The urban fire ant paradox: native fire ants persist in an urban refuge while invasive fire ants dominate natural habitats

In contrast to the widespread extirpation of native fire ants (Solenopsis geminata) across southern US following the invasion by imported red fire ants (S. invicta), some residential areas of Austin form unexpected refuges for native fire ants. Ironically, these urban environments provide refuges for the native fire ants while adjacent natural habitats have been overrun by invasive fire ants. Resistance to invasive fire ants in these urban areas occurs mainly in older residential properties constructed prior to the S. invicta invasion, while more recent construction has allowed establishment by S. invicta. The invasive ability of S. invicta is often attributed to escape from parasitoids and efficient dispersal of polygyne multiple queen colonies. Here we also show the importance of landscape parameters in the invasion process, where low levels of disturbance and continuous plant cover in older residential areas form possible barriers to colonization. Dense leaf cover (high NDVI) was also found to be associated with native ant refuges. Long term residential land ownership may have resulted in lower recent disturbance levels and increased plant cover that support refuges of native fire ants.     

The role of ants and mammals in dispersal and post-dispersal seed predation of the shrubs Grevillea (Proteaceae)

The role seed predators play in influencing the dynamics of plant populations has been little studied in Australia. The interaction of ant dispersal and seed predation on the soil seedbank in six shrubby species of Grevillea from the Sydney region of southeastern Australia was examined in selective exclusion experiments, seed array trials and placement of single seeds on the ground.Two distinct seed types in Grevillea were examined and different seed dispersal and post-dispersal seed predation patterns were associated with each: (a) seeds lacking an elaiosome were not attractive to ants and annual seed losses of between 82 and 95% were found in vegetation unburnt for greater than 8 years. Native rodents, Rattus fuscipes, and macropods, Wallabia bicolor, were responsible for these seed losses; (b) seeds with an elaiosome were rapidly handled by ants. Two functional types of ants were recognised. Most encounters were by ants that were small (Local) relative to seed size and these ants simply removed the elaiosome in situ or moved seeds only small distances (<20 cm). Some 0–24% of ant/seed encounters were by large (Removalist) ant species that were capable of moving seeds back to nests. In addition, Rattus fuscipes and Wallabia bicolor consumed at least 32–68% of seeds of Grevillea species with an elaiosome.Ants may reduce the overall levels of seed predation where seeds moved by Removalist ant species escape predation and are deposited in safe sites, hence allowing more seeds to reach the persistent soil seedbank. Mammals do not consume all seeds when ants are excluded, allowing for the potential for some seed escape from predation after seeds are discarded by Local ant species.     

Contrasting effects of resource availability and plant mortality on plant community invasion by Bromus tectorum L.

The positive effect of disturbance on plant community invasibility is one of the more consistent results in invasion ecology. It is generally attributed to a coincident increase in available resources (due to the disturbance) that allows non-resident plant species to establish (Davis MA, Grime JP Thompson K, J Ecol 88:528–534, 2000). However, most research addressing this issue has been in artificial or highly modified plant communities. Our goal in this study was to investigate the interactive effects of resource availability and plant mortality disturbance on the invasion of natural plant communities. We conducted a series of experiments that examined the response of Bromus tectorum L., a highly invasive annual grass, to experimentally created gradients of resource availability [nitrogen (N) and water] and resident plant species mortality. We found that B. tectorum biomass was co-limited by N and water. Biomass at the end of the growing season was a saturating function (i.e., increased to a maximum) of water, which determined maximum biomass, and N, which determined the rate at which maximum biomass was attained. Despite that fact that plant mortality increased N availability, it had a negative impact on invasion success. Plant mortality also decreased foliar cover, standing dead biomass, and soil cover by litter. In harsh environments, removing foliar and soil cover may increase germination and seedling stress by increasing soil temperatures and water loss. Across all treatments, B. tectorum success decreased with decreasing foliar cover and standing dead biomass. This, in combination with the strong limitation of B. tectorum biomass by water in this experiment, suggests that our plant mortality disturbance removed soil cover that may have otherwise aided B. tectorum invasion into this semi-arid plant community by reducing water stress.     

Butterfly seed predation: effects of landscape characteristics,plant ploidy level and population structure

Polyploidization has been suggested as one of the most common mechanisms for plant diversification. It is often associated with changes in several morphological, phenological and ecological plant traits, and therefore has the potential to alter insect–plant interactions. Nevertheless, studies evaluating the effect of plant polyploidy on interspecific interactions are still few. We investigated pre-dispersal seed predation by the butterfly Anthocharis cardamines in 195 populations of two ploidy levels of the herb Cardamine pratensis (tetraploid ssp. pratensis, 2n = 30 vs. octoploid ssp. paludosa, 2n = 56–64). We asked if differences in incidence and intensity of predation among populations were related to landscape characteristics, plant ploidy level and population structure. The incidence of the seed predator increased with increasing plant population size and decreasing distance to nearest population occupied by A. cardamines. The intensity of predation decreased with increasing plant population size and was not affected by isolation. Probability of attack decreased with increasing shading, and intensity of predation was higher in grazed than in non-grazed habitats. The attack intensity increased with increasing mean flower number of plant population, but was not affected by flowering phenology. Individuals in tetraploid populations suffered on average from higher levels of seed predation, had higher mean flower number, were less shaded and occurred more often in grazed habitats than octoploid populations. When accounting for differences in habitat preferences between ploidy levels there was no longer a difference in intensity of predation, suggesting that the observed differences in attack rates among populations of the two ploidy levels are mediated by the habitat. Overall, our results suggest that polyploidization is associated with differentiation in habitat preferences and phenotypic traits leading to differences in interspecific interaction among plant populations. This, in turn, may facilitate further divergence of ploidy levels.     

Chemical differences between seeds and elaiosomes indicate an adaptation to nutritional needs of ants

Ant-dispersed plants usually produce seeds with appendages (elaiosomes) as reward for ants. Plants that produce high-quality elaiosomes benefit because ants preferentially disperse their diaspores. We therefore hypothesized that seeds and elaiosomes differ in chemical composition in ways that make elaiosomes of high nutritional quality for ants, capable of providing essential dietary components that explain the increased fitness and higher gyne production documented for colonies with elaiosome consumption. To test the hypothesis we analysed the content and composition of lipids, amino acids, soluble carbohydrates, proteins and starch in seeds and elaiosomes of 15 central European ant-dispersed plants. After separating the different fractions, total lipids were determined gravimetrically, fatty acids and soluble carbohydrates were detected by gas chromatography (GC) and GC–mass spectrometry, free amino acids by an amino acid analyser while starch and protein were analysed photometrically. Seeds accumulated high molecular weight compounds such as proteins and starch, whereas elaiosomes accumulated more easily digestible low molecular weight compounds such as amino acids and monosaccharides. Analysis of similarities and similarity percentages analysis demonstrated that the composition of fatty acids, free amino acids and carbohydrates differed markedly between elaiosomes and seeds. The most important difference was in total amino acid content, which was on average 7.5 times higher in elaiosomes than in seeds. The difference was especially marked for the nitrogen-rich amino acid histidine. The availability of essential nutrients and, in some species, the higher nitrogen content in elaiosomes suggest that their nutritional value for larvae plays a key role in this interaction. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Effects of ants,ground beetles and the seed-fall patterns on myrmecochory of Erythronium japonicum Decne. (Liliaceae)

Erythronium japonicum (Liliaceae) inhabits deciduous mesic forests of Hokkaido, northern Japan. Myrmecochory of this species was investigated, especially the dispersal frequency, the effect of seed predators and the seed fall pattern. In the quadrat census using marked seeds of E. japonicum, the ant Myrmica kotokui frequently transported the seeds. However, the frequency of seed removal was low and most seeds were dispersed as little as 1 m or less. The spatial distribution of E. japonicum individuals was nearly random and most seedlings were established 5–20 cm away from the fertile plants, indicating that even this small scale of seed dispersal contributes to avoiding crowding of seedlings. Some arthropods, e.g. springtails, spiders and ticks, hindered seed dispersal by devouring elaiosomes and seeds. Although ground beetle species also damaged seeds and elaiosomes, a few of them exhibited seed removal behaviour. E. japonicum dropped their seeds not all at once but bit by bit, taking 3–6 days to drop all seeds. This seed-fall pattern was effective in raising the frequency of seed removal by ants and reducing seed predation by some arthropods.     

Uncoupling the effects of seed predation and seed dispersal by granivorous ants on plant population dynamics

Secondary seed dispersal is an important plant-animal interaction, which is central to understanding plant population and community dynamics. Very little information is still available on the effects of dispersal on plant demography and, particularly, for ant-seed dispersal interactions. As many other interactions, seed dispersal by animals involves costs (seed predation) and benefits (seed dispersal), the balance of which determines the outcome of the interaction. Separate quantification of each of them is essential in order to understand the effects of this interaction. To address this issue, we have successfully separated and analyzed the costs and benefits of seed dispersal by seed-harvesting ants on the plant population dynamics of three shrub species with different traits. To that aim a stochastic, spatially-explicit individually-based simulation model has been implemented based on actual data sets. The results from our simulation model agree with theoretical models of plant response dependent on seed dispersal, for one plant species, and ant-mediated seed predation, for another one. In these cases, model predictions were close to the observed values at field. Nonetheless, these ecological processes did not affect in anyway a third species, for which the model predictions were far from the observed values. This indicates that the balance between costs and benefits associated to secondary seed dispersal is clearly related to specific traits. This study is one of the first works that analyze tradeoffs of secondary seed dispersal on plant population dynamics, by disentangling the effects of related costs and benefits. We suggest analyzing the effects of interactions on population dynamics as opposed to merely analyzing the partners and their interaction strength.     

Effects of management and adjacent forest on the heathland bryophyte layer

Heathlands are severely threatened by decreasing habitat quantity and quality. In these habitats, bryophytes form an important component of plant diversity. Therefore, the aims of this study were to investigate the effects of management and adjacent forest land use on the heathland bryophyte layer. Furthermore, effects of these factors on invasive bryophyte species were studied.

Bryophyte communities of 11 dry heathland patches bordering forest were surveyed. In these heathlands, management turned out to be an important factor explaining community composition of bryophytes. Furthermore, significant effects of adjacent land use on bryophyte community composition and species richness were found. However, these effects were rather limited in extent, occurring only close to habitat borders, i.e., up to maximum 5 m into the heathland patch. Also for individual bryophyte species, effects of adjacent forest could only be observed within maximum 5 m from the edge. One remarkable edge effect was the increased dominance of the invasive species Campylopus introflexus near the forest edge, especially at grazed sites. Effects of adjacent land use seemed to predominate management effects. Consequently, management cannot serve as a tool to mitigate edge effects on bryophyte species in heathland patches.     

Soil CO2 efflux in uneven-aged managed forests: temporal patterns following harvest and effects of edaphic heterogeneity

Increased nutrient availability can have a large impact in Australian woodland ecosystems, many of which are very poor in nutrients, particularly phosphorus. A study was conducted in an urban Banksia woodland remnant in Perth, southwest Western Australia to test the hypothesis that the soil nutrient status in areas in good condition (GC), poor condition invaded by the perennial grass Ehrharta calycina (PCe), and poor condition invaded by the perennial herb Pelargonium capitatum (PCp), is reflected in the nutrient status of the native and introduced species. Leaf concentrations of P, K, N, Na, Ca, Mg, S, B, Cu, Fe, Mn and Zn of six native (Banksia attenuata, Banksia menziesii, Allocasuarina humilis, Melaleuca systena, Macrozamia fraseri and Conostylis aculeata) and four introduced species (Ehrharta calycina, Pelargonium capitatum, Gladiolus caryophyllaceus and Briza maxima), were measured. Soil pH, electrical conductivity, N (total), P (total), available P, K, S and organic C were assessed beneath all species on all sites. Significantly higher concentrations of soil P (total) and P (HCO₃) were found at PCe and PCp sites than GC sites, while PCp sites also had significantly higher soil concentrations for N (total) and S. Principal Components Analysis of the leaf analyses showed (a) individual species have characteristically different nutrient concentrations; (b) the introduced species Ehrharta calycina and Pelargonium capitatum clustered separately from each other and by vegetation condition. Leaf concentrations of P were significantly (P<0.05) higher, and K and Cu were significantly lower in PCe and PCp sites compared with those at GC sites. Introduced species leaf nutrient concentrations were significantly greater than native species for all nutrients except Mn which was significantly lower, with no differences for Mg and B. The results indicate a key role for P in the Banksia woodland, and we conclude that higher levels of available P at invaded sites are having a detrimental impact on the ecosystem. These results provide new knowledge to enhance conservation practices for the management of the key threatening process of invasion within a biodiversity hot spot. Section Editor: T. Kalapos     

Influence of litter removal and mineral soil disturbance on the spread of an invasive grass in a Central Hardwood forest

Soil and litter disturbances within Central Hardwood forests may facilitate exotic plant species invasion of interior forest areas. Microstegium vimineum is an annual exotic grass that has become common throughout the Southeastern United States. Three replicates of three different mineral soil and litter disturbance treatments, plus a control with no disturbance, were established on the leading edge of M. vimineum patches prior to seed fall. All patches were located in areas with similar forest canopy structure and slope in three Central Hardwood forest stands prior to seed fall. At the beginning of the following growing season, each individual M. vimineum seedling was mapped within the treatment plots. The mean number of M. vimineum individuals that established within each treatment did not differ significantly from the control. The distance at which 90% of the individuals had spread, and the overall mean distance spread were significantly farther for the litter removal treatment than the control. The farthest individual seedling from the boundary of existing patches in both the litter removal and the mineral soil disturbance and litter removal treatments were significantly farther than the control. The individuals that spread the farthest are of most concern due to the large number of viable seed that a single M. vimineum plant can produce. These results suggest that disturbance of the litter layer may increase the spread rate of M. vimineum and facilitate its invasion of new habitats, and that leaving litter layers intact may slow the spread of M. vimineum.     

Composition of soil seed banks in southern California coastal sage scrub and adjacent exotic grassland

Soil seed banks are important to many plant communities and are recognized as an important component of management plans. Understanding seed bank composition and density is especially important when communities have been invaded by exotic species and must be managed to promote desirable species. We examined germinable soil seed banks in southern California coastal sage scrub (CSS) that is heavily invaded by exotic grasses and in adjacent exotic grassland. Soils from both communities had similar seed banks, dominated by high densities of exotic grass and forb species. Up to 4,000 exotic grass seeds and at least 400 exotic forb seeds/m² were found in most soils, regardless of aboveground vegetation type. Native forbs averaged 400 seeds/m² in grass-dominated areas and about 800 in shrub-dominated soils. Shrub seed density was <1 and <10 seeds/m² in grass- and shrub-dominated areas, respectively, indicating that the shrub seed bank is not persistent compared to annuals. We also compared pre- and post-burn soil seed banks from one location that burned in October 2003. Late-season burning in both grass- and CSS-dominated areas disproportionately reduced exotic grass seed densities relative to native seed densities. The similarity of the seed banks in adjacent grass and shrub communities suggests that without intervention, areas currently dominated by CSS may become more similar to grass-dominated areas in terms of aboveground vegetation. In such areas, the first growing season following a wildfire is a window of opportunity for increasing native diversity at a time when density of exotic grass seeds is low. At time of research, Robert D. Cox was graduate student.     

Extensive contiguous north–south range expansion of the original population of an invasive lizard in Florida

Florida has one of the two worst non-indigenous invasive species problems in the United States, and all such species are considered by statute to be a potential ecological problem. Unfortunately, little information is available about most of Florida's invasive species. Here, we provide information on range expansion of a population of one of the lesser-known species. The Palm Beach County, Florida, population of the northern curly-tailed lizard, Leiocephalus carinatus armouri, was examined for north–south expansion of its previously documented range. Observations were conducted as contiguous expansion outward from the previously known range. We found a substantial contiguous range expansion by a minimum of 46.3 km to the south and 34.1 km to the north. This species appears to be closely associated with coastal habitat degradation in the form of human infrastructure, especially ageing pavement, sea walls, buildings, and bridge bases.     

Positive feedbacks between plant invasions and fire regimes: <Emphasis Type=Italic>Teline monspessulana</Emphasis> (L.) K. Koch (Fabaceae) in central Chile

Invasive species can increase fire frequency and intensity, generating favorable conditions for their self-perpetuation. Mediterranean south-central Chile may be especially prone to the effects of invasive species on fire regimes because it is less adapted to fire and it contains a highly endemic flora. Teline monspessulana (L.) K. Koch (syn. Cytisus monspessulanus L.; Genista monspessulana (L.) L.A.S. Johnson) is an introduced shrub that forms monotypic stands or is present as an understory species in native forests as well as in forestry plantations. Dense T. monspessulana stands are completely destroyed by fire, generating the conditions for it seeds to germinate and establish an abundant regeneration, with up to 900 plants/m². We report key evidence on abundance and biomass in adult stands, and patterns of seed bank and regeneration after fire in stands of T. monspessulana around the city of Concepción, Chile. We estimated living biomass in pure stands and underneath Eucalyptus plantations. In burned areas, we assessed T. monspessulana seed bank and studied regeneration patterns. We found that T. monspessulana densities reaches 52,778 plants/ha and 8.92 ton/ha in pure stands and 34,223 plants/ha and 2.31 ton/ha underneath Eucalyptus plantations. T. monspessulana generates small caliper fuel and acts as a ladder-fuel. Large soil seed banks allow for abundant regeneration after fire, with mean densities of 877,111 plants/ha, but an overall mortality of 37.2% in the first year after the fire. The high values of regeneration compared to final densities in adult stands suggest that density-dependent mortality. Our results indicate that T. monspessulana regeneration is not only favored by fires, but also that the species creates favorable conditions for intense and continuous fires, both under pure conditions, but also associated to exotic tree plantations. To understand the implications of positive feedbacks between invaders and fire, we recommend focusing in the mechanisms by which they increases fuel accumulation and fuel flammability, and how higher fire frequency and intensity favors invasive species recruitment over native species. Comprehension of this dynamics will allow for better management and control of these invasions which have major ecological, economical and social implications.     

The role of the permanent soil seed bank in early stages of a post-agricultural succession in the Inland Pampa,Argentina

We studied the soil seed bank composition in four old fields of different ages, after abandonment from agriculture. Complete seed bank composition was assessed by direct seed separation from soil samples and identification to species. Most species found in the seed bank were not important in the present seral communities. Seed of the species that dominated the early succession were generally not found. Additionally, there were very few propagules rather than on the germination of in situ propagules. We suggest that pampean grasses evolved under that the course of post-agricultural succession will depend strongly on the pattern of arrival of exogenous propagules rather than the germination of in situ propagules. We suggest that pampean grasses evolved under disturbances of low intensity and/or a disturbance regime dominated by small gaps, in which open areas could be rapidly colonized from the edges and/or by remnant vegetative propagules. The changes produced by the introduction of agriculture triggered the invasion by exotic species adapted to the new disturbance regime.     

Size and lipid content of nonmyrmecochorous diaspores: effects on the interaction with litter-foraging ants in the Atlantic rain forest of Brazil

Ants are often attracted to diaspores not adapted for dispersal by ants. These diaspores may occasionally benefit from this interaction. We selected six nonmyrmecochorous plant species (Virola oleifera, Eugenia stictosepala, Cabralea canjerana, Citharexylum myrianthum, Alchornea glandulosa and Hyeronima alchorneoides) whose diaspores differ in size and lipid content, and investigated how these features affect the outcome of ant-diaspore interactions on the floor of a lowland Atlantic forest of Southeast Brazil. A total of 23 ant species were seen interacting with diaspores on the forest floor. Ants were generally rapid at discovering and cleaning the diaspore pulp or aril. Recruitment rate and ant attendance were higher for lipid-rich diaspores than for lipid-poor ones. Removal rate and displacement distance were higher for small diaspores. The large ponerine ant Pachycondyla striata, one of the most frequent attendants to lipid-rich arillate diaspores, transported the latter into their nests and discarded clean intact seeds on refuse piles outside the nest. Germination tests with cleaned and uncleaned diaspores revealed that the removal of pulp or aril may increase germination success in Virola oleifera, Cabralea canjerana, Citharexylum myrianthum and Alchornea glandulosa. Gas chromatography analyses revealed a close similarity in the fatty acid composition of the arils of the lipid-rich diaspores and the elaiosome of a typical myrmecochorous seed (Ricinus communis), corroborating the suggestion that some arils and elaiosomes are chemically similar. Although ant-derived benefits to diaspores – secondary dispersal and/or increased germination – varied among the six plant species studied, the results enhanced the role of ant-diaspore interactions in the post-dispersal fates of nonmyrmecochorous seeds in tropical forests. The size and the lipid-content of the diaspores were shown to be major determinants of the outcome of such interactions.     

Diversity of understory plants in undisturbed and disturbed tropical lowland forests of Little Andaman Island, India

Species richness and density of understory plants were investigated in eight 1 ha plots, distributed one each in undisturbed and disturbed tropical evergreen, semi-evergreen, deciduous and littoral forests of Little Andaman island, India, which falls under one of the eight hottest hotspots of Biodiversity in the world viz. the Indo-Burma. One hundred 1 m⁻² quadrats were established in each 1 ha plot, in which all the understory plants (that include herbs, undershrubs, shrubs and herbaceous climbers) were enumerated. The total density of understory plants was 6,812 individuals (851 ha⁻¹) and species richness was 108 species, representing 104 genera and 50 families. Across the four forest types and eight study plots, the species richness ranged from 10 to 39 species ha⁻¹. All the disturbed sites harbored greater number of species than their undisturbed counterparts. Herbs dominated by species (63%) and density (4,259 individuals). The grass Eragrostis tenella (1,860 individuals; IVI 40), the invasive climber Mikania cordata (803; IVI 20) and the shrub Anaxagorea luzonensis (481; IVI 17.5) were the most abundant species. Poaceae, Asteraceae, Acanthaceae, Orchidaceae and Euphorbiaceae constituted the species-rich families represented by 6 species each. The species-area curves attained an asymptote at 0.8 ha level except in sites DD and DL, indicating 1 ha plot is not sufficient to capture all the understory species in disturbed forests. The alien weeds formed about one-fourth of the species richness (31 species; 28%) and density (1,926 individuals; 28.3%) in the study sites, indicating the extent of weed invasion and the attention required for effective conservation of the native biodiversity of the fragile island forest ecosystem.     

Landslides significantly alter land cover and the distribution of biomass: an example from the Ninole ridges of Hawai'i

In the Ninole ridges of Hawai'i we investigated how landslides influence ecosystem development and modify land cover and the distribution of biomass. We estimated above and below-ground biomass, and N and P concentration in leaves (Metrosideros polymorpha) and very fine roots (all species), for vegetation developing on landslides of three age classes (young, < 18 yr; intermediate, 42 yr; and old ca. 124 yr) and on undisturbed soils (ca. 430 yr). The undisturbed soils were derived from ash underlain by basalt. To quantify changes in land cover and the distribution of biomass we combined our estimates of biomass with estimates of the area covered by each vegetation class. The latter estimates were obtained from the analysis and classification of color-infrared aerial photographs. Average above- and below-ground biomass for the herbaceous vegetation (young landslides) was 10.4 and 3.2 t/ha, whereas for the ohia-non ash forest (intermediate and old landslides) was 37.5 and 5.2 t/ha, respectively. For the ohia-ash forest (undisturbed sites), average above and below-ground biomass was 354.6 and 9.5 t/ha, respectively. Average foliar N for the herbaceous and ohia-non ash forest ranged between 0.80–0.84%, whereas root P between 0.056–0.040%, respectively. For the ohia-ash forest, average foliar and root P was 0.918% and 0.036%, respectively. Based on changes in vegetation cover during the last 430 yr, we estimated rate of disturbance at 15% per century or equivalently that 53 t/ha biomass per century exited through the system. The removal of ash-derived soils by landslides significantly alters successional trajectories and by doing so may be transforming the Ninole ecosystems in irreversible ways.