Habitat complexity facilitates coexistence in a tropical ant community

The role of habitat complexity in the coexistence of ant species is poorly understood. Here, we examine the influence of habitat complexity on coexistence patterns in ant communities of the remote Pacific atoll of Tokelau. The invasive yellow crazy ant, Anoplolepis gracilipes (Smith), exists in high densities on Tokelau, but still coexists with up to seven other epigeic ant species. The size-grain hypothesis (SGH) proposes that as the size of terrestrial walking organisms decreases, the perceived complexity of the environment increases and predicts that: (1) leg length increases allometrically with body size in ants, and (2) coexistence between ant species is facilitated by differential habitat use according to body size. Analysis of morphological variables revealed variation inconsistent with the morphological prediction of the SGH, as leg length increased allometrically with head length only. We also experimentally tested the ability of epigeic ants in the field to discover and dominate food resources in treatments of differing rugosity. A. gracilipes was consistently the first to discover food baits in low rugosity treatments, while smaller ant species were consistently the first to discover food baits in high rugosity treatments. In addition, A. gracilipes dominated food baits in planar treatments, while smaller ant species dominated baits in rugose treatments. We found that the normally predictable outcomes of exploitative competition between A. gracilipes and other ant species were reversed in the high rugosity treatments. Our results support the hypothesis that differential habitat use according to body size provides a mechanism for coexistence with the yellow crazy ant in Tokelau. The SGH may provide a mechanism for coexistence in other ant communities but also in communities of other terrestrial, walking insects that inhabit a complex landscape.Electronic Supplementary Material Supplementary material is available to authorised users in the online version of this article at .     

Supercolonies of the invasive yellow crazy ant, Anoplolepis gracilipes, on an oceanic island: Forager activity patterns, density and biomass

Summary. Ants have the capacity to reach unusually high densities, mostly in their introduced ranges. Numerical dominance is often cited as key to the ability of exotic ants to displace native ant species, reduce the abundance of invertebrates and negatively impact upon bird, land crab and other vertebrate populations. On Christmas Island, Indian Ocean, the yellow crazy ant, Anoplolepis gracilipes (Jerdon), forms supercolonies, where extremely high densities of foraging ants have contributed to ‘invasional meltdown’ in rainforest areas. Densities of up to 2254 foraging ants per m² and a biomass of 1.85 g per m² were recorded, and nest densities reached 10.5 nest entrances per m². Populations of A. gracilipes can overcome and kill red endemic land crabs (Gecarcoidea natalis) over 100 times their own biomass. This is the highest recorded density of foraging ants, and adds another element to the definition of ‘supercolony’ of unicolonial ants. This paper documents one extreme in a continuum of densities of unicolonial, invasive ant species and highlights the need to incorporate forager densities into invasive ant research.Received 17 November 2004; revised 14 February 2005, accepted 21 February 2005.     

Impact of the invasive crazy ant Anoplolepis gracilipes on Bird Island, Seychelles

The crazy ant (Anoplolepis gracilipes) invaded Bird island, Seychelles, in the 1980s. In 1997, its range expanded and population densities increased. The impacts of this change were studied in 2001 using a combination of arthropod collecting methods. The ant population excluded larger invertebrates (principally the large ant Odontomachus simillimus and the crabs, principally Ocypode spp.). Cockroaches, however, remained abundant in ant-infested areas and tree-nesting birds (Lesser Noddy Anous tenuirostris) appear to be able to breed successfully in the presence of the crazy ant. The ants are only abundant in areas of deep shade which provide cool nesting areas, yet enabling them to forage in the open when ground temperatures fall. The expansion of the ants was correlated with the regeneration of woodland on the island. Recommendations are made for the management of the woodland which may reduce the impacts of the crazy ant.     

Invasive ants compete with and modify the trophic ecology of hermit crabs on tropical islands

Invasive species can dramatically alter trophic interactions. Predation is the predominant trophic interaction generally considered to be responsible for ecological change after invasion. In contrast, how frequently competition from invasive species contributes to the decline of native species remains controversial. Here, we demonstrate how the trophic ecology of the remote atoll nation of Tokelau is changing due to competition between invasive ants (Anoplolepis gracilipes) and native terrestrial hermit crabs (Coenobita spp.) for carrion. A significant negative correlation was observed between A. gracilipes and hermit crab abundance. On islands with A. gracilipes, crabs were generally restricted to the periphery of invaded islands. Very few hermit crabs were found in central areas of these islands where A. gracilipes abundances were highest. Ant exclusion experiments demonstrated that changes in the abundance and distribution of hermit crabs on Tokelau are a result of competition. The ants did not kill the hermit crabs. Rather, when highly abundant, A. gracilipes attacked crabs by spraying acid and drove crabs away from carrion resources. Analysis of naturally occurring N and C isotopes suggests that the ants are effectively lowering the trophic level of crabs. According to δ¹⁵ N values, hermit crabs have a relatively high trophic level on islands where A. gracilipes have not invaded. In contrast, where these ants have invaded we observed a significant decrease in δ¹⁵ N for all crab species. This result concurs with our experiment in suggesting long-term exclusion from carrion resources, driving co-occurring crabs towards a more herbivorous diet. Changes in hermit crab abundance or distribution may have major ramifications for the stability of plant communities. Because A. gracilipes have invaded many tropical islands where the predominant scavengers are hermit crabs, we consider that their competitive effects are likely to be more prominent in structuring communities than predation. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Community level impacts of an ant invader and food mediated coexistence

Ant invasions exert a range of effects on recipient communities, from displacement of particular species to more complex community disruption. While species loss has been recorded for a number of invasion events, a little examined aspect of these invasions is the mechanisms for coexistence with resident ant species.The yellow crazy ant, Anoplolepis gracilipes (Smith), is considered one of the world’s worst ant invaders and has recently undergone rapid population growth in Tokelau. We surveyed the ground-dwelling ant fauna in two plots on each of five invaded and three uninvaded islands across two atolls in Tokelau to examine community characteristics of the ant fauna in areas with and without yellow crazy ants. We also used three types of food bait (tuna, jam and peanut butter) to experimentally test if species are able to coexist by consuming different food resources. Anoplolepis gracilipes was found to coexist with two to six other ant species at any one site, and coexisted with a total of 11 ant species. Four species never co-occurred with A. gracilipes. Non-metric multidimensional scaling showed significant differences in community composition and the relative abundance of species between areas that had, and had not, been invaded by A. gracilipes. The number of other ant species was significantly lower in communities invaded by the yellow crazy ant, but did not decline with increasing A. gracilipes abundance, indicating that impacts were independent of population density. The yellow crazy ant dominated all tuna and jam baits, but had a low attendance on peanut butter, allowing four other ant species to access this resource. Our results demonstrate community level impacts of an ant invader on a tropical oceanic atoll and suggest that differing use of food resources can facilitate coexistence in ant communities. Received 11 September 2006; revised 15 January 2007; accepted 22 February 2007.     

Intercontinental union of Argentine ants: behavioral relationships among introduced populations in Europe, North America, and Asia

Many invasive ants, including the Argentine ant Linepithema humile, form expansive supercolonies, within which intraspecific aggression is absent. The behavioral relationships among introduced Argentine ant populations at within-country or within-continent scales have been studied previously, but the behavioral relationships among intercontinental populations have not been examined. The present study investigated the levels of aggression among intercontinental Argentine ant populations by transporting live ants from Europe and California to Japan and conducting aggression tests against Japanese populations. Workers from the dominant supercolonies of Europe and California did not show aggressive behavior toward workers from the dominant supercolony of Japan, whereas they fought vigorously against workers from minor supercolonies. The three massive supercolonies, together with Argentine ants from Macaronesia, may be the largest non-aggressive unit formed by a social insect species in which intraspecific aggression exists. Absence or low levels of aggression at transcontinental scale, which may have derived from low genetic variation, may help introduced Argentine ants maintain expansive supercolonies. The lack of aggression implies possible frequent exchanges of individuals among the intercontinental populations mediated by human activities.     

Interference and exploitation competition of three nectar-thieving invasive ant species

Summary. Plant and insect exudates are known to play a key role in structuring tropical ant communities, but less is known about the utilization of these resources in communities dominated by invasive ants. Invasive ants are thought to require large amounts of carbohydrates such as honeydew or nectar to maintain their high abundances. Invasive ants that consume floral nectar may compete with legitimate floral visitors through interference or exploitation competition. I compared the nectar-thieving behavior of three widespread invasive ant species: long-legged ants (Anoplolepis gracilipes), Argentine ants (Linepithema humile), and big-headed ants (Pheidole megacephala) in inflorescences of the native Hawaiian ‘ōhi’a tree, an important food source for native fauna. A. gracilipes was least likely to leave inflorescences unvisited and visited inflorescences in higher numbers than both L. humile and P. megacephala. A. gracilipes and L. humile visited more flowers in an inflorescence and were less likely to retreat from a flower with a competitor than P. megacephala. A. gracilipes was able to take 5.5 and 11.3 times the amount of nectar than L. humile and P. megacephala, respectively. Thus, A. gracilipes may be effective at both interference and exploitation competition against other nectarivores, L. humile may be effective at interference competition, and P. megacephala may be relatively weak at both types of competition against other nectarivores. Ascertaining the competitive abilities of invasive ants against legitimate floral visitors will be especially important in agricultural and other systems that are nectar or pollinator limited.Received 6 December 2004; revised 13 January 2005; accepted 14 January 2005.     

Twenty four new microsatellite markers in two invasive pavement ants, <Emphasis Type="Italic">Tetramorium</Emphasis> sp.E and <Emphasis Type="Italic">T. tsushimae</Emphasis> (Hymenoptera: Formicidae)

Invasive species trigger biodiversity losses and alter ecosystem functioning, with life history shaping invasiveness (Sakai et al., Annu Rev Ecol Syst 32:305–332, 2001). However, pinpointing the relation of a specific life history to invasion success is difficult. One approach may be comparing congeners. The two Palearctic pavement ants, Tetramorium sp.E (widely known as T. caespitum, Schlick-Steiner et al., Mol Phylogenet Evol 40:259–273, 2006) and T. tsushimae have invaded North America (Steiner et al., Biol Invasions 8:117–123, 2006). Their life histories differ in that T. sp.E has separate single-queened colonies but T. tsushimae multi-queened colonies scattered over large areas (Sanada-Morimura et al., Insect Soc 53:141–148, 2006; Schlick-Steiner et al., Mol Phylogenet Evol 40:259–273, 2006; Steiner et al., Biol Invasions 8:117–123, 2006). Comparison of the genetic diversity in the entire native and non-native ranges will elucidate the invasion histories. Here, we present 13 and 11 microsatellites, developed for T. sp.E and T. tsushimae, respectively, and characterize all for both species. Florian M. Steiner, Wolfgang Arthofer and Birgit C. Schlick-Steiner contributed equally to this work.     

Detrimental effects of two widespread invasive ant species on weight and survival of colonial nesting seabirds in the Hawaiian Islands

Invasive ants are a significant conservation concern and can have far-reaching effects in ecosystems they invade. We used the experimental control of ant numbers on two pairs of small (<5 ha) offshore islets dominated by either the big-headed ant, Pheidole megacephala or the tropical fire ant, Solenopsis geminata to investigate the influence of these species on seabird hatching success, fledging success and weight. Limited unpublished observations of both ant species attacking nesting seabirds exist, but the frequency of attacks or how they affect seabird growth and survival are unknown. Island-wide treatments with hydramethylnon resulted in the eradication of P. megacephala and the temporary reduction of S. geminata densities. No difference in hatching success, growth, or fledging success of Wedge-tailed Shearwaters (Puffinus pacificus), a common colonial nesting seabird in the Hawaiian Islands was observed on the pair of islets dominated by P. megacephala. On islets dominated by S. geminata, ant control resulted in a temporary increase in fledging success. Injury frequency increased dramatically on the untreated islet (8.3–100%) while remaining the same on the treated islet (27–38%). Severely injured chicks (i.e., chicks that lost >20% of tissue on their feet) weighed significantly less than uninjured chicks and did not fledge. It is unclear if the chicks were being preyed upon or stung in defense of nearby ant colonies. Radical changes in invasive ant populations have been noted, and booming ant populations could cause short-term, but widespread damage to seabird colonies. The negative effects of invasive ants on seabirds may be difficult to detect, and therefore unknown or underestimated throughout the world where the two groups overlap.     

Differences in life history strategies between an invasive and a competing resident seed predator

Competitive interactions may arise from biological invasion if a successful invasive species requires the ecological niche of a resident one. Life-history traits that make a species a successful invader are of particular interest in elucidating both invasion success and how interspecific competition may emerge. In southeastern France, the invasion of cedar forests by the seed chalcid Megastigmus schimitscheki generated competitive relationships with the resident M. pinsapinis for the exploitation of the seed resource. Among the numerous ecological traits allowing these seed predators to exploit their niche successfully, the timing of adult emergence, initial egg load and age-specific realized fecundities of females were investigated to help understanding the issue of such interspecific relationships. Spring adult emergence of M. schimitscheki under natural conditions was significantly earlier than that of M. pinsapinis, suggesting that an advantage for the access to the seed resource for the invasive species may be associated with this trait. Initial egg load was significantly higher in M. schimitscheki than in M. pinsapinis and the analysis of age-specific realized fecundity in semi-natural conditions indicated that both M. schimitscheki and M. pinsapinis females lay a large proportion of their eggs during the early days of their lives. In the light of these findings, both earlier phenology and higher reproduction abilities of M. schimitscheki may have the potential to confer an advantage within a competitive context with M. pinsapinis through enhanced seed resource preemption. This may potentially explain the invasiveness of M. schimitscheki in southern France despite the presence of its closely related competitor M. pinsapinis.     

Invasion history, habitat preferences and phenology of the invasive ladybird Harmonia axyridis in Belgium

Harmonia axyridis Pallas (Coleoptera: Coccinellidae) is native to Asia, and was widely introduced as a biocontrol agent of aphids and coccids in Europe and North America. In Europe, H. axyridis is considered to be an invasive alien species because of its potential to disrupt native ladybird communities. Since 1999, the Belgian Ladybird Working Group mapped all Belgian Coccinellidae and recorded data on substratum plants and habitat. The first feral H. axyridis population in Belgium was recorded in 2001, but the expansion rate is decreasing because it now colonised the whole country. Recorded occupancy in Belgium showed an average rate of increase of 189% between 2002 and 2006. In Belgium, H. axyridis occurred in a wide range of habitats, including those of high conservation value. However, habitat and land cover analysis showed that H. axyridis is more frequently found in urbanised landscapes than in semi-natural landscapes. A niche overlap analysis based on plant use data showed that the potential to affect native species is higher for generalist, deciduous and coniferous tree ladybird species than for heathland and wetland specialist species. Phenology data showed that H. axyridis is able to reproduce later in the year than native species. Based on recorded distribution, ecology and phenology, we discuss the potential of H. axyridis to disrupt native ladybird assemblages in Belgium.     

Differences in the effects of parental age on offspring life history between tropical and temperate populations of milkweed bugs (Oncopeltus spp.)

Summary We compare the effects of parental age on several offspring life history traits in two milkweed bug populations: a typically univoltine population ofOncopeltus fasciatus from the Sacramento Valley of California and a typically multivoltine population ofO. cingulifer from Monteverde, Costa Rica. Reared under identical conditions (27°C, 12 h light: 12 h dark photoperiod), these bugs exhibit significant differences in the effects of parental age on offspring life history. As they age,O. fasciatus females from Sacramento lay clutches of eggs of decreasing average weight and a decreasing proportion of their eggs produce offspring that survive to adulthood. Those offspring which do survive have a significantly faster developmental rate and females have a larger body size at adult eclosion. AsO. cingulifer from Monteverde age, they also produce lighter eggs, but there is no significant change in the offspring developmental rate, survival or female adult body size. We suggest that these results are largely explicable as the consequence of different parental investment strategies associated with the predictable relationship between parental age and time of season inO. fasciatus but not inO. cingulifer. AsO. fasciatus from Sacramento age, they may be increasing their investment per developing offspring so as to increase the probability that nymphs hatching late in the season will reach a prereproductive adult diapause before the first killing frost.     

Reproductive strategy of an invasive thistle: effects of adults on seedling survival

Invasive species are known for their ability to form monocultures that exclude native species, yet intraspecific interactions among invasives have not been well studied. Cynara cardunculus (L.) is an invasive perennial thistle that establishes high-density populations in coastal California grasslands. We examined the natural distribution of C. cardunculus seedlings in an established population and found that nearly 100% of seedlings grew within 2 m of adults despite an expected distribution peak at 3 m from source plants based on measured dispersal distances. We then investigated the role of mature plants in seedling survival and establishment with regard to live vegetation, litter, and seedling distance by planting seedlings at increasing distances around adults and applying removal treatments to the focal adult rosettes. We applied control (no removal), adult rosette removal (live leaves), litter removal (dead leaves), and adult rosette plus litter removal (all aboveground plant material) treatments. Seedlings experienced a higher rate of survival, measured by senescence date, and establishment, measured by return rate the following year, with all adult rosette removal treatments. Inhibition by adult rosettes was reduced with distance to 60–80 cm from the rosette, and there was little effect of adult plants between 80 and 200 cm. These results suggest that adult rosettes may inhibit conspecific seedlings at very close distances but provide a favorable environment for seedlings within nearby interspaces. This pattern may contribute to the creation and maintenance of high-density populations in C. cardunculus. Land managers seeking to control this species may improve long-term effectiveness by expanding management efforts to include a 2 m radius around adult plants and treating within 5 months of seedling emergence to prevent recruitment rather than treating adults alone.     

An evaluation of the impact of Melaleuca quinquenervia invasion and management on plant community structure after fire

Two questions were asked in this study: after a fire, does the choice of invasive plant management strategy, namely herbicidal or biological, alter (1) plant community assemblages and (2) the re-invasion potential of the Australian tree Melaleuca quinquenervia? Plant species richness was highest in the non-invaded and herbicide sites compared to the biological site with 10.5, 10.8, and 8.25 species m⁻² found in each site, respectively. Although the total count of live and dead seedlings was highest in the biologically controlled site at 22.8 and 13.6 plants m⁻², respectively, M. quinquenervia seedlings were recruited in all sites. While the ultimate goal of management programs is to restore ecosystem integrity, this work provides evidence that passive restoration may not be enough to restore plant community structure in this system.     

The effects of light on foliar chemistry,growth and susceptibility of seedlings of a canopy tree to an attine ant

Summary Seedlings of Inga oerstediana Benth. (Mimosaceae) growing in three different light environments (the understory, tree-fall gaps and full sun) were tested for differences in chemistry (nutrients and tannins), wound-induced increases in tannins, growth, and susceptibility to leaf-cutter ants, Atta cephalotes (L.) (Formicidae: Attini). I hypothesized that seedlings of I. oerstediana would contain higher concentrations of tannins when growing in high light conditions and, therefore, would be less susceptible to leaf-cutter ants.Foliar concentrations of condensed tannins were much higher in plants growing in full sun compared to those growing in the understory. The concentrations of condensed tannins did not increase following damage. Despite higher concentrations of condensed tannins in sun foliage, leaf-cutter ants found these leaves more acceptable. The preference for sun leaves was consistent with higher concentrations of foliar nutrients. I suggest that the magnitude of the increase in condensed tannins was not great enough to override the benefits of increased concentrations of foliar nutrients. Finally, based on these results and those of others, I suggest that foraging by leaf-cutter ants may be an important factor determining patterns of succession in early successional habitats.     

Parsimony dictates a human introduction: on the use of genetic and other data to distinguish between the natural and human-mediated invasion of the European snail Littorina littorea in North America

Two centuries of historical, archaeological, paleontological, geological, oceanographical and biological data conclusively indicate that the periwinkle snail Littorina littorea was introduced to North America from Europe either by Norse explorers 1000 years ago or by European colonists after 1840. Available genetic data do not indicate ancient divergence of North American and European L. littorea and thus do not challenge all other evidence that it was introduced from Europe by humans.     

Concordance between life history traits, invasion history, and allozyme diversity of the Everglades invader Melaleuca quinquenervia

During the century following its initial introduction in 1886, the Australian tree Melaleuca quinquenervia (Myrtaceae) dispersed from a few introduction points to occupy over 200,000 ha, primarily in historic Everglades wetlands of southern Florida. Cellulose acetate gel electrophoresis (CAGE) was used to investigate the allozyme diversity and population genetic structure of 208 individuals in a dozen populations resulting from this invasion. The analyses showed that these populations have a high (82%) rate of polymorphic loci and an average of 2 alleles/locus. There was substantial heterozygosity (mean H_e = 0.356), which concords well with recent studies reporting a greater number of introduction events and sources than generally recognized. The introduction history and distributional patterns within Florida have led to geographic structuring (G_ST = 0.419) in which the Gulf Coast metapopulation has a greater effective number of alleles and greater heterozygosity than the Atlantic Coast metapopulation. The gene diversity in M. quinquenervia was comparable to other tropical woody species. Its strong population divergence was reminiscent of pioneer species and consistent with its status as a plant invader in Florida.     

Old pythons stay fit; effects of haematozoan infections on life history traits of a large tropical predator

We document the impact of blood parasite infections caused by Hepatozoon sp. on water python (Liasis fuscus) life history traits such as growth rates, condition, reproductive output and survival. Individual snakes maintained similar among-year parasite loads. Hepatozoon infections affected python growth rate, i.e. snakes suffering from high infection levels exhibited significantly slower growth compared to individuals with low parasite loads. Our results suggest that the parasites also affected the pythons nutritional status (condition), as snakes with low condition scores suffered from higher parasite infection levels than snakes with high scores. Furthermore, our data suggest that parasitaemia may affect female reproductive output, as reproductive female pythons harboured lower parasite loads compared to non-reproductive adult females. High levels of parasite infections also affected juvenile python survival, as recaptured snakes harboured significantly lower parasite loads compared to non-recaptured yearling pythons. In our study area, water python have very few natural predators and, hence, experience low mortality rates and commonly reach an age of >15 years. In contrast to results obtained in other studies, parasite loads in larger/older pythons were lower compared to younger snakes, suggesting that only snakes harbouring lower levels of parasitaemia were able to survive to old age. We suggest that a possible cause for the opposing results regarding parasite prevalence and host age may be due to different levels of extrinsic mortality rates and longevity. Long-lived organisms, such as water pythons, may invest relatively more into crucial self-maintenance functions such as parasite defence, compared to short-lived organisms.     

Direct and indirect impacts of predation by fish on the zooplankton community: an experimental analysis using tanks

The impact of Pseudorasbora parva, a common zooplanktivorous fish species in Japan, on a zooplankton community was analyzed in experimental tanks, half of which were stocked with the fish. Different zooplankton species showed different responses to the introduction of the fish. In the presence of the fish, the populations of the large cladoceran Ceriodaphnia and the predatory copepod Mesocyclops were reduced, but the population of the herbivorous copepod Eodiaptomus and the small cladocerans Bosmina fatalis and Bosminopsis deitersi increased relative to the controls. The increase of Mesocyclops seen in the control tanks might have suppressed the populations of the small cladocerans, which are vulnerable to invertebrate predation. The results suggest that the population densities of the large prey items preferred by the fish, Ceriodaphnia and Mesocyclops, were controlled directly by fish predation, but the population densities of the smaller and less preferred zooplankton were controlled indirectly through the food-web cascade.     

Competitive impacts and responses of an invasive weed: dependencies on nitrogen and phosphorus availability

Changes in competitive interactions under conditions of enhanced resource availability could explain the invasion success of some problematic plant species. For one invader of North American grasslands, Centaurea diffusa (diffuse knapweed), we test three hypotheses: (1) under ambient (high resource) conditions, C. diffusa is better able to tolerate competition from the resident community (competitive response), (2) under ambient conditions, C. diffusa strong impacts the competitive environment (competitive effect), and (3) reductions in nitrogen and/or phosphorus availability diminish these advantages. In support of our first hypothesis, C. diffusa was the most tolerant to neighbor competition of the four focal species under current resource conditions. In opposition to our second hypothesis, however, neighborhoods that contained C. diffusa and those where C. diffusa had been selectively removed did not differ in their impact on the performance of target transplant individuals or on resource conditions. Reduction in resource availability influenced competitive tolerance but not competitive impact, in partial support of our last hypothesis. Reduction in soil nitrogen (via sucrose carbon addition) enhanced the degree of neighbor competition experienced by all species but did not change their relative rankings; C. diffusa remained the best competitor under low nitrogen conditions. Reduction of soil phosphorus (via gypsum addition) weakened the ability of C. diffusa to tolerate neighbor competition proportionately more than the other focal species. Consequently, under low phosphorus conditions, C. diffusa lost its competitive advantage and tolerated neighbor competition similarly to the other focal species. We conclude that C. diffusa invasion may be double-edged: C. diffusa is less limited by nitrogen than the other focal species and is better able to utilize phosphorus to its competitive advantage.