Diversity and species composition of West African ungulate assemblages: effects of fire,climate and soil

Aim Anthropogenic fires are a major component of the ecology of rangelands throughout the world. To assess the effects of these fires on the diversity patterns of herbivores, we related gradients in fire occurrence, climate and soil fertility to patterns in alpha and beta diversity of African ungulates. Location West Africa. Methods We used a survey‐based approach for ungulates in 37 protected areas in desert, savanna and rain forest habitats throughout West Africa, combined with satellite images of fire occurrence and digital maps of actual evapotranspiration and soil fertility. Alpha diversity was related to the environmental variables using conventional and spatial regression models. We investigated beta diversity using partial Mantel tests and ordination techniques, and by partitioning the variance in assemblage composition into environmental and spatial components. Results The species richness of grazers showed a quadratic relationship with actual evapotranspiration, whereas that of browsers and frugivores showed a linear relationship. However, in the multiple regression models fire occurrence was the only variable that significantly correlated with the species richness of grazers. Soil fertility was weakly related to overall beta diversity and the species richness of browsers, but was non‐significant in the multiple regression models. Fire occurrence was the most important variable explaining species composition of the overall species set and of grazers, whereas the assemblage composition of browsers and frugivores was explained mostly by actual evapotranspiration. Main conclusions In contrast to previous studies, our analyses show that moisture and nutrients alone fail to adequately predict the diversity patterns of grazing ungulates. Rather, the species richness and assemblage composition of grazers are largely governed by anthropogenic fires that modify the quality and structure of the grass sward. Diversity patterns of browsers and frugivores are markedly different from grazers and depend mainly on the availability of moisture, which is positively correlated with the availability of foliage and fruits. Our study highlights the importance of incorporating major human‐induced disturbances or habitat alterations into analyses of diversity patterns.     

Increased temperature alters feeding behavior of a generalist herbivore

Temperature can regulate a number of important biological processes and species interactions. For example, environmental temperature can alter insect herbivore consumption, growth and survivorship, suggesting that temperature‐driven impacts on herbivory could influence plant community composition or nutrient cycling. However, few studies to date have examined whether rising temperature influences herbivore preference and performance among multiple plant species, which often dictates their impact at the community level. Here, we assessed the effects of temperature on the performance and preference of the generalist herbivore Popillia japonica among nine plant species. We show that, on average, consumption rates and herbivore performance increased at higher temperatures. However, there was considerable variation among plant species with consumption and performance increasing on some plant species at higher temperatures but decreasing on others. Plant nutritional quality appeared to influence these patterns as beetles increased feeding on high‐nitrogen plants with increasing temperature, suggesting stronger nitrogen limitation. In addition to changes in feeding rates, feeding preferences of P. japonica shifted among temperatures, a pattern that was largely explained by differential deterrence of plant chemical extracts at different temperatures. In fact, temperature‐induced changes in the efficacy of plant chemical extracts led P. japonica to reduce its diet breadth at higher temperatures. Our results indicate that rising temperatures will influence herbivore feeding behavior by altering the importance of plant nutritional and chemical traits, suggesting that climate change will alter the strength and sign of plant–insect interactions.     

Short-term soil inorganic N pulse after experimental fire alters invasive and native annual plant production in a Mojave Desert shrubland

Post-fire changes in desert vegetation patterns are known, but the mechanisms are poorly understood. Theory suggests that pulse dynamics of resource availability confer advantages to invasive annual species, and that pulse timing can influence survival and competition among species. Precipitation patterns in the American Southwest are predicted to shift toward a drier climate, potentially altering post-fire resource availability and consequent vegetation dynamics. We quantified post-fire inorganic N dynamics and determined how annual plants respond to soil inorganic nitrogen variability following experimental fires in a Mojave Desert shrub community. Soil inorganic N, soil net N mineralization, and production of annual plants were measured beneath shrubs and in interspaces during 6 months following fire. Soil inorganic N pools in burned plots were up to 1 g m⁻² greater than unburned plots for several weeks and increased under shrubs (0.5–1.0 g m⁻²) more than interspaces (0.1–0.2 g m⁻²). Soil NO₃ ⁻−N (nitrate−N) increased more and persisted longer than soil NH₄ ⁺−N (ammonium−N). Laboratory incubations simulating low soil moisture conditions, and consistent with field moisture during the study, suggest that soil net ammonification and net nitrification were low and mostly unaffected by shrub canopy or burning. After late season rains, and where soil inorganic N pools were elevated after fire, productivity of the predominant invasive Schismus spp. increased and native annuals declined. Results suggest that increased N availability following wildfire can favor invasive annuals over natives. Whether the short-term success of invasive species following fire will direct long-term species composition changes remains to be seen, yet predicted changes in precipitation variability will likely interact with N cycling to affect invasive annual plant dominance following wildfire.     

Prairie vegetation and soil nutrient responses to ungulate carcasses

The impact of large ungulate carcasses on grassland dynamics was investigated by monitoring vegetation and soil nutrients in 50-cm circular zones around the center of bison (Bos bison), cattle (B. taurus), and deer (Odocoileus virginianus) carcasses. An ungulate carcass creates an intense localized disturbance that varies with animal size and the season of death. Unlike other natural disturbances, carcasses deposit a concentrated pulse of nutrients into the soil. One year after death, inorganic nitrogen concentrations were significantly higher in the inner 50 cm at both adult and juvenile carcass sites than in surrounding prairie. Areas around a carcass became zones of fertility that favored different components of the vegetation and stimulated biomass production. Species richness and diversity at the center of carcass sites were lowest 1 year after death, but increased significantly in subsequent years. However, warm-season perennial grasses declined near the center of carcass sites and did not recover. Five years after death, ungulate carcass sites remained disturbed patches that harbored vegetation characteristically different in composition and stature from surrounding prairie. By providing a niche for species not normally found in undisturbed prairie, carcasses increased community heterogeneity and may play an important role in adding spatial complexity to grassland ecosytems. Received: 12 April 1999 / Accepted: 7 September 1999     

The jasmonate pathway alters herbivore feeding behaviour: consequences for plant defences

The jasmonate pathway is a highly conserved defensive cascade in plants that regulates the induction of resistance against herbivores; however, its role in herbivore feeding behaviour remains unknown. We used a mutant tomato plant (def‐1) deficient in the production of jasmonate‐related defensive proteins to test the hypothesis that genotypes with a reduced ability to induce resistance have a higher and more concentrated pattern of herbivore damage. Wild‐type and def‐1 plants received either damage by Spodoptera exigua (Hübner) (Lepidoptera: Noctuidae) caterpillars or no damage. After treatment, we tested for systemic responses by allowing a free roaming S. exigua caterpillar to feed on the undamaged portions of plants. Weight‐gain and leaf consumption of S. exigua were highest on def‐1 plants, regardless of prior herbivore damage. Def‐1 plants also had fewer numbers of leaves and leaflets eaten, and fewer feeding holes, which was indicative of a more concentrated distribution of damage on mutant compared to wild‐type plants. Following these results, we mimicked the amount and distribution of feeding damage that wild‐type or jasmonate‐deficient plants would receive on wild‐type plants to test whether changes in feeding behaviour may feedback to influence the expression of induced resistance. We mimicked the distribution of damage in wild‐type and jasmonate‐deficient plants by allowing caterpillars to feed on either one (leaf 1 or 2) or two leaves (leaf 1 and 2). Increased herbivore damage resulted in higher proteinase inhibitor (PI) activity, a jasmonate‐regulated defensive protein, and lower S. exigua performance on wild‐type but not jasmonate‐deficient plants. Compared to undamaged plants, a concentrated pattern of herbivore damage increased systemic resistance; these induced responses were greater on leaflets with stronger vascular connections to the damaged leaf. A more dispersed pattern of caterpillar damage altered the expression of induced responses, but the outcome depended on the specific pattern of damage. When leaf 1 was damaged and then leaf 2, the undamaged (third) leaf (which is more strongly connected to leaf 1 than 2) expressed reduced the PI activity compared to plants receiving concentrated damage to leaf 1; whereas in plants where leaf 2 was first damaged and then leaf 1, there were no differences in PI activity in leaf 3 compared to plants receiving concentrated damage to leaf 2. Thus, induction of the jasmonate pathway may not only determine the amount and distribution of feeding damage by herbivores, but this may feedback to affect the subsequent expression of plant defence.     

Native herbivore exerts contrasting effects on fire regime and vegetation structure

Although native herbivores can alter fire regimes by consuming herbaceous vegetation that serves as fine fuel and, less commonly, accumulating fuel as nest material and other structures, simultaneous considerations of contrasting effects of herbivores on fire have scarcely been addressed. We proposed that a colonial rodent, vizcacha (Lagostomus maximus), reduces and increases fire intensity at different stages in its population cycle in the semiarid scrub of Argentina. Specifically, we hypothesized that, when colonies are active, vizcachas create natural fire-breaks through intense grazing, generating over time patches of large unburned shrubs in grazed zones. In contrast, when colonies are abandoned, recovery of fine fuels and previous accumulation of coarse wood on colonies during territorial displays increases fire intensity, creating patches of high shrub mortality. To test these hypotheses, we estimated stem age of the dominant shrub (Larrea divaricata) and measured aboveground biomass in zones actively grazed by vizcachas and in ungrazed zones, and compared densities of live and dead shrubs on abandoned colonies and adjacent zones following fire. In active colonies, age and biomass of shrubs were much greater in grazed than ungrazed zones. In abandoned colonies that had been burnt, density of dead, burned shrubs was higher and density of live shrubs was lower than in adjacent zones. These results support our hypotheses and reveal a new interaction between native herbivores and fire, in which herbivores augment fire intensity by gathering fuel. Our findings indicate that, through opposing effects on fire, native herbivores enhance the heterogeneity of vegetation in woody-dominated ecosystems.     

Plant species influence on soil microbial short-term response after fire simulation

Aims

Plant species can influence fire intensity and severity causing different immediate and long-term responses on the soil microbial community. The main objective of this work was to determine the role of two representative Mediterranean plant species as soil organic matter sources, and to identify their influence on microbial response before and after heat exposure.

Methods

A laboratory heating experiment (300 °C for 20 min) was performed using soil collected under Pinus hallepensis (PIN) and Quercus coccifera (KER). Dried plant material was added before heating for a total of six different treatments: non-heated control samples amended with the original plant material (PIN₀ and KER₀); PIN samples heated with pine (PIN_p) or kermes oak litter (PIN_k); KER samples heated with kermes oak (KER_k) or pine litter (KER_p). Heated soils were inoculated with the original fresh soil and different microbial parameters related to abundance, activity and possible changes in microbial community composition and chemical soil parameters that could be conditioning microbial response were measured for 28 days after inoculation.

Results

The effect of heating on the soil microbial parameters studied was influenced to a small extent by the plant species providing fuel, being evident in soil samples taken under pine influence. Nevertheless heating effect showed marked differences when plant species influence on soil origin was analyzed.

Conclusions

In general, samples taken under pine appear to be more negatively affected by heating treatment than samples collected under kermes oak, highlighting the importance of vegetation as a fresh organic matter source in soil ecosystems before and after fire.     

Long-term responses in arctic ungulate dynamics to changes in climatic and trophic processes

 Following predictions from climatic general circulation models, the effects of perturbations in global climate are expected to be most pronounced in the Northern Hemisphere. Elaborating on a recently developed plant–herbivore–climate model, we explore statistically how different winter climate regimes and density-dependent processes during the past century have affected population dynamics of two arctic ungulate species. Our analyses were performed on the dynamics of six muskox and six caribou populations. In muskoxen, variation in winter climate, mediated through the North Atlantic Oscillation (NAO), explained up to 24% of the variation in interannual abundance, whereas in caribou up to 16% was explained by the NAO. Muskoxen responded negatively following warm and snowy winters, whereas caribou responded negatively to dry winters. Direct and delayed density dependence was recorded in most populations and explained up to 32% and 90% of variations in abundance of muskoxen and caribou, respectively. Received: November 19, 2001 / Accepted: May 28, 2002     

Introduced fire ants can exclude native ants from critical mutualist-provided resources

After over 30 years of research, it was recently shown that nectar amino acids increase female butterfly fecundity. However, little attention has been paid to the effect of nectar amino acids on male butterfly reproduction. Here, we show that larval food conditions (nitrogen-rich vs. nitrogen-poor host plants) and adult diet quality (nectar with or without amino acids) affected the amount of consumed nectar in Coenonympha pamphilus males. Furthermore, amino acids in the nectar diet of males increased progeny’s larval hatching mass, irrespective of paternal larval reserves. Our study takes the whole reproductive cycle of male butterflies into account, and also considers the role of females in passing male nutrients to offspring, as males’ realized reproduction was examined indirectly via nuptial gifts, by female performance. With this comprehensive approach, we demonstrate for the first time that nectar amino acids can improve male butterfly reproduction, supporting the old postulate that nectar amino acids generally enhance butterfly fitness.     

Simulated climate warming alters phenological synchrony between an outbreak insect herbivore and host trees

As the world’s climate warms, the phenologies of interacting organisms in seasonally cold environments may advance at differing rates, leading to alterations in phenological synchrony that can have important ecological consequences. For temperate and boreal species, the timing of early spring development plays a key role in plant–herbivore interactions and can influence insect performance, outbreak dynamics, and plant damage. We used a field-based, meso-scale free-air forest warming experiment (B4WarmED) to examine the effects of elevated temperature on the phenology and performance of forest tent caterpillar (Malacosoma disstria) in relation to the phenology of two host trees, aspen (Populus tremuloides) and birch (Betula papyrifera). Results of our 2-year study demonstrated that spring phenology advanced for both insects and trees, with experimentally manipulated increases in temperature of 1.7 and 3.4 °C. However, tree phenology advanced more than insect phenology, resulting in altered phenological synchrony. Specifically, we observed a decrease in the time interval between herbivore egg hatch and budbreak of aspen in both years and birch in one year. Moreover, warming decreased larval development time from egg hatch to pupation, but did not affect pupal mass. Larvae developed more quickly on aspen than birch, but pupal mass was not affected by host species. Our study reveals that warming-induced phenological shifts can alter the timing of ecological interactions across trophic levels. These findings illustrate one mechanism by which climate warming could mediate insect herbivore outbreaks, and also highlights the importance of climate change effects on trophic interactions.     

Stoichiometric tracking of soil nutrients by a desert insect herbivore

Abstract Biogeochemistry and population biology have developed independently, with few attempts at linkage, almost none of which were mechanistically based. We hypothesize that biogeochemical cycling is linked to herbivore population dynamics through the influence of soil nutrient availability on foliar nutrient content, which constrains herbivore investment in phosphorus (P)‐rich molecules necessary for growth. We show that variation in desert soil P availability is linked to abundance of an insect herbivore (Sabinia setosa) through the influence of soil P on the C:P ratio of the host plant (Prosopis velutina). Low P availability increases C:P ratio of Prosopis leaves, leading to a decline in body %P, %RNA and abundance of Sabinia. Tight association between soil, plant and herbivore P provides the first evidence of a mechanistic pathway linking P biogeochemistry to terrestrial food webs by altering the supply of dietary P to herbivores, thus limiting their capacity for growth by constraining the production of P‐rich cellular ribosomal RNA (rRNA).     

Jasmonic acid signalling and herbivore resistance traits constrain regrowth after herbivore attack in Nicotiana attenuata

Because traits conferring resistance on herbivores can reduce fitness-associated traits, trade-offs may occur between tolerance and resistance responses. We examined these trade-offs in genotypes of Nicotiana attenuata that were transformed to silence trypsin proteinase inhibitor (TPI) production (AS-Natpi), an antiherbivore defense associated with (14%) reductions in seed production, and the jasmonate signal cascade that elicits these defenses (AS-Nalox3), by measuring stalk and axillary branch growth and seed production after two defoliation regimes and Manduca sexta larval attack to bottom or middle and top stalk leaves. Larval attack and defoliation at middle and top leaves depressed seed production and increased axillary branching more than at bottom leaves. AS-Nalox3 and AS-Natpi plants produced significantly longer (two- to fourfold) branches than did wild-type (WT) plants, results that are consistent with resource-based trade-offs between resistance and regrowth. Methyl jasmonate (MeJA) treatment of AS-Nalox3 plants restored WT branch growth, suggesting that jasmonic acid (JA) signalling suppresses regrowth and contributes to apical dominance. These results are consistent with the existence of JA- and resource-mediated trade-offs between regrowth and herbivore resistance traits.     

Topographic and ungulate regulation of soil C turnover in a temperate grassland ecosystem

A major obstacle for predicting the effects of climate and land use changes on global soil carbon (C) stores is the very limited knowledge about the long timescale dynamics of the relatively stable fraction of soil C, which represents the bulk of soil C and the primary determinant of the long‐term C balance of terrestrial ecosystems. In this study, we examined how variable topo‐edaphic conditions and herds of native migratory ungulates influenced turnover of the stable pool (total minus active fraction) of soil C in grasslands of Yellowstone National Park (YNP). Soil C properties were determined for grasslands located inside and outside long‐term ungulate exclosures established 1958–1962 at seven variable topographic positions. Active C pool sizes, estimated with soil laboratory incubations, and soil radiocarbon measures were used to parameterize a process‐based model to determine turnover of the stable C pool at the sites. Stable C turnover ranged 37–653 and 89–869 years for 0–10 and 0–20 cm soils, respectively. Among ungrazed communities, there was a trend for stable soil C turnover to slow along topographic gradients of increasing soil moisture, soil C content, and shoot biomass from hilltop to slope‐bottom positions. This was likely a result of an increasing amount of support tissue resulting in greater concentrations of lignin and cellulose as shoot biomass increased down slope. In contrast, across the grazed landscape, stable C turnover sped up from hilltop to slope‐bottom positions, which was likely a consequence of grazer effects on plant species composition along the topographic gradient. These findings indicated that despite topography playing the primary role in controlling such important site characteristics as soil moisture, soil C content, and plant production in YNP grassland, the long‐term turnover of the stable C pool was determined by herbivores. The results demonstrate the important regulatory role of herbivores in controlling the C balance of this semiarid grassland ecosystem.     

Recycling of nitrogen in herbivore feces: plant recovery,herbivore assimilation,soil retention,and leaching losses

Herbivores directly and indirectly affect ecosystem functioning in forests. Feces deposition is a direct effect that supplies ephemeral N pulses to soils. Herbivore-mediated changes in plant N allocation and uptake are indirect effects that can also influence soil N availability. These effects may interact if defoliation influences the ability of plants to recover fecal N, and this may affect subsequent generations of herbivores. We added ¹⁵N-enriched insect feces (frass) to a series of replicated red oak, Quercus rubra, mesocosms that had been damaged experimentally and then followed the frass N over the course of 2 years. In the first season, some frass N was mineralized in the soil and leached in organic form from the mesocosms within 1 week of deposition. Within 1 month, frass N had been acquired by the oaks and enriched the foliage; late-season herbivores assimilated the frass N within the same growing season. In the second season, herbivore damage from the previous year lowered total leaf N contents and ¹⁵N recovered in the foliage. A subsequent cohort of early-season herbivores fed on this foliage consequently derived less of their N from the previous year’s frass, and feral leaf rollers colonized fewer of these saplings. The 0- to 5-cm soil fraction was the largest N sink measured, and 42% of the frass N was recovered in the soil. The results demonstrate that: (1) some frass N can be recycled rapidly into foliage and assimilated by successive cohorts of herbivore within the same season; (2) damage can affect N allocation in the following year’s foliage, influencing N availability to and host selection by herbivores; and (3) leaching losses occur soon after deposition but are buffered by soil pools, which are the largest sinks for frass N.     

Succession processes in a food web of a two autotroph--one herbivore system

This paper deals with the succession process of a food web model consisting of one herbivore, two autotrophs and available nutrient in the environment in a closed nutrient flux. The model provides a way of describing successional changes in the form of species replacement with increasing nutrient levels. It is shown that distinct threshold (with upper and lower) values of nutrient are required for progression of succession process.     

Afforestation alters community structure of soil fungi

Relatively little is known about the effect of afforestation on soil fungal communities. This study demonstrated that afforestation altered fungal community structure and that changes were correlated to pools of soil C. Pasture at three locations on the same soil type was afforested with Eucalyptus globulus or Pinus pinaster. The structure of fungal communities under the three land uses was measured after 13y using automated ribosomal intergenic spacer analysis (ARISA). Afforestation significantly altered the structure of fungal communities. The effect of location on the structure of fungal communities was limited to pasture soils; although these contained the same plant species, the relative composition of each species varied between locations. Differences in the structure of fungal communities between pasture, E. globulus and P. pinaster were significantly correlated with changes in the amount of total organic C and microbial biomass-C in soil. Afforestation of patches of agricultural land may contribute to conserving soil fungi in agricultural landscapes by supporting fungal communities with different composition to agricultural soils.