Chimpanzees detect ant-inhabited dead branches and stems: a study of the utilization of plant–ant relationships in the Mahale Mountains, Tanzania

Chimpanzees in the Mahale Mountains of Tanzania consume several species of stem- and branch-inhabiting ants throughout the year, without tools. Those ants are cryptic species, and it was unknown how to find them constantly. There has been little research on how the chimpanzees locate these ants. In this study, I use behavioral observations of the chimpanzee predators and surveys of the ant fauna and plants across different habitats to test the hypothesis that chimpanzees use plant species as a cue to efficiently locate ant colonies in litter units (dead parts of the plant). Ants were found to be associated with live plants and with spaces within litter units which provide nesting places. Such ant–plant litter relationships were not necessarily as strong as the mutualism often observed between live plants and ants. The proportion of available litter units inhabited by ants was 20 %, and litter units of three plant species (Vernonia subligera, Dracaena usambarensis, and Senna spectabilis) were well occupied by ants in the home range of the chimpanzees. The ant-inhabited ratio in chimpanzee-foraged litter units was higher than that in the available units in the home range. Chimpanzees fed more often on Crematogaster spp. than on other resident ants and at a higher rate than expected from their occurrence in the litter units. Above three plant species were well occupied by Crematogaster sp. 3 or C. sp. 18. It is concluded that chimpanzees locate ants by selecting litter units of plant species inhabited by ants.     

Contribution of cocoa plantations to the conservation of native ants (Insecta: Hymenoptera: Formicidae) with a special emphasis on the Atlantic Forest fauna of southern Bahia,Brazil

By maintaining a forest-like structure, shaded cocoa plantations contribute to the conservation of ants that usually live in the soil, leaf litter or canopy of tropical forests. Here we synthesize the available information on the diversity and community structure of ants in shaded cocoa plantations in the Atlantic forest region of Brazil, compare ant assemblages in cocoa agroforests with forests and other forms of agriculture, and discuss how these shaded plantations contribute to the conservation of the ants in the Atlantic Forest region. We also discuss ants of economical importance and of special interest, including Camponotus, Dolichoderus, Gnamptogenys, Pachycondyla, Pseudomyrmex and other litter dwelling genera. We discuss the situation of the tramp ant Wasmannia auropunctata in the Bahian cocoa-producing region where it is considered as native, and that of the two cryptobiotic genera Thaumatomyrmex and Typhlomyrmex, as well as that of proven and possible endangered army ant and Ponerini species. A total of 192 ant species from four strata were found in extensive sampling of a cocoa plantation with a relatively simple shade canopy (comprised primarily of Erythrina). Species richness in the cocoa plantations corresponded roughly to that of low diversity native forests, and species composition of cocoa plantations was most similar to native habitats (forest and mangroves) while ant composition in other agricultural habitats was most similar to that of urban areas. Although occurrences of Wasmannia auropunctata were similar in cocoa plantations and forests, abundance of Thaumatomyrmex and Typhlomyrmex, generally thought to be rare ants, was relatively high in cocoa plantations. These results, from cocoa plantations with relatively simple shade, demonstrate the importance of cocoa for ant conservation in the Atlantic forest region of Brazil. It is likely that cocoa plantations with a greater number of vegetation strata and higher tree species richness (such as traditional cabruca plantations) provide even more important habitat for ants generally and for ant species of conservation concern.     

Macrofauna assemblage composition and soil moisture interact to affect soil ecosystem functions

Changing climatic conditions and habitat fragmentation are predicted to alter the soil moisture conditions of temperate forests. It is not well understood how the soil macrofauna community will respond to changes in soil moisture, and how changes to species diversity and community composition may affect ecosystem functions, such as litter decomposition and soil fluxes. Moreover, few studies have considered the interactions between the abiotic and biotic factors that regulate soil processes. Here we attempt to disentangle the interactive effects of two of the main factors that regulate soil processes at small scales - moisture and macrofauna assemblage composition. The response of assemblages of three common temperate soil invertebrates (Glomeris marginata Villers, Porcellio scaber Latreille and Philoscia muscorum Scopoli) to two contrasting soil moisture levels was examined in a series of laboratory mesocosm experiments. The contribution of the invertebrates to the leaf litter mass loss of two common temperate tree species of contrasting litter quality (easily decomposing Fraxinus excelsior L. and recalcitrant Quercus robur L.) and to soil CO₂ fluxes were measured. Both moisture conditions and litter type influenced the functioning of the invertebrate assemblages, which was greater in high moisture conditions compared with low moisture conditions and on good quality vs. recalcitrant litter. In high moisture conditions, all macrofauna assemblages functioned at equal rates, whereas in low moisture conditions there were pronounced differences in litter mass loss among the assemblages. This indicates that species identity and assemblage composition are more important when moisture is limited. We suggest that complementarity between macrofauna species may mitigate the reduced functioning of some species, highlighting the importance of maintaining macrofauna species richness.     

云南橡胶林和天然次生林枯落物层蚂蚁物种多样性、群落结构差异及指示种

橡胶树Hevea brasiliensis是云南省重要的经济林木, 但对其生态服务功能尚存在争议。本研究以天然次生林为对照, 使用Winkler法对橡胶林枯落物层蚂蚁进行初步研究, 探讨橡胶林枯落物层蚂蚁的生态状况。于2012年10月和2013年4月采用Winkler袋法调查了云南省绿春县大黑山乡橡胶林和牛孔乡天然次生林枯落物层蚂蚁群落的物种多样性、 群落结构差异及指示种。结果表明： 橡胶林枯落物层蚂蚁多度（转换后）、 物种丰富度S和ACE值显著低于无干扰的天然次生林（P<0.05）; 蚂蚁多度（转换后）显著低于有干扰的天然次生林（P<0.05）, 而物种丰富度S和ACE值差异不显著。橡胶林枯落物层蚂蚁群落结构与两种天然次生林都不相似（F=3.93, df=12, P<0.01）。橡胶林中流浪种大头蚁属Pheidole的蚂蚁种类与天然次生林相比, 物种丰富度增加了100%。天然次生林枯落物层中蚂蚁指示种有3种, 分别为刘氏隆头蚁Strumigenys lewisi、 黄足厚结猛蚁Pachycondyla luteipes和女娲角腹蚁Recurvidris nuwa, 而橡胶林枯落物层中指示种仅为菱结大头蚁Pheidole nodus。枯落物层蚂蚁物种多样性与枯落物厚度呈显著正相关, 而枯落物盖度仅与蚂蚁多度（转换后）有相关性。结果说明, 橡胶林经过长期的经营管理, 生态环境趋于稳定, 对枯落物层蚂蚁群落具有一定的保护作用, 但与天然次生林相比, 蚂蚁多度（转换后）及群落结构仍显示出明显的不同。     

Oil palm expansion into rain forest greatly reduces ant biodiversity in canopy,epiphytes and leaf-litter

Oil palm cultivation is expanding rapidly into many of the world's most biodiverse tropical regions. One of the most functionally important and ecologically dominant animal groups in these environments is the ants. Here, we quantify the overall impacts of clear-felling lowland dipterocarp rainforest and conversion into oil palm plantation on ant diversity. At study sites in Sabah, Malaysia we collected ants from three microhabitats: 1 – the canopy, 2 – bird's nest ferns (Asplenium nidus complex, a common epiphyte in forest and oil palm), and 3 – leaf litter. We also measured temperature, humidity and light at collection sites to assess their impacts on ant community composition. Total ant species richness decreased from 309 to 110 (?64%) between forest and oil palm plantation. However, this impact was not the same across all microhabitats, with bird's nest ferns maintaining almost the same number of ant species in oil palm compared to forest (forest-oil palm, ferns: 36–35 (3% loss), canopy: 120–58 (52% loss), leaf litter: 216–56 (74% loss)). Relative abundance distributions remained the same for fern-dwelling ants, but became less even for oil palm ants in both the canopy and the leaf litter. These differences may be due in part to the ability of bird's nest ferns to provide a stable microclimate in hot, dry plantations. We also found that non-native ant species were more abundant in oil palm than in forest, and few forest ant species survived in plantations in any of the microhabitats. Only 59 of the 309 forest species persisted in oil palm plantations, corresponding to an 81% loss of forest species resulting from habitat conversion. Although oil palm supports many more ant species than has been previously reported, converting forest into plantation still leads to a dramatic reduction in species richness. The maintenance of forested areas is therefore vital for the conservation of ant biodiversity.     

Drought and presence of ants can influence hemiptera in tropical leaf litter

Climate change is predicted to impact tropical rain forests, with droughts becoming more frequent and more severe in some regions. We currently have a poor understanding of how increased drought will change the functioning of tropical rain forest. In particular, tropical rain forest invertebrates, which are numerous and biologically important, may respond to drought in different ways across trophic levels. Ants are a diverse group that carry out important ecosystem processes, shaping ecosystem structure and function through predation and competition, which can influence multiple trophic levels. Hemiptera are a mega-diverse order, abundant in tropical rain forests and are ecologically important. To understand the roles of ants in exerting predation and competition pressure on invertebrates in tropical rain forests during drought and a post-drought period, we established a large-scale ecosystem manipulation experiment in Maliau Basin Conservation Area in Malaysian Borneo, suppressing the activity of ants on four 0.25 ha plots over a two-year period. We sampled hemipterans found in the leaf litter during a drought (July 2015) and a post-drought period (September 2016) period. We found significant shifts in the assemblage of hemipterans sampled from the leaf litter following ant suppression. Specifically, for ant-suppression plots, the species richness and abundance of herbivorous hemipterans increased only during the post-drought period. For predatory hemipterans, abundance increased with ant-suppression regardless of drought conditions, and we found marginal evidence for a species richness increase during the post-drought period with little or no change in the drought period. These results illustrate how ants in tropical forests structure invertebrate communities and how these effects may vary with climatic variation.     

Ant assemblages on rehabilitated tropical landfills

Ant assemblages have been used widely throughout the world except in Asia to assess land restoration. In this study, ant assemblages were studied on three rehabilitated landfills (closed for 11–26 years) and a mature woodland as reference in Hong Kong. Grassland and woodland areas on each landfill were examined separately. Ants were sampled by soil extraction, litter extraction and pitfall trapping to maximize capture efficiency. Simple vegetation measurements were also taken. A total of 64 ant species from 33 genera were recorded. Ant species richness in the three landfills was similar (25–30 species), which did not correlate with the age of rehabilitation. A relationship between ant species richness and various vegetation parameters was not observed. Cluster analysis separated the landfill sites from the reference site, and grassland sites from woodland sites. Composition of ants based on functional grouping differed on the grassland and woodland sites, and functional group composition also separated the landfill sites from the reference site. Ant functional groups but not species richness showed a successional pattern with the ecological development of landfills. However, even after more than 25 years of rehabilitation, the ant assemblages on landfills were still very different from that on the reference site.     

Evaluating community effects of a Keystone Ant,Azteca sericeasur,on Inga micheliana leaf litter decomposition in a shaded coffee agro-ecosystem

Our research examined the effect of Azteca sericeasur, a keystone arboreal ant, on the decomposition of leaf litter of the shade tree, Inga micheliana, in coffee agro-ecosystems. This interaction is important in understanding spatial heterogeneity in decomposition. We hypothesized that A. sericeasur could affect leaf litter decomposition by excluding other ants, which could release decomposers, like collembolans, from predation pressure. Determining the relative strengths of these interactions can illuminate the importance of A. sericeasur in decomposition and nutrient cycling processes. We assessed the ant and arthropod communities surrounding 10 pairs of trees, where each pair included one shade tree with an established A. sericeasur nest. Tuna baits were used in conjunction with pitfall traps to assess the ant and arthropod community, and litterbags with I. micheliana leaf litter were used to assess rates of decomposition. The species richness of ants did not change in proximity to A. sericeasur nests, though the ant communities were distinct. Abundance of Collembola and community composition of other invertebrates did not change in the presence of A. sericeasur nests, and there were no differences in leaf litter decomposition rates. This contradicts past studies that suggest A. sericeasur reduces ant species richness in its territory. We suggest that other ants may avoid A. sericeasur by moving within and beneath the leaf litter. Our results indicate that there is no net effect of A. sericeasur on leaf litter decomposition.     

Can ants be used as indicators of environmental impacts caused by arsenic?

We evaluated ants as bioindicators of environmental impacts caused by arsenic residuals in the soil. We tested the hypotheses that the presence of arsenic in the soil affects: (1) estimates of resources and habitat condition for arboreal and epigaeic ants; (2) species richness of arboreal and epigaeic ants and (3) arboreal and epigaeic ant species composition. Ants were sampled at an inactivated raticide factory in Nova Lima, Minas Gerais, Brasil, which used arsenic as one of its main byproducts. The following environmental variables were measured: bioavailable arsenic concentration in the soil, the number and density of tree species, plant cover and leaf litter depth. The species richness of arboreal ants decreased with increased bioavailable arsenic concentration whilst epigaeic ants increased with arsenic. Arboreal ants were positively related to the number of tree species, which in turn were negatively affected by arsenic. We verified which ants are good bioindicators of arsenic. Independent verification of the influence of arsenic on background environmental variables was fundamental in defining responses of ant communities, as well as in identifying the most effective pathways for the recovery of biological communities in degraded areas.     

Changes in Species Richness,Abundance, and Composition of Arboreal Twig‐nesting Ants Along an Elevational Gradient in Coffee Landscapes

The distribution, diversity, and assembly of tropical insects have long intrigued ecologists, and for tropical ants, can be affected by competitive interactions, microhabitat requirements, dispersal, and availability and diversity of nesting sites. Arboreal twig‐nesting ants are limited by the number of hollow twigs available, especially in intensive agricultural systems. Ant diversity and abundance may shift along elevation gradients, but no studies have examined if the proportion of occupied twigs or richness of arboreal twig‐nesting ants vary with elevation. In coffee agroecosystems, there are over 40 species of arboreal twig‐nesting ants. We examined communities of twig‐nesting ants in coffee plants along an elevational gradient to answer the following questions: (1) Do species richness and colony abundance decline with elevation or show a mid‐elevation peak? (2) Does community composition change with elevation? (3) Is elevation an important predictor of change in ant abundance, richness, and relative abundance of common species? We surveyed 42 10 × 10 m plots in 2013 from 450 to1550 m elevation across a coffee landscape in Chiapas, Mexico. We sampled a total of 2211 hollow coffee twigs, 77.1 percent of which were occupied by one of 28 species of ants. Pseudomyrmex simplex was more abundant in lower elevations, whereas Pseudomyrmex ejectus dominated in high elevations. Species richness and the percent of occupied hollow twigs both peaked at mid‐elevations (800–1050 m). In sum, we found that species richness, abundance, and composition of arboreal twig‐nesting ants shift with elevation. These findings may provide important insights for understanding ant communities in coffee agroecosystems.     

Ants on the Move: Resource Limitation of a Litter‐nesting Ant Community in Costa Rica1

The leaf litter of tropical wet forests is replete with itinerant ant nests. Nest movement may help ants evade the constraints of stress and disturbance and increase access to resources. I studied how nest relocation and environmental factors may explain the density, size, and growth of leaf litter ant nests. I decoupled the relationships among litter depth, food abundance, and nest availability in a 4‐mo manipulation of food and leaf litter in a community of litter‐nesting ants in a lowland wet forest in Costa Rica. Over 4 mo, 290 1 m² treatment and control plots were sampled without replacement. Nest densities doubled in response to food supplementation, but did not decrease in response to litter removal or stress (from litter trampling). The supplementation of food increased the utilization of less favored nesting materials. In response to food supplementation and litter trampling, arboreal ants established nests in the litter, and growth rates of the most common ants (Pheidole spp.) increased. Colony growth was independent of colony size and growth rates of the most abundant ants. In general, I conclude that litter‐nesting ant density is driven primarily by food limitation, that nest relocation behavior significantly affects access to resource and the demographic structure of this community, and that nest fission may be a method to break the growth–reproduction trade‐off.     

Impact of Flooding on the Species Richness, Density and Composition of Amazonian Litter-Nesting Ants

Litter-nesting ants are diverse and abundant in tropical forests, but the factors structuring their communities are poorly known. Here we present results of the first study to examine the impact of natural variation in flooding on a highly diverse (21 genera, 77 species) litter-nesting ant community in a primary Amazonian forest. Fifty-six 3 × 3 m plots experiencing strong variation in flooding and twenty-eight 3 × 3 m terra firme plots were exhaustively searched for litter-nesting ants to determine patterns of density, species richness and species composition. In each plot, flooding, litter depth, twig availability, canopy cover, plant density, percent soil nitrogen, carbon, and phosphorus were measured. Degree of flooding, measured as flood frequency and flood interval, had the strongest impact on ant density in flooded forest. Flooding caused a linear decrease in ant abundance, potentially due to a reduction of suitable nesting sites. However, its influence on species richness varied: low-disturbance habitat had species richness equal to terra firme forest after adjusting for differences in density. The composition of ant genera and species varied among flood categories; some groups known to contain specialist predators were particularly intolerant to flooding. Hypoponera STD10 appeared to be well-adapted to highly flooded habitat. Although flooding did not appear to increase species richness or abundance at the habitat scale, low-flooding habitat contained a mixture of species found in the significantly distinct ant communities of terra firme and highly flooded habitat.
     

Size matters: nest colonization patterns for twig‐nesting ants

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Wood-nesting ants and their parasites in forests and coffee agroecosystems

Agricultural intensification is linked to reduced species richness and may limit the effectiveness of predators in agricultural systems. We studied the abundance, diversity, and species composition of wood-nesting ants and frequency of parasitism of poneromorph ants in coffee agroeco systems and a forest fragment in Chiapas, Mexico. In three farms differing in shade management and in a nearby forest fragment, we surveyed ants nesting in rotten wood. We collected pupae of all poneromorph ants encountered, and incubated pupae for 15 d to recover emerging ant parasites. If no parasites emerged, we dissected pupae to examine for parasitism. Overall, we found 63 ant morphospecies, 29 genera, and 7 subfamilies from 520 colonies. There were no significant differences in ant richness or abundance between the different sites. However, there were significant differences in the species composition of ants sampled in the four different sites. The parasitism rates of ants differed according to site; in the forest 77.7% of species were parasitized, and this number declined with increasing intensification in traditional polyculture (40%),commercial polyculture (25%), and shade monoculture (16.6%). For three of four poneromorph species found in >1 habitat, parasitism rates were higher in the more vegetatively complex sites. The result that both ant species composition and ant parasitism differed among by site indicates that coffee management intensification affects wood-nesting ant communities. Further, coffee intensification may significantly alter interactions between ants and their parasites, with possible implications for biological control in coffee agroecosystems.     

Limitation of nesting resources for ants in Colombian forests and coffee plantations

Abstract.  1. This study examines limitation of nesting resources for leaf-litter and twig-nesting ants as a mechanism of diversity loss across an intensification gradient of coffee production in Colombia. Twelve farms were selected and classified into four management types: forest, polygeneric shade coffee, monogeneric shade coffee, and sun coffee (unshaded coffee monocultures).
2. At each of the farms, four treatment subplots were established at the corners of each of 10 25 m² plots: (i) twig augmentation (adding 10 empty bamboo twigs); (ii) litter augmentation (tripling existing litter profile); (iii) twig and litter augmentation; and (iv) no manipulation control, for a total of 480 subplots. A twig addition experiment was also performed on coffee bushes.
3. The results showed significantly more ant colonies in the forest and monogeneric shade coffee litter augmentation plots after 4 months. Litter-nesting ant species richness was higher in all three shade systems than in the sun coffee. The identities of ants nesting on coffee bushes were different from those in the soil level litter. Fewer species nested in bamboo twigs placed in litter in the most intensive systems.
4. More ants nested in the resource addition treatments, and more ant species were found in forested habitats; however, a single mechanism cannot explain the observed patterns. It was concluded that a combination of bottom-up and top-down effects might lead to the loss of associated fauna with the intensification of these agroecosystems.     

The influence of nest availability on the abundance and diversity of twig-dwelling ants in a Papua New Guinea forest

Tropical ant communities are frequently diverse, but highly patchy in nature. The availability of suitable nest sites may be a regulating force in structuring litter ant communities. Our aim was to examine ant resource utilization in naturally occurring twigs, and to modify the availability of these resources in order to quantify the influence of nest availability on ant communities in a Papua New Guinean forest. First, we compared ant communities that assemble in artificial twigs (drilled, wooden dowels), naturally occurring twigs, and the leaf litter. A total of 55 ant species were captured: 33 from the leaf litter, 29 from naturally occurring twigs, and only 12 from artificial nests. Significantly different communities formed in each of the three nest types. Second, we examined how the density of natural or artificial nest material influenced the ant abundance and species richness. Plots had between 5 and 96 potential nest sites. An average of only 11.2% of these twigs was colonized. Both species richness and the total abundance of adult ants were significantly positively correlated with increasing naturally occurring twig density. Conversely, increasing the availability of artificial nests from 5 to 20 per plot had no significant effect on the proportion of artificial nests colonized, species richness, or the colony size. We observed that ant species richness and abundance increased with natural twig density, at least for naturally occurring communities. But why so many twigs remain vacant and available for ant colonization remains unknown. Other biotic and abiotic factors likely influence the use of nesting habitat in these ant communities.     

Effect of an exotic Acacia (Fabaceae) on ant assemblages in South African fynbos

Ant assemblages in South African fynbos invaded by Acacia saligna were compared with ant assemblages in undisturbed fynbos to determine whether ant assemblages change under exotic plants that produce ant‐dispersed seeds. Overall, no differences in the species richness of ants were found between weed‐infested and native sites but there were differences in both ant abundance and the composition of the ant assemblage. Ants were much less abundant in weed‐infested sites. To investigate whether changes in ant assemblages in weed‐infested areas could be due to a preference for native seeds over exotic seeds, seeds of a range of species were offered to ants and ants that handled seeds were identified. Thirteen species of ants handled A. saligna seeds and there was no evidence to suggest that the ant assemblage as a whole preferred native seeds to A. saligna seeds. Hypotheses that may account for this pattern are discussed.     

Impacts of Argentine ants on invertebrate communities with below-ground consequences

The Argentine ant, Linepithema humile (Mayr), is an invasive species that has been associated with various negative impacts in native communities around the world. These impacts, as for other invasive ants, are principally towards native ant species, and impacts on below-ground processes such as decomposition remain largely unexplored. We investigated the relationship between Argentine ants and invertebrate fauna, litter decomposition and soil microbial activity between paired invaded and uninvaded sites at two locations in Auckland, New Zealand, where there has been no research to date on their impacts. We examined the diversity and composition of invertebrate and microorganisms communities, and differences in soil and litter components. The composition of invertebrates (Order-level, ant and beetle species) was different between invaded and uninvaded sites, with fewer ants, isopods, amphipods, and fungus-feeding beetles at the invaded sites, whereas Collembola were more abundant at the invaded sites. There were significant differences in soil chemistry, including higher carbon and nitrogen microbial biomass at uninvaded sites. Several litter components were significantly different for Macropiper excelsum. The fibre content of litter was higher, and key nutrients (e.g. nitrogen) were lower, at invaded sites, indicating less breakdown of litter at invaded sites. A greater knowledge of the history of invasion at a site would clarify variation in the impacts of Argentine ants, but their persistence in the ground litter layer may have long-term implications for soil and plant health in native ecosystems.     

Effects of experimental rainfall manipulations on Chihuahuan Desert grassland and shrubland plant communities

Aridland ecosystems are predicted to be responsive to both increases and decreases in precipitation. In addition, chronic droughts may contribute to encroachment of native C₃ shrubs into C₄-dominated grasslands. We conducted a long-term rainfall manipulation experiment in native grassland, shrubland and the grass–shrub ecotone in the northern Chihuahuan Desert, USA. We evaluated the effects of 5 years of experimental drought and 4 years of water addition on plant community structure and dynamics. We assessed the effects of altered rainfall regimes on the abundance of dominant species as well as on species richness and subdominant grasses, forbs and shrubs. Nonmetric multidimensional scaling and MANOVA were used to quantify changes in species composition in response to chronic addition or reduction of rainfall. We found that drought consistently and strongly decreased cover of Bouteloua eriopoda, the dominant C₄ grass in this system, whereas water addition slightly increased cover, with little variation between years. In contrast, neither chronic drought nor increased rainfall had consistent effects on the cover of Larrea tridentata, the dominant C₃ shrub. Species richness declined in shrub-dominated vegetation in response to drought whereas richness increased or was unaffected by water addition or drought in mixed- and grass-dominated vegetation. Cover of subdominant shrubs, grasses and forbs changed significantly over time, primarily in response to interannual rainfall variability more so than to our experimental rainfall treatments. Nevertheless, drought and water addition shifted the species composition of plant communities in all three vegetation types. Overall, we found that B. eriopoda responded strongly to drought and less so to irrigation, whereas L. tridentata showed limited response to either treatment. The strong decline in grass cover and the resistance of shrub cover to rainfall reduction suggest that chronic drought may be a key factor promoting shrub dominance during encroachment into desert grassland.