Myrmecochory in Australia's seasonal tropics: Effects of disturbance on distance dispersal

Abstract Successful ecosystem restoration requires the re-establishment of fundamental ecological processes, many of which involve plant-animal interactions. Myrmecochory (seed dispersal by ants) is a particularly important plant-animal mutualism in Australia, but little is known about its response to either disturbance or restoration following disturbance. Here we investigate the effects of disturbance on seed dispersal by ants, and the extent to which the ant-seed relationship has re-established at sites undergoing rehabilitation, at Ranger uranium mine in the seasonal tropics of Australia's Northern Territory. We focused on the composition of seed-dispersing ant assemblages, rates of seed removal by ants, and the dispersal curves generated by ants, as determined by observations of removal from seed depots. Ten sites were studied, comprising four ‘natural’ (undisturbed) sites representing a range of savanna habitats occurring in the region, four disturbed sites representing a range of habitat disturbance but with intact soil, and two waste rock sites subject to preliminary revegetation trials. A total of 22 ant species from 10 genera were observed during 154 observations of seed removal, most commonly Rhytidoponera aurata (53 records), Monomorium (rothsteini gp) sp. 1 (14), Iridomyrmex sanguineus (13), Iridomyrmex pallidus (12) and Pleidole sp. 3 (10). Removal rates (over 3 h) averaged 29% across all sites and time periods, varying markedly both between and within sites. However, mean rates of removal were similar between natural, disturbed and waste rock sites (29%, 28% and 31%, respectively). A high incidence (62% of all depots) of'aril robbing’ by ants (primarily Monomorium spp.) eating arils in situ, without removal, was observed. Dispersal distances varied markedly between ant species, with Iridomyrmex sanguineus having both the highest mean (7.25m) and maximum (13.08 m) dispersal distances. Species of Pheidole typically dispersed seeds less than 0.5 m, and Meranoplus, Monomorium and Tetramorium spp. only ever moved seeds a few centimetres, usually dropping and abandoning them before reaching the nests. The dispersal curves characteristic of each site varied markedly due to the different composition of seed-dispersing ants. The mean dispersal distance at disturbed sites (3.91 m) was significantly higher than at natural sites (2.19 m), and the curves were strongly skewed in the former, but relatively uniform in the latter. The implications of these differences for recovery following disturbance are unclear. At rehabilitated waste rock sites, all observed removals involved distances less than 0.5 m, with a mean of 17 cm. This lack of effective ant-seed relationships might represent a barrier to further vegetation development at rehabilitated sites.     

Mutualism between co-introduced species facilitates invasion and alters plant community structure

Generalized mutualisms are often predicted to be resilient to changes in partner identity. Variation in mutualism-related traits between native and invasive species however, can exacerbate the spread of invasive species (‘invasional meltdown’) if invasive partners strongly interact. Here we show how invasion by a seed-dispersing ant (Myrmica rubra) promotes recruitment of a co-introduced invasive over native ant-dispersed (myrmecochorous) plants. We created experimental communities of invasive (M. rubra) or native ants (Aphaenogaster rudis) and invasive and native plants and measured seed dispersal and plant recruitment. In our mesocosms, and in laboratory and field trials, M. rubra acted as a superior seed disperser relative to the native ant. By contrast, previous studies have found that invasive ants are often poor seed dispersers compared with native ants. Despite belonging to the same behavioural guild, seed-dispersing ants were not functionally redundant. Instead, native and invasive ants had strongly divergent effects on plant communities: the invasive plant dominated in the presence of the invasive ant and the native plants dominated in the presence of the native ant. Community changes were not due to preferences for coevolved partners: variation in functional traits of linked partners drove differences. Here, we show that strongly interacting introduced mutualists can be major drivers of ecological change.     

Soil disturbance effects on the composition of seed-dispersing ants in roadside environments

Myrmecochory (the dispersal of seeds by ants) is a significant ecological process in sclerophyll woodlands, but habitat disturbance is known to alter the extent and success of this mutualism. We investigated the influence of soil disturbance on the composition of the seed-dispersing ant community. Surveys were conducted in roadside verges where soils are regularly disturbed by road maintenance activities. Using a ‘cafeteria’ bait station approach, we selected 24 roads of different widths to investigate ant composition and abundance in relation to soil disturbance. We found ant species richness was greater in non-disturbed than disturbed zones, where road verge width significantly influenced results. The composition and abundance of individual seed-dispersing ant species varied between disturbed and non-disturbed zones. Rhytidoponera metallica were more abundant in non-disturbed sites, whereas Melophorus bruneus and Monomorium rothseini were more frequently recorded in disturbed areas. Commonly found Iridomyrmex purpureus was significantly more abundant in disturbed zones in narrow roadsides and vice versa in wide roadsides, and strongly influenced total community composition. Variation in the abundance of commonly recorded Iridomyrmex and Monomorium genera were related more to site conditions (roadside width and habitat) than soil disturbance. The rich composition of seed dispersing ants in roadside environments, and the effects of soil disturbances on these ant communities that we describe, provide a key insight to important seed dispersal vectors occurring in fragmented rural landscapes.     

Seed bank,seed size and dispersal in moisture gradients of temporary pools in Southern France

Plants reduce risk of extinction due to unpredictable rainfall by soil seed banks, dispersal or large seeds. However, seed size also increases independently in dry habitats, and since seed size is in a trade-off with seed number, size of seed banks is expected to increase in moister habitats. Therefore, we wanted to test if seed abundance in soil increases in wet habitats, if seed size increases in dry habitats, and if spread of seeds along the gradient is higher for plants of intermediate habitats in local moisture gradients.We studied 15 temporary pools in three biogeographically separated wetlands in Southern France. For each pool we studied five different moisture levels, totalling 75 local plant communities. We quantified soil seed bank by the seedling emergence method, seed size and an index of spatial spread of seeds in the soil for every species. We also quantified water levels for each plot.We found increasing abundance of seeds in the soil with increasing water levels but lower seed size and higher spread at intermediate water levels. When we controlled for niche position, we found no trade-off between seed size, spread and abundance in the soil seed bank.Type and importance of risk reduction strategies thus appeared to be strongly driven by the plant species’ moisture niche and the spatial arrangement of water levels.     

Variegated desert vegetation: Covariation of edaphic and fire variables provides a framework for understanding mulga‐spinifex coexistence

Abstract Mulga (Acacia aneura Mimosaceae) and spinifex (Triodia spp. Poaceae) habitats together characterize a large part of arid central Australia. Often very abrupt boundaries form between these two habitats, giving rise to a mosaic pattern of contrasting shrub‐grass alterations across the landscape. Reasons for such patterning remain poorly understood though current niche‐based views relate species' distributions to spatial resource gradients or to fire effects. Field survey work was conducted on central Australian mountain ranges to further quantify floristic, regeneration traits, and structural patterning across mulga‐spinifex transitions and to test resource‐ and disturbance‐models that explain these patterns. Compositional analysis demonstrated variability in transition type – in certain cases boundaries denoted true floristic discontinuity and in others, somewhat more of a structural shift. Moreover, it was shown that minimal between‐habitat floristic overlap coincided with the occurrence of distinct edaphic changes, while greater compositional commonality occurred when soil gradients were more diffuse. This indicated that floristic patterning cannot be ascribed to any one single process. In the case of strong soil gradients, between‐habitat segregation most likely resulted from resource‐based niche differentiation; for weaker gradients, fire‐frequency assumed greatest importance. Disturbance theory most readily accounted for the distribution of woody species' post‐fire regeneration traits across habitat boundaries. The results also suggested that biotic factors –viz competition, facilitation and animal‐mediated dispersal – may be of additional consequence for mulga‐spinifex coexistence. Overall, the study served to emphasize the importance of multi‐factor explanation for within‐ and between‐habitat patterning in these mosaics. It also highlighted the need for experimentation to facilitate distinction between cause and correlation.     

Effect of neighboring seeds associated with habitats on the seed removal of two coexisting myrmecochorous plants in central China

Although neighboring plants can influence animal-seed dispersal interactions, little is known about the effect of neighboring seeds and the influence of habitat on seed dispersal by ants. Here we investigated the influence of neighboring seeds on seed removal in two coexisting myrmecochorous species (Epimedium pubescens and Helleborus thibetanus) from temperate deciduous forests of Qinling Mountains, central China, by examining (1) the potential role of ants and rodents and (2) whether the neighboring seed effect differed between forest edge and interior. We found that, presence of the higher-ESMR (elaiosome: seed mass ratio) E. pubescens did not significantly affect seed removal of the lower-ESMR H. thibetanus. By contrast, the presence of H. thibetanus decreased the seed removal rates of E. pubescens, with only a significant effect in rodent exclusion (ant alone) rather than in both ant exclusion (rodent alone) and full access (rodent + ant). Moreover, we found that those effects were not significantly different between the forest edge and the interior, which may be attributed to a similar pattern of overall seed-dispersing ant abundance in the two habitats. This suggested that neighboring seeds could influence seed removal of the focal plant depending on the absence of rodents; when rodents were present, the interaction of rodents and ants rendered no influence of neighboring seeds on seed removal. Our results show that the neighborhood effect was regulated by both dispersers and predators, and this effect was not context-dependent at a small spatial scale. This study highlights the importance to understand the effect of shared seed-removing animals and habitat context to assess the neighboring seed effect on plant-animal interactions.     

Response of ants to human‐altered habitats with reference to seed dispersal of the myrmecochore Corydalis giraldii Fedde (Papaveraceae)

Anthropogenic habitat disturbance has potential consequences for ant communities. However, there is limited information on the effects of ant responses on associated ecological processes such as seed dispersal. We investigated the effect of disturbance on the abundance, richness, and composition of ant communities and the resulting seed‐dispersal services for a herbaceous myrmecochore, Corydalis giraldii (Papaveraceae), in an undisturbed habitat (forest understory), moderately disturbed habitat (abandoned arable field), and highly disturbed habitat (road verge) on Qinling Mountains, China. In total, we recorded 13 ant species, and five out of these were observed to transport seeds. The community composition of dispersers was significantly different amongst habitats. The richness of the dispersers did not differ among the habitats, but their total abundance varied significantly across habitats and was 21% lower in the road verge than in the abandoned arable fields. The major seed‐dispersing ant species in both the forest understory and the abandoned arable field were large‐bodied (Myrmica sp. and Formica fusca, respectively), whereas the major seed‐dispersing ants found in the road verge were the small‐bodied Lasius alienus. This difference resulted in lower seed removal rates and dispersal distances in the road verge than in the other two habitats. The different dispersal patterns were attributed primarily to differences in dispersing ant abundance and identity, most likely in response to habitats with different degree of anthropogenic disturbance. The possible influence of disturbance on the ecological specialization of ant‐seed dispersal interaction is also discussed.     

HOW DOES POLLEN VERSUS SEED DISPERSAL AFFECT NICHE EVOLUTION?

In heterogeneous landscapes, the genetic and demographic consequences of dispersal influence the evolution of niche width. Unless pollen is limiting, pollen dispersal does not contribute directly to population growth. However, by disrupting local adaptation, it indirectly affects population dynamics. We compare the effect of pollen versus seed dispersal on the evolution of niche width in heterogeneous habitats, explicitly considering the feedback between maladaptation and demography. We consider two scenarios: the secondary contact of two subpopulations, in distinct, formerly isolated habitats, and the colonization of an empty habitat with dispersal between the new and ancestral habitat. With an analytical model, we identify critical levels of genetic variance leading to niche contraction (secondary contact scenario), or expansion (new habitat scenario). We confront these predictions with simulations where the genetic variance freely evolves. Niche contraction occurs when habitats are very different. It is faster as total gene flow increases or as pollen predominates in overall gene flow. Niche expansion occurs when habitat heterogeneity is not too high. Seed dispersal accelerates it, whereas pollen dispersal tends to retard it. In both scenarios very high seed dispersal leads to extinction. Overall, our results predict a wider niche for species dispersing seeds more than pollen.     

吉林西部草原地区蚂蚁种类及分布

在吉林西部草原地区蚂蚁是大型土壤动物的优势类群,数量巨大,但种类少,仅发现3亚科9属14种。放牧场、撂荒地、农田、人工林、乡间土路、院落6种生境中分布最多的蚂蚁是红林蚁和铺道蚁,撂荒地和人工林的蚂蚁种类和巢口密度最大。在割草场分布的蚂蚁主要是玉米毛蚁、黄墩蚁和铺道蚁,它们均修建明显的地上蚁丘,蚁丘呈环带状分布,即主要分布在草地与碱斑的交界处,并且从草场边缘向中心,蚁丘的数量逐渐减少。利用拥挤度指数对蚁丘的分布格局进行分析,表明蚁丘呈聚集分布,并对可能影响蚁巢分布的因素进行了探讨。     

Structure and succession of ant assemblages in a north European sand dune area

Ant assemblages of a successional dune area (Tvärminne, south Finland) were analysed at three levels: the distribution of species among habitats, the distribution of colonies within habitats and the spatio-temporal distribution of individuals. The distribution of ant species among habitat patches representing different stages of succession is correlated with the composition of the epigeic fauna, the number and condition of dead twigs on the ground and the vegetation architecture. The composition of ant assemblages seems relatively independent of the plant species composition, and there are uncoordinated successional steps between the vegetation and ant communities.
In the early successional stages interference competition is weak and insignificant, as indicated by the random spatial arrangement of colonies, the absence of postcompetitive niche segregation and the low encounter rates of ant workers on the ground and at baits. In the aggressive behavioural hierarchy, Formica cinerea is a submissive species as compared with either Formica sanguinea or Lasius alienus.     

Ant-nest soil and seedling growth in a neotropical ant-dispersed herb

Summary A major hypothesis concerning the benefits of myrmecochory, seed dispersal by ants, to plants is that ant nests are nutrient-enriched microsites that are beneficial to seedling growth. We experimentally test this hypothesis for a neotropical myrmecochore, Calathea ovandensis, asking two questions: 1) is soil of nests of a seed-dispersing ant chemically or structurally distinct from surrounding soils, and 2) do seedlings grow better in soil collected from ant nests than in randomly collected soil? We found that although ant-nest soil was significantly enriched in nitrate-nitrogen, magnesium, iron, manganese, cadmium and percent organic matter compared to randomly collected soil, seedling growth was not significantly improved by ant-nest soil.     

Strong plant-soil associations in a heterogeneous subtropical broad-leaved forest

The relative importance of deterministic and neutral processes on community assembly is currently a topic of much debate among ecologists. Analyzing species-environment associations is an effective way to assess the importance of deterministic process such as niche differentiation, but both habitat association and dispersal limitation can produce similar patterns of spatial aggregation in species. Therefore, it is crucial to control for the impact of dispersal limitation on species distributions when analyzing species-environment associations. We sampled soil with high resolutions in a 24 ha stem-mapped subtropical forest and tested plant-soil associations. We controlled for the influence of dispersal limitation by employing the homogeneous Thomas process to simulate the effect of dispersal limitation on the aggregation of tree species. After controlling for the effect of dispersal limitation, we found that the spatial heterogeneity of soil properties was associated with distributions of 88.2% (90 of 102 species) of tree species in this subtropical forest. Furthermore, not only did soil properties influence the distribution of tree species, but also tree species tended to affect properties of the soil around them. The soil factors most strongly influencing species distributions were TC, TN, TP, K, Mg, Si, soil moisture, and bulk density. We found the spatial heterogeneity of soil properties to be strongly associated with tree species distributions. Niche partitioning of soil gradients contributed substantially to species coexistence in this subtropical forest.     

Argentine ant invasion associated with loblolly pines in the southeastern United States: minimal impacts but seasonally sustained

Invasive ants are notorious for directly displacing native ant species. Although such impacts are associated with Argentine ant invasions (Linepithema humile) worldwide, impacts within natural habitat are less widely reported, particularly those affecting arboreal ant communities. Argentine ants were detected in North Carolina mixed pine-hardwood forest for the first time but were localized on and around loblolly pines (Pinus taeda), probably because of association with honeydew-producing Hemiptera. We explored the potential impacts of L. humile on arboreal and ground-foraging native ant species by comparing interspersed loblolly pines invaded and uninvaded by Argentine ants. Impacts on native ants were assessed monthly over 1 yr by counting ants in foraging trails on pine trunks and in surrounding plots using a concentric arrangement of pitfall traps at 1, 2, and 3 m from the base of each tree. Of floristics and habitat variables, higher soil moisture in invaded plots was the only difference between plot types, increasing confidence that any ant community differences were caused by Argentine ants. Overall patterns of impact were weak. Composition differed significantly between Argentine ant invaded and uninvaded trunks and pitfalls but was driven only by the presence of Argentine ants rather than any resulting compositional change in native ant species. Native ant abundance and richness were similarly unaffected by L. humile. However, the abundance of individual ant species was more variable. Although numbers of the arboreal Crematogaster ashmeadi (Myrmicinae) declined on and around invaded pines, epigeic Aphaenogaster rudis (Myrmicinae) remained the most abundant species in all plots. Argentine ant densities peaked in late summer and fall, therefore overlapping with most native ants. Unexpected was their continued presence during even the coldest months. We provide evidence that Argentine ants can invade and persist in native North Carolina forests, probably mediated by pine-associated resources. However, their localized distribution and minimal impact on the native ant fauna relative to previously described invasions requires further resolution.     

Forest edges and fire ants alter the seed shadow of an ant-dispersed plant

Exotic species invade fragmented, edge-rich habitats readily, yet the distinct impacts of habitat edges and invaders on native biota are rarely distinguished. Both appear detrimental to ant-dispersed plants such as bloodroot, Sanguinaria canadensis. Working in northeastern Georgia (USA), an area characterized by a rich ant-dispersed flora, fragmented forests, and invasions by the red imported fire ant, Solenopsis invicta , I monitored the interactions between ants and S. canadensis seeds in uninvaded forest interiors, uninvaded forest edges, invaded forest interiors, and invaded forest edges. I observed 95% of the seed dispersal events that occurred within the 60-min observation intervals. Seed collection rates were similar among all four (habitat × invasion) groups. The presence of invasive ants had a strong effect on seed dispersal distance: S. invicta collected most seeds in invaded sites, but was a poorer disperser than four of five native ant taxa. Habitat type (interior versus edge) had no effect on seed dispersal distance, but it had a strong effect on seed dispersal direction. Dispersal towards the edge was disproportionately rare in uninvaded forest edges, and ants in those habitats moved the average dispersed seed approximately 70 cm away from that edge. Dispersal direction was also skewed away from the edge in uninvaded forest interiors and invaded forest edges, albeit non-significantly. This biased dispersal may help explain the rarity of myrmecochorous plants in younger forests and edges, and their poor ability to disperse between fragments. This is the first demonstration that forest edges and S. invicta invasion influence seed dispersal destination and distance, respectively. These forces act independently.     

Niche properties of Central European spiders: shading, moisture and the evolution of the habitat niche

Aim  Niche theory emphasizes the importance of environmental conditions for the distribution and abundance of species. Using a macroecological approach our study aimed at identifying the important environmental gradients for spiders. We generated numerical values of niche position and niche width. We also investigated relationships between these niche properties as well as the degree of phylogenetic conservatism in order to draw conclusions about the evolution of the habitat niche.
Location  Central Europe: lowlands of Austria, Belgium, Germany, Luxembourg, the Netherlands and Switzerland.
Methods  We analysed 244 published spider communities from 70 habitat types by correspondence analysis. The resulting community scores were used to test for correlations with habitat characteristics. Species scores were used to derive niche position (mean scores) and niche width (standard deviation of scores). To test for niche conservatism we estimated variance components across the taxonomic hierarchy.
Results  The first two axes of the correspondence analysis were correlated with shading and moisture, respectively. Niche width had a hump-shaped relationship to both environmental gradients. β-diversity was strikingly higher in open habitats than in forests. Habitat niche conservatism was lower than phylogenetic conservatism in body size.
Main conclusions  Environmental factors are important drivers for the β-diversity of spiders, especially across open habitats. This underlines the importance of preserving the whole range of moisture conditions in open habitats. Narrow niches of species occurring at the ends of both environmental gradients indicate that adaptations to extreme habitats lead to constraints in ecological flexibility. Nevertheless, the habitat niche of species seems to evolve much faster than morphological or physiological traits.     

Functional structure of ant and termite assemblages in old growth forest,logged forest and oil palm plantation in Malaysian Borneo

Forested tropical landscapes around the world are being extensively logged and converted to agriculture, with serious consequences for biodiversity and potentially ecosystem functioning. Here we investigate associations between habitat disturbance and functional diversity of ants and termites—two numerically dominant and functionally important taxa in tropical rain forests that perform key roles in predation, decomposition, nutrient cycling and seed dispersal. We compared ant and termite occurrence and composition within standardised volumes of soil and dead wood in old growth forest, logged forest and oil palm plantation in Sabah, Malaysian Borneo. Termites occurred substantially less frequently in converted habitats than in old growth forest, whereas ant occurrences were highest in logged forest and lowest in old growth forest. All termite feeding groups had low occurrence in disturbed habitats, with soil feeders occurring even less frequently than wood feeders. Ant functional groups showed more variable associations, with some opportunist and behaviourally dominant groups being more abundant in degraded habitats. The importance of ants and termites in tropical ecosystems and such differing patterns of assemblage variation suggest that ecosystem functioning may be significantly altered in converted habitats.     

Perturbed partners: opposite responses of plant and animal mutualist guilds to inundation disturbances

Mutualists have been suggested to play an important role in the assembly of many plant and animal communities, but it is not clear how this depends on environmental factors. Do, for instance, natural disturbances increase or decrease the role of mutualism? We focused on entire guilds of mutualists, studying seed‐dispersing ants and ant‐dispersed plants along gradients of inundation disturbances. We first studied how abundance and richness of the mutualists, relative to non‐mutualists, change along 35 small‐scale gradients of inundation disturbances. We found that at disturbed sites, mutualistic plant species, those that reproduce by seeds dispersed by ants, increased in abundance and in consequences in richness, relative to other herbaceous plants. In contrast, we found that among the epigeic arthropods the abundance of mutualists declined, even more so than other arthropods. Correspondingly, distributions of plant and animal mutualists became increasingly discordant at disturbed sites: most plant mutualists were spatially separated from most animal mutualists. We finally found that high abundances of plant mutualists did not translate into a high nutrition service rendered to ants: at disturbed sites, many of the plants of ant‐dispersed species did not produce seeds, which coincided with a decline in seed dispersal by ants and a changing searching behavior of the ants. Overall, the small‐scale natural disturbances we studied were correlated to a major change in the assembly of mutualist guilds. However, the correlation was often opposite between interacting plant and animal mutualist guilds and may thus reduce the potential interaction between them.     

Seed dispersal in Erythronium dens-canis L. (Liliaceae): variation among habitats in a myrmecochorous plant

Erythronium dens-canis is a geophyte which produces a single flower each season. The fruits produce small seeds with relatively large elaiosomes. We performed experiments to investigate primary and secondary seed dispersal mechanisms of this species in different habitats in the western part of the Cantabrian Range in northwest Spain. Sticky traps were used to measure primary dispersal of seeds up to 0.5 m from mother plants. Seed cafeteria experiments were performed in different habitats to examine the role of ants and rodents in secondary seed transport and seed predation. Our results indicate that: (a) primary seed dispersal is positively skewed (99% of seeds fall within 20 cm of the mother plant) and seed dispersal distances vary significantly among plants; (b) secondary dispersal is exclusively by myrmecochory, although the proportion of seeds removed by ants differs significantly among habitats; (c) ant species composition and abundances vary among habitats; and (d) freshly dropped seeds are more likely to be removed than seeds that have begun to dry out. We conclude that secondary dispersal of seeds is greatly influenced by habitat but not by small-scale microhabitat. This revised version was published online in June 2006 with corrections to the Cover Date.     

Seed handling behaviours of native and invasive seed‐dispersing ants differentially influence seedling emergence in an introduced plant

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