Impacts of seedling herbivory on plant competition and implications for species coexistence

Background and Aims

Although the causes and consequences of seedling herbivory for plant community composition are well understood, the mechanisms by which herbivores influence plant species recruitment to the established phase remain less clear. The aim was to examine how variation in the intensity of seedling herbivory interacts with growth-defence trade-offs and herbivore feeding preferences to affect plant community development.

Methods

Using 14-d-old seedlings of Trifolium pratense and T. repens, relative growth and susceptibility to herbivory by the snail Helix aspersa was quantified to elucidate putative growth-defence trade-offs for these species. Then mixed assemblages of 14-d-old Trifolium seedlings were exposed to herbivory by zero, two, five or ten snails and determined how variation in the intensity of herbivory affected competitive interactions into the mature phase (as measured by total plant biomass at 120 d old).

Key Results

In the absence of herbivory, communities were dominated by T. pratense; a result expected on the basis that it yielded larger and presumably more competitive seedlings. However, when seedlings were exposed to herbivory, the balance of competition shifted. At low levels of herbivory (two snails), both Trifolium species contributed equally to total plant biomass. More intense herbivory (five snails) resulted in almost total mortality of T. pratense and dominance of the mature community by T. repens. The most intense herbivory (ten snails) effectively removed all seedlings from the experimental community.

Conclusions

The study illustrates a mechanism whereby spatio-temporal fluctuations in seedling herbivory, when coupled with species-specific variation in competitive ability and sensitivity to herbivore attack, can differentially influence plant recruitment into the mature phase. This mechanism may be a key element in our attempts to understand plant species coexistence, since fluctuations in plant recruitment are fundamental to the many theories that view coexistence as a consequence of a spatio-temporal lottery for dominance over regeneration micro-sites.Key words: Growth-defence trade-off, lottery models, plant–animal interactions, plant size variability, seedling acceptability, seedling defence, spatio-temporal niches, Trifolium pratense, Trifolium repens     

Hydraulic lift among native plant species in the Mojave Desert

Hydraulic lift was investigated among native plants in the Mojave Desert using in situ thermocouple psychrometers. Night lighting and day shading experiments were used to verify the phenomenon. Hydraulic lift was detected for all species examined: five shrub species with different rooting depths and leaf phenologies and one perennial grass species. This study was the first to document hydraulic lift for a CAM species, Yucca schidigera. The pattern of diel flux in soil water potential for the CAM species was temporally opposite to that of C₃ species: for the CAM plant, soil water potential increased in shallow soils during the day when the plant was not transpiring and decreased at night when transpiration began. Because CAM plants transport water to shallow soils during the day when surrounding C₃ and C₄ plants transpire, CAM species that hydraulically lift water may influence water relations of surrounding species to a greater extent than hydraulically lifting C₃ or C₄ species. A strong, negative relationship between the percent sand in the study site soils at the 0.35 m soil depth and the frequency that hydraulic lift was observed at that depth suggests that the occurrence of hydraulic lift is negatively influenced by coarse-textured soils, perhaps due to less root–soil contact in sandy soils relative to finer-textured soils. Differences in soil texture among study sites may explain, in part, differences in the frequency that hydraulic lift was detected among these species. Further investigations are needed to elucidate species versus soil texture effects on hydraulic lift. This revised version was published online in June 2006 with corrections to the Cover Date.     

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.     

Interspecific competition: A mechanism for rodent succession after fire in wet heathland

Abstract A competitive interaction between two species of sympatric native rodents was demonstrated experimentally in Myall Lakes National Park (NSW, Australia). The previously documented replacement of Pseudomys gracilicaudatus by Rattus lutreolus in areas of wet heath in this region implied that competition may be the mechanism facilitating this succession. We present experimental evidence to support this suggestion. Removal of the larger R. lutreolus caused a significant increase in abundance of P. gracilicaudatus on five experimental sites in comparison to five unmanipulated control sites. The sites used in this experiment were chosen from three ages of wet heath regenerating after fire. These three ages represent key stages in the replacement of P. gracilicaudatus by R. lutreolus and the occurrence of competition in each of these ages indicates that competition plays an important role in this succession. In the initial stages of the experiment when R. lutreolus were removed they were replaced by new R. lutreolus individuals. This has been interpreted as evidence of strong intraspecific competition. After removal of R. lutreolus, P. gracilicaudatus expanded its habitat range into microhabitats that had formerly been occupied by R. lutreolus and reduced its range in microhabitats previously occupied by P. gracilicaudatus. This leads us to believe that P. gracilicaudatus was occupying inferior microhabitat before the removal of R. lutreolus, which had excluded it from more preferable microhabitat.     

Effects of herbivory and competition on an introduced plant in decline

Anthemiscotula was introduced to Denmark 500 years ago, and its distribution is presently limited and in decline. A manipulative field experiment was performed to investigate the effects of native plant competitors and native invertebrate herbivores on its performance. Generally, both herbivory and competition treatments had great impact, and when both factors were operating, the effects were additive for all variables except plant height. Although A. cotula showed plasticity in growth, resource allocation and flowering timing, it was unable to adjust to competition and compensate for losses due to herbivory sufficiently to ensure and restore its achene production. This vulnerability, combined with improved cereal cleaning techniques and thus fewer reintroductions of A. cotula seeds, may be the cause of its current decline. A. cotula responded to herbivory by prolonging its flowering period, a “bet-hedging” strategy. In Denmark this strategy is unreliable since risks of sub-optimal conditions are much greater in August–October. Received: 12 December 1997 / Accepted: 14 October 1998     

Changing dominance of key plant species across a Mediterranean climate region: implications for fuel types and future fire regimes

Herbaceous and woody plants represent different fuel types in flammable ecosystems, due to contrasting patterns of growth and flammability in response to productivity (moisture availability). However, other factors, such as soil type, fire regimes and competitive interactions may also influence the relative composition of herbaceous and woody plants within a community. The Mediterranean climate region of south eastern Australia is transitional between two contrasting fuel systems; herbaceous dominated in the dry north, versus woody plant dominated shrublands in the relatively moist south. Across the rainfall gradient of the region, there are confounded changes in dominant soil types and fire frequency. We used model-subset selection using Akaike’s Information Criterion to examine potential driving mechanisms of community compositional change from herbaceous (e.g. Triodia scariosa, Austrostipa sp.) to woody plants (e.g. Beyeria opaca, Leptospermum coriaceum, Acacia ligulata) by measuring relative cover across combinations of rainfall, time since the last fire (TSF) and soil type. We examined the relative influence of environmental versus competitive interactions on determining the cover of perennial hummock grass, T. scariosa, and co-occurring woody shrubs. Rainfall and soil types, rather than competition, were the over-arching determinants of the relative cover of grasses and shrubs. Given the sensitivity to rainfall, our results indicate there is strong potential for the nature of fuel, flammability and fire regimes to be altered in the future via climate change in this region.     

Plant diversity,herbivory and resistance of a plant community to invasion in Mediterranean annual communities

Several components of the diversity of plant communities, such as species richness, species composition, number of functional groups and functional composition, have been shown to directly affect the performance of exotic species. Exotics can also be affected by herbivores of the native plant community. However, these two possible mechanisms limiting invasion have never been investigated together. The aim of this study was to investigate the relationships between plant diversity, herbivory and performance of two annual exotics, Conyza bonariensis and C. canadensis, in Mediterranean annual communities. We wanted to test whether herbivory of these exotics was influenced either by species richness, functional-group richness or functional-group composition. We also studied the relationship between herbivory on the exotic species and their performance. Herbivory increased with increasing species and functional-group richness for both Conyza species. These patterns are interpreted as reflecting a greater number of available herbivore niches in a richer, more complex, plant community. The identities of functional groups also affected Conyza herbivory, which decreased in the presence of Asteraceae or Fabaceae and increased in the presence of Poaceae. Increasing herbivory had consequences for vegetative and demographic parameters of both invasive species: survival, final biomass and net fecundity decreased with increasing herbivory, leading to a loss of reproductive capacity. We conclude that communities characterised by a high number of grass species instead of Asteraceae or Fabaceae may be more resistant to invasion by the two Conyza species, in part due to predation by native herbivores.     

Fire regimes and shifting community patterns: a case study and method for species with complex fire requirements

Anthropogenic alteration of fire regimes is implicated in the extinction or decline of species across the globe. Active management of fire regimes using specified guidelines can help sustain diversity and counter this loss. However, some species occur across multiple communities with differing fire regime requirements or may exploit the dynamics of associated ecotones. These complexities are not easily understood via a simple consideration of life history traits or unravelled by a simple experimental framework. Population modelling, however, may provide valuable insights in these instances, but to date this remains unexplored. A population model based on detailed demographic, habitat and fire regime data was developed for the species Prostanthera askania. The species occurs across rainforest/eucalypt forest ecotones subject to long-term alteration, fragmentation and invasion by exotic species. Modelling revealed that current recommended fire guidelines will not sustain this species under these novel conditions. Fire regimes that minimised extinction risk in some scenarios had double or five times the fire frequencies of recommended regimes. Managing fire was also more important in specific habitat (gully habitat rather than on slopes or ridges). Currently applied management actions for Prostanthera askania do not include any fire management and the regimes that apply to most habitat in which the species occurs (no fire or fire at up to 50-year intervals in gully habitat) is not optimal for the species. Management will be substantially more effective if specific fire treatments revealed by this study are employed (e.g. fire intervals of 20–30 years in gully habitat). The study demonstrates that a conceptualisation and consideration of communities as both temporally and spatially dynamic can substantially contribute to better fire management outcomes. The approach used herein can be readily adapted and applied to a plethora of species via a range of software packages and codes.     

Stoichiometry, herbivory and competition for nutrients: simple models based on planktonic ecosystems

Models are examined in which two prey species compete for two nutrient resources, and are preyed upon by a predator that recycles both nutrients. Two factors determine the effective relative supply of the nutrients, hence competitive outcomes: the external nutrient supply ratio, and the relative recycling of the two nutrients within the system. This second factor is governed by predator stoichiometry--its relative requirements for nutrients in its own biomass. A model with nutrient resources that are essential for the competing prey is detailed. Criteria are given to identify the limiting nutrient for a food chain of one competitor with the predator. Increased supply of this limiting nutrient increases predator density and concentration of this nutrient at equilibrium, while decreasing the concentration of a non-limiting nutrient. Changes in supply or recycling of a non-limiting nutrient affect only the concentration of that nutrient. Criteria for the invasion of a second prey competitor are presented. When different nutrients limit growth of the resident prey and the invader, increased supply or recycling of the invader's limiting nutrient assists invasion, while increased supply or recycling of the resident's limiting nutrient hinders invasion. If the same nutrient limits both resident and invader, then changes in supply and recycling have complex effects on invasion, depending on species properties. In a parameterized model of a planktonic ecosystem, green algae and cyanobacteria coexist over a wide range of nitrogen:phosphorus supply ratios, without predators. When the herbivore Daphnia is added, coexistence is eliminated or greatly restricted, and green algae dominate over a wide range of supply conditions, because the effective supply of P is greatly reduced as Daphnia rapidly recycles N.     

Disease, parasitism and herbivory: Multidimensional challenges in plant evolution

Plants provide feeding sites for a broad range of parasites, commensals and symbionts. In the process, they can be subjected to selection whenever feeding choices are based upon heritable traits of the plants and the feeding affects plant fitness. The outcome of such selection depends on the correlation between choices made by various taxa. Recent work suggests that this multispecies selection, although common in natural communities, now needs to be incorporated more fully into ecological and evolutionary perspectives.     

Spatial interplay of plant competition and consumer foraging mediate plant coexistence and drive the invasion ratchet

Indirect effects may play an important role in structuring plant communities. Using a spatially explicit model of consumer foraging and plant competition, we demonstrate how the relationship between the spatial area over which plants compete and the spatial scale of consumer behaviour can determine the outcome of competition when one plant species provides a refuge for mobile consumers (i.e. refuge-mediated apparent competition). Once an initial population of the invader is established, complete invasion may be inevitable because of an ever-advancing invasion front ratchets forward driven by a feeding front of mobile consumers. Because the spatial extent of apparent competition determines the area available for colonization, consumers may also dictate the rate at which an invasion occurs. We find that, as long as refuge-mediated apparent competition is sufficiently localized, invasion is possible even in systems characterized by low overall levels of consumer pressure. Moreover, we show that a stable equilibrium can result in which both resident and invading plants coexist, suggesting that spatial heterogeneity created by refuge-mediated apparent competition may be important in mediating coexistence in plant communities. The spatial interplay of consumer behaviour and plant competition may be an underappreciated mechanism affecting the composition, diversity and spatial pattern of plant communities.     

Interference competition by Argentine ants displaces native ants: implications for biotic resistance to invasion

The Argentine ant Linepithema humile (Dolichoderinae) is one of the most widespread invasive ant species in the world. Throughout its introduced range, it is associated with the loss or reduced abundance of native ant species. The mechanisms by which these native species are displaced have received limited attention, particularly in Australia. The role of interference competition in the displacement of native ant species by L. humile was examined in coastal vegetation in central Victoria (southeastern Australia). Foragers from laboratory colonies placed in the field consistently and rapidly displaced the tyrant ant Iridomyrmex bicknelli, the big-headed ant Pheidole sp. 2, and the pony ant Rhytidoponera victoriae from baits. Numerical and behavioural dominance enabled Argentine ants to displace these ants in just 20 min; the abundance of native species at baits declined 3.5–24 fold in direct relation to the rapid increase in L. humile. Most precipitous was the decline of I. bicknelli, even though species in this typically dominant genus have been hypothesized to limit invasion of L. humile in Australia. Interspecific aggression contributed strongly to the competitive success of Argentine ants at baits. Fighting occurred in 50–75% of all observed interactions between Argentine and native ants. This study indicates that Argentine ants recruit rapidly, numerically dominate, and aggressively displace from baits a range of Australian native ant species from different subfamilies and functional groups. Such direct displacement is likely to reduce native biodiversity and indirectly alter food web structure and ecosystem processes within invaded areas. Biotic resistance to Argentine ant invasion from native ants in this coastal community in southeastern Australia is not supported in this study.     

Three New Species of Nothacrobeles (Nemata: Cephalobidae) from the Mojave Desert, California

Three new species of Nothacrobeles are described from localities in the Mojave Desert, southern California. Nothacrobeles triniglarus n. sp. is characterized by the presence of a long post-vulval sac and three tubular adoral projections. Both N. spatulatus n. sp. and N. nanocorpus n. sp. are smaller than any other known species within the genus. Nothacrobeles spatulatus n. sp. has labial probolae that are short and spatulate without a basal ridge, whereas those of N. nanocorpus n. sp. are flattened and plate-like. Furthermore, N. nanocorpus n. sp. is unique by its extremely short esophageal corpus (less than 25 µm long in adult females) and the small size of its guard processes. An emended diagnosis of the genus is given to accommodate distinctive characteristics of these new species. A table comparing the 11 valid species of Nothacrobeles is presented.     

Status competition,inequality, and fertility: implications for the demographic transition

The role that social status plays in small-scale societies suggests that status may be important for understanding the evolution of human fertility decisions, and for understanding how such decisions play out in modern contexts. This paper explores whether modelling competition for status—in the sense of relative rank within a society—can help shed light on fertility decline and the demographic transition. We develop a model of how levels of inequality and status competition affect optimal investment by parents in the embodied capital (health, strength, and skills) and social status of offspring, focusing on feedbacks between individual decisions and socio-ecological conditions. We find that conditions similar to those in demographic transition societies yield increased investment in both embodied capital and social status, generating substantial decreases in fertility, particularly under conditions of high inequality and intense status competition. We suggest that a complete explanation for both fertility variation in small-scale societies and modern fertility decline will take into account the effects of status competition and inequality.