating whereas unsuitable patches became more suitable in 2003. This inversion results from a broad shift with shrubs being a limiting habitat element in 1987 to sandy patches being the limiting element in 2003. Future changes are believed to negatively impact sand lizards. The habitat suitability model has proven to be a useful tool to functionally interpret changes in coastal dune vegetation heterogeneity from an animal’s perspective. Further research should aim to include multiple species operating on different scale levels to fully capture the natural landscape dynamics.     

Modelling trade-offs between livestock grazing and wader conservation in a grassland agroecosystem

The objective of this study was to model the trade-offs between ecological and productive performance of a grassland agroecosystem. We developed a dynamic model linking grass dynamics controlled by livestock grazing to stochastic population dynamics of two wader bird species with high conservation value. Bird dynamics were driven by both direct and indirect effects of grazing. Viable control framework was used to predict grazing strategies ensuring production and conservation and to generate the whole trade-off curve between ecological and productive performance. Bird population size was used as an indicator of ecological performance and the total number of grazing days defined productive performance. Model results show that conciliating ecological and productive performance implies a temporal shift in grazing sequences. The best ecological performance was obtained at intermediate levels of productive performance. Without grazing or with too low a grazing intensity, it was not possible to maintain any of the bird populations due to the indirect effects of grazing on habitat quality. However, too high a level of grazing implied fast population decline for both populations due to direct negative effects of grazing on nest survival. Field data on current grazing regimes and grass height showed a low proportion of suitable fields in our case study area. This result indicates an antagonism between direct and indirect effects of grazing on wader birds, implying the need of very specific management of grassland. Our results illustrate the fact that European grasslands are anthropic habitats which are highly dependent on human activity. In such habitats, trade-off curves are not expected to be strictly decreasing or increasing.     

Influence of a dominant consumer species reverses at increased diversity

Theory and experiments indicate that changes in consumer diversity affect benthic community structure and ecosystem functioning. Although the effects of consumer diversity have been tested in the laboratory and the field, little is known about effects of consumer diversity in the subtidal zone, one of the largest marine habitats. We investigated the grazing effects of sea urchins on algal abundance and benthic community structure in a natural subtidal habitat of the Galápagos Islands. Three species of urchins (Eucidaris, Lytechinus, and Tripneustes) were manipulated in inclusion cages following a replacement design with three levels of species richness (one, two, and three species) with all possible two-species urchin combinations. Identity was the main factor accounting for changes in the percentage of substrate grazed and benthic community structure. Two out of the three two-species assemblages grazed more than expected, suggesting a richness effect, but analyses revealed that this increased grazing was due to a sampling effect of the largest and commercially valued urchin species, Tripneustes. Benthic community structure in treatments with Eucidaris, Lytechinus, and Tripneustes alone was significantly different at the end of the experiment, suggesting that resource use differentiation occurred. Communities in Tripneustes enclosures were characterized by abundant crustose coralline algae and grazed substrate, while those without it contained abundant green foliose algae (Ulva sp.). An unexpected emergent property of the system was that the most species-rich urchin assemblage underyielded, grazing less than any other assemblage with Tripneustes, effectively reversing its dominant influence observed in the two-species treatments. While further experiments are needed to discern the mechanisms of underyielding, it may be related to changing interspecific interactions as richness increases from two to three species or to density-dependent Tripneustes grazing. This study highlights the general importance of evaluating consumer richness effects across the entire range of species richness considered, as the performance of the most species-rich consumer assemblage could not be predicted by manipulations of intermediate levels of consumer species richness.     

Self-organized spatial patterns of vegetation in alpine grasslands

There is increased interest in vegetation spatial pattern as an indicator of transition shifts following catastrophes. Much, however, remains unknown about the mechanisms that underlie spatial pattern formations. In this study, we examined how the spatial heterogeneity of species distributions in the grasslands of the Central Pyrenees and Middle Atlas Mountains is associated with plant species diversity and the importance of self-organizing processes in the control of pattern formations. In the grasslands of the Central Pyrenees and Middle Atlas, the spatial heterogeneity of species distributions increased along a habitat degradation gradient defined by an increase in bare soil. In Central Pyrenees grasslands, however, the increase in heterogeneity was associated with self-organizing bare soil formations, rather than the self-organizing distribution of plant species. In Middle Atlas grasslands, the increased heterogeneity of species spatial distributions was a consequence of the self-organizing capacity of the composing species; the increase in bare soil was randomly distributed. In the more heavily grazed grasslands (Middle Atlas), but not in the more lightly grazed and better preserved ecosystem (Central Pyrenees), plant species richness and diversity declined significantly with an increase in grazing pressure because fewer species were able to colonize empty space. On the contrary, the colonization of bare soil by new species increased the diversity and spatial organization of new colonizing species in Central Pyrenees grassland.     

Anthropogenic threats to evolutionary heritage of angiosperms in the Netherlands through an increase in high-competition environments

Present biodiversity comprises the evolutionary heritage of Earth's epochs. Lineages from particular epochs are often found in particular habitats, but whether current habitat decline threatens the heritage from particular epochs is unknown. We hypothesized that within a given region, humans threaten specifically habitats that harbor lineages from a particular geological epoch. We expect so because humans threaten environments that dominated and lineages that diversified during these epochs. We devised a new approach to quantify, per habitat type, diversification of lineages from different epochs. For Netherlands, one of the floristically and ecologically best-studied regions, we quantified the decline of habitat types and species in the past century. We defined habitat types based on vegetation classification and used existing ranking of decline of vegetation classes and species. Currently, most declining habitat types and the group of red-listed species are characterized by increased diversification of lineages dating back to Paleogene, specifically to Paleocene-Eocene and Oligocene. Among vulnerable habitat types with large representation of lineages from these epochs were sublittoral and eulittoral zones of temperate seas and 2 types of nutrient-poor, open habitats. These losses of evolutionary heritage would go unnoticed with classical measures of evolutionary diversity. Loss of heritage from Paleocene-Eocene became unrelated to decline once low competition, shade tolerance, and low proportion of non-Apiaceae were accounted for, suggesting that these variables explain the loss of heritage from Paleocene-Eocene. Losses of heritage from Oligocene were partly explained by decline of habitat types occupied by weak competitors and shade-tolerant species. Our results suggest a so-far unappreciated human threat to evolutionary heritage: habitat decline threatens descendants from particular epochs. If the trends persist into the future uncontrolled, there may be no habitats within the region for many descendants of evolutionary ancient epochs, such as Paleogene.