Effects of experimental small-scale grassland fragmentation on spatial distribution, density, and persistence of ant nests

Abstract.  1. Grassland fragmentation is expected to influence the abundance of different invertebrate species to a different extent. Fragmentation-related effects are of particular importance in species that interact with many other species.
2. The density and spatial distribution of nests of 15 ant species in experimentally fragmented calcareous grasslands at three sites in the Northern Swiss Jura mountains were examined. Fragments of different size (0.25 m², 2.25 m², and 20.25 m²) were isolated by a 5-m wide strip of frequently mown vegetation. Control plots of corresponding size were situated in adjacent undisturbed grassland.
3. Three years after initiation of the experiment, ant nest density did not differ between fragments and control plots. Six years after initiation of the experiment, however, ant nest density and forager abundance were higher in large fragments than in large control plots. Ant nests tended to occur more frequently along the edge of fragments than in the core area. Persistence time of nests of the most abundant species, Lasius paralienus , tended to be shorter in fragments than in control plots. Furthermore, persistence time was longer in nests situated close to the fragment edge than in nests in the core area.
4. Effects on nest density, edge effects on the spatial distribution of nests, and the relationships between nest density and environmental factors were more pronounced when only nests of L. paralienus were considered. The implications of these findings for plant and other invertebrate species are discussed.     

Habitat fragmentation,EFN‐bearing trees and ant communities: Ecological cascades in Atlantic Forest of northeastern Brazil

Habitat fragmentation has a marked impact on the functional composition of tropical forest tree assemblages, and such change is likely to cascade through other trophic levels. Here, we investigate how habitat fragmentation affects extrafloral nectary (EFN)‐bearing plants and ant functional groups known to attend EFNs in a fragmented landscape of the Atlantic Forest. Extrafloral nectary‐bearing trees were identified in 50 0.1‐ha plots located in forest fragments, edge and interior patches. Ants were surveyed in 30 1‐m² litter samples in each of 17 forest fragments and in forest interior. Extrafloral nectary‐bearing plants accounted for 19.9% of individuals and 10.5% of species and included both pioneer and shade‐tolerant species similarly rich in the three habitat types. However, shade‐tolerant individuals accounted for >80% of EFN‐bearing plants in forest interior, compared with 2% in forest edge and 29% in fragments. Forest edge and fragment plots had a third fewer EFN‐bearing individuals and species compared with forest interior. This appeared to have important implications for local ant communities as the density of EFN‐bearing trees was the most important variable explaining the species richness of arboreal dominant ants. Our results show that plant loser–winner replacements promoted by forest fragmentation can cascade through higher trophic levels, with implications for forest dynamics and biodiversity conservation.     

Meat ants as dominant members of Australian ant communities: an experimental test of their influence on the foraging success and forager abundance of other species

Meat ants (Iridomyrmex purpureus and allies) are perceived to be dominant members of Australian ant communities because of their great abundance, high rates of activity, and extreme aggressiveness. Here we describe the first experimental test of their influence on other ant species, and one of the first experimental studies of the influence of a dominant species on any diverse ant community. The study was conducted at a 0.4 ha savanna woodland site in the seasonal tropics of northern Australia, where the northern meat ant (I. sanguineus) represented 41% of pitfall catches and 73% of all ants at tuna baits, despite a total of 74 species being recorded. Meat ants were fenced out of experimental plots in order to test their influence on the foraging success of other species, as measured by access to tuna baits. The numbers of all other ants and ant species at baits in exclusion plots were approximately double those in controls (controlling for both the fences and for meat ant abundance), and returned rapidly to control levels when fences were removed after 7 weeks. Individual species differend markedly in their response to the fencing treatment, with species of Camponotus and Monomorium showing the strongest responses. Fencing had no effect on pitfall catches of species other than the meat ant, indicating that the effect of meat ants at baits was directly due to interference with foraging workers, and not regulation of general forager abundance. Such interference by meat ants has important implications for the sizes and densities of colonies of other ant species, and ultimately on overall ant community structure.     

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 .     

Diversity and structure of ant communities associated with oak,pine, eucalyptus and arable habitats in Portugal

Summary In central Portugal, 28 species of ants were found in oak, 25 in pine, 35 in eucalyptus and 12 in arable habitats, a total of 43 species. They include five numerically dominant species, the exotic Argentine ant.Linepithema (Iridomyrmex) humile and four native species,Crematogaster scutellaris, Pheidole pallidula, Tapinoma nigerrimum andTetramorium hispanicum. L. humile occurred in 34% of sampled sites. When present, it was abundant in all three arboreal habitats, where it was associated with strikingly decreased ant species richness and equitability. Such diversity was greater inL. humile-colonised arable habitats, where the ant was much less abundant than in the arboreal habitats.WhereL. humile was absent,C. scutellaris andP. pallidula were usually co-dominants in oak and pine, andT. nigerrimum andT. hispanicum in the arable habitat, but all were absent or rare in eucalyptus. Ants were numerically least abundant in eucalyptus but, in the absence ofL. humile, species richness was greater and the community more equitable than in the other habitats.L. humile-absent arable habitats supported the poorest and least equitable ant community.     

Dissecting components of population-level variation in seed production and the evolution of masting behavior

Theory predicts that at higher trophic levels species are especially vulnerable to habitat fragmentation due to small population size and dependence on particular prey species. Using aphids as model organism, we tested the hypothesis that herbivore abundance increases in fragmented habitats as a result of reduced predator and parasitoid densities. In a 3 year-study, we examined the population dynamics of aphids with respect to host plant abundance and ant nest density in experimentally fragmented calcareous grasslands at two sites in the northern Jura mountains. Fragments of different size (area: 20.25 m², 2.25 m² and 0.25 m²) were isolated by a 5 m wide strip of frequently mown vegetation and corresponding control plots were situated in the adjacent undisturbed grassland. Aphid density was higher in fragments than in control plots. This was a combined result of a higher frequency of aphid-infested plants and larger aphid colonies in fragments than in control plots. Furthermore, a larger proportion of aphid colonies was ant-attended in fragments than in control plots. Aphid colonies were also more frequently visited by ants in fragments than in control plots in one of the 3 years. Parasitoid pressure on aphids was not influenced by the experimental fragmentation. Neither were aphid species richness and diversity affected by the fragmentation. Our study shows that even small-scale habitat fragmentation can have profound effects on the abundance of herbivorous insects. The effect on aphid density was consistent over 3 years and two sites with slightly different aphid communities.     

Differential and delayed response of two ant species to habitat fragmentation via the introduction of a pine matrix

1. Ants are a ubiquitous and crucial component of Australian Eucalyptus forests, but responses to long‐term habitat fragmentation remain poorly understood. Two ant species were followed across a 21‐year history of pine plantation establishment and maturation in a southeast Australian Eucalyptus forest. 2. At Wog Wog in southeastern Australia, a Native Eucalyptus forest was clear‐cut to make way for plantation establishment and 12 remnant patches of forest were left intact and subsequently surrounded by a pine matrix. Pitfall traps were placed in the continuous native forest, remnant Eucalyptus patches, and the pine matrix between fragments, and were stratified based on proximity to remnant patch edges and habitat type. Two ant species are focused on that represent the only remaining data for the early years of the experiment. 3. While Leptomyrmex erythrocephalus (Fabricius), the rarer of the two species, was not affected by fragmentation in the short term, 21 years after fragmentation, it was less likely to occur in both the mature pine matrix and fragments than in continuous forest controls. Aphaenogaster longiceps (Smith F.) was equally likely to occur in the fragments, continuous forest, and pine matrix early in the experiment but by year 21 post‐fragmentation was less likely to occur in the pine matrix than fragments or controls. 4. Importantly, we only detected negative impacts of fragmentation on ant occurrence as the pine plantation matrix matured and isolated ant populations on fragments. 5. It is concluded that changes in matrix suitability and specific habitat characteristics influence ant persistence in Eucalyptus fragments.     

Edge effects and intraguild predation in native and introduced centipedes: evidence from the field and from laboratory microcosms

Human alteration of habitat has increased the proportion of forest edge in areas of previously continuous forest. This edge habitat facilitates invasion of exotic species into remaining fragments. The ability of native species to resist invasion varies and may depend on intrinsic variables such as dispersal and reproductive rates as well as external factors such as rate of habitat change and the density of populations of introduced species in edge habitat. We examined the distributional and competitive relationships of two members of the class Chilopoda, Scolopocryptops sexspinosus, a centipede native to the eastern US, and Lithobius forficatus, an exotic centipede introduced from Europe. We found that L. forficatus was most abundant in edge habitat and S. sexspinosus was most abundant in the interior habitat at our field sites. Although L. forficatus was present in habitat interiors at 11 of 12 sites, there was no correlation between fragment size and numbers of L. forficatus in interior habitat. The native centipede was rarely found occupying fragment edges. We used laboratory microcosms to examine potential competitive interactions and to indirectly assess prey preferences of the two species. In microcosms both species consumed similar prey, but the native centipede, S. sexspinosus, acted as an intraguild predator on the introduced centipede. Native centipedes were competitively superior in both intraspecific and interspecific pairings. Our results suggest that intraguild predation may aid native centipedes in resisting invasion of introduced centipedes from edge habitat.     

Effect of habitat complexity on biological control by the red imported fire ant (Hymenoptera: Formicidae) in collards

A potentially important and understudied biological control agent in US agroecosystems is the red imported fire ant, Solenopsis invicta Buren. Red imported fire ants may be particularly important biological control agents because we can manipulate their abundance with changes in habitat complexity. The effect of habitat complexity on biological control by fire ants was determined using plots of collards intercropped with white clover (complex habitat) and simple collard monocrops. The most economically significant pests of collards are larvae of the diamondback moth (DBM), Plutella xylostella (L.). Predation of DBM larvae by fire ants was more rapid and efficient in the intercrop than the monocrop. Red imported fire ants were 23% less abundant in the intercrop than the monocrop, however, suggesting that fire ants had a greater per capita effect on DBM survival in the complex habitat. Red imported fire ant predation of DBM larvae was significantly affected by larval density. Red imported fire ants also reduced the survival of leaf beetles, another economically significant pest taxa, by 45%. Furthermore, collard leaf damage tended to be inversely related to fire ant density and fire ants were more effective at reducing crop damage in the complex intercrop. Our study indicates the ability of red imported fire ants to be effective biological control agents and suggests that increasing habitat complexity can enhance red imported fire ant efficacy and herbivore control.     

Influence of livestock grazing on meadow pipit foraging behaviour in upland grassland

Changes in grazing management are believed to be responsible for declines in populations of birds breeding in grassland over the last decades. The relationships between grazing management regimes, vegetation structure and composition and the availability of invertebrate food resources to passerine birds remain poorly understood. In this study, we investigated the foraging site selection of meadow pipits (Anthus pratensis L.) breeding in high intensity sheep-grazed plots or low intensity mixed (i.e. sheep and cattle)-grazed plots. We sampled above-ground invertebrates, measured vegetation height and density and conducted a vegetation survey in areas where meadow pipits were observed to forage and areas that were randomly selected. Birds foraged in areas with a lower vegetation height and density and in areas containing a lower proportion of the dominant, tussock-forming grass species Molinia caerulea. They did not forage in areas with a total higher invertebrate biomass but at areas with preferred vegetation characteristics invertebrate biomass tended to be higher in foraging sites than random sites. The foraging distance of meadow pipits was higher in the intensively grazed plots. Our findings support the hypothesis that resource-independent factors such as food accessibility and forager mobility may determine patch selection and are of more importance as selection criteria than food abundance per se. Food accessibility seems to become an even more important selection criterion under high grazing intensity, where prey abundance and size decrease. In our upland grazing system, a low intensity, mixed grazing regime seems to provide a more suitable combination of sward height, plant diversity, structural heterogeneity and food supply for meadow pipit foraging activity compared to a more intensive grazing regime dominated by sheep.     

其它可利用食物对小花宽瓣黄堇(Corydalis giraldii Fedde)种子散布的影响

作为蚁播植物种子的重要传播者,蚂蚁不但取食种子上附着的油质体,也喜食其它富含蛋白质、脂类、糖和维生素等的食物,因此环境中其它可利用食物的存在可能会影响蚂蚁对种子的搬运进而影响种子散布,但目前对于这种影响是如何发生的仍不清楚。在野外研究了蚂蚁对小花宽瓣黄堇(Corydalis giraldii Fedde)种子、肉、蜂蜜、苹果、馒头等食物的趋性和偏好程度,以及添加食物后蚂蚁对种子的拜访频率和搬运效率,以揭示其它可利用食物如何影响蚂蚁觅食和取食偏好,进而影响小花宽瓣黄堇种子散布。结果显示,在所诱捕的8种蚂蚁中,玉米毛蚁(Lasius alienus(Foerster))和丝光蚁(Formica fusca Linnaeus)是小花宽瓣黄堇种子的主要搬运者,不同食物诱捕的玉米毛蚁数量无显著性差异(P0.05),但蜂蜜和苹果诱捕的丝光蚁数量均显著大于种子(P0.05)。玉米毛蚁和丝光蚁均为杂食性,在觅食中分别行使群体募集和简单协作性募集。在仅有种子的对照处理中,玉米毛蚁和丝光蚁对种子的拜访频率分别为(38.73±4.57)头和(30.8±2.87)头(40min,n=15),两种蚂蚁对种子的拜访频率差异不显著(P0.05);玉米毛蚁和丝光蚁搬运种子的效率分别为(33.87±4.22)粒和(16.27±3.35)粒(40min,n=15),玉米毛蚁的搬运效率显著高于丝光蚁(P0.05)。与对照相比,添加馒头、苹果和蜂蜜后丝光蚁对种子的拜访频率显著降低(P0.05),分别为(15.6±3.61)头、(9.07±1.4)头和(7.67±1.58)头(40min,n=15);添加苹果和蜂蜜后丝光蚁对种子的搬运效率显著降低(P0.05),分别为(3.47±1.17)粒和(2.87±0.9)粒(40min,n=15);添加不同食物后玉米毛蚁对种子的拜访频率和搬运效率均无显著变化(P0.05)。研究结果表明行使群体募集的玉米毛蚁比行使简单协作募集的丝光蚁有更高的种子搬运效率,添加食物后影响丝光蚁对种子的拜访频率和搬运效率,这说明其它可利用食物对小花宽瓣黄堇种子散布的影响与搬运蚂蚁的种类及其觅食的募集方式有关。研究结果可为进一步研究蚂蚁与植物(种子)间的互利共生关系及其影响因素提供资料。     

Ant community structure on a small Pacific island: only one native species living with the invaders

In most studies about ant communities, species are grouped into competitive hierarchies where top dominants drive the majority of other species away from resources. Nevertheless, in some ecosystems high ground temperatures may disrupt this hierarchical organization. Other changes in community structure are caused by the arrival of invasive ant species, which rapidly disassemble local communities. We studied the effects of competition and temperature on ant community organization on Surprise Island (New Caledonia). Four different habitats were distinguished: a central plain, a sea shore Argusia shrubland, a dense Scaveola shrub, and an arboreal Pisonia strata. Eight ant species were identified from pitfall traps (seven introduced and only one native species, Pheidole oceanica). Ant assemblages in each habitat had a different ecologically dominant species, and a dominant species in one habitat could be non-dominant and less abundant in another. From interactions at baits, we built a competitive hierarchy where the top dominant species was the native Ph. oceanica. Daily foraging activity rhythms of the different species mostly overlapped. The relationship between bait occupation and ground temperature followed a negative linear pattern at all sites and for most species, except for the relatively thermophilous Monomorium floricola. Indices of co-occurrence in pitfall traps indicated that species co-occurred randomly with respect to one another. Conversely, species appeared to be segregated when we examined co-occurrence at baits at the sites where Ph. oceanica was abundant. Oceanic islands are very susceptible to alien species, but on Surprise Island it seems that the sole native species dominates in some habitats when confronted by invasive species.     

Diverse Effects of a Seven-Year Experimental Grassland Fragmentation on Major Invertebrate Groups

Habitat fragmentation is a major driver of biodiversity loss, but observed effects vary and may depend on the group examined. Time since fragmentation may explain some differences between taxonomical groups, as some species and thus species composition respond with a delay to changes in their environment. Impacts of drivers of global change may thus be underestimated in short-term studies. In our study we experimentally fragmented nutrient-poor dry calcareous grasslands and studied the response of species richness, individual density and species composition of various groups of invertebrates (gastropods, ants, ground beetles, rove beetles, orthoptera, spiders, woodlice) in 12 small (1.5 m * 1.5 m) and 12 large (4.5 m * 4.5 m) fragments and their corresponding control plots after 7 years. We further examined responses to fragmentation in relation to body size and habitat preferences. Responses to fragmentation varied between taxonomical groups. While spider species richness and individual density were lower in fragments, the opposite was true for an orthopteran species and woodlice. Species composition and β-diversity differed between fragments and control plots for some groups. However, the interaction treatment*plot size was rarely significant. Species with high occupancy rates in undisturbed control plots responded more negatively to the fragmentation, while species with large body size were relatively more abundant in fragments in some groups. No effect of the fragmentation was found for ants, which may have the longest lag times because of long-lived colonies. However, relationships between abundance and the species’ preferences for environmental factors affected by edge effects indicate that ant diversity too may be affected in the longer-term. Our results show the importance of considering different groups in conservation management in times of widespread fragmentation of landscapes. While species richness may respond slowly, changes in abundance related to habitat preferences or morphology may allow insights into likely long-term changes.     

Delayed response in a plant–pollinator system to experimental grassland fragmentation

The fragmentation of natural habitat is considered to be a major threat to biodiversity. Decreasing habitat quality and quantity caused by fragmentation may lead to a disruption of plant–pollinator interactions and to a reduction in sexual reproduction in plant species. We conducted a 6-year field experiment to investigate the effects of small-scale fragmentation on plant–pollinator interactions and genetic diversity in the self-compatible Betonica officinalis. We examined the abundance and composition of pollinators, the foraging behaviour of bumblebees and the performance, outcrossing rate and genetic diversity of B. officinalis after 2 and 6 years in experimentally fragmented nutrient-poor, calcareous grassland in the northern Swiss Jura mountains. Fragments of different size (2.25 and 20.25 m²) were isolated by a 5-m-wide strip of frequently mown vegetation. Control plots of corresponding size were situated in adjacent undisturbed grassland. Experimental grassland fragmentation altered the composition of B. officinalis pollinators and reduced their flower visitation rate. Furthermore, the foraging behaviour of bumblebees was changed in the fragments. After 6 years of fragmentation seed weight was higher in fragments than in control plots. However, the densities of B. officinalis rosettes and inflorescences, plant height and inflorescence length were not affected by fragmentation. The outcrossing frequency of B. officinalis growing in fragments was reduced by 15% after 2 years and by 33% after 6 years of experimental fragmentation. This resulted in a significant reduction of the genetic diversity in seedlings emerging in fragments after 6 years. Our study shows that small-scale habitat fragmentation can disturb the interaction between B. officinalis and pollinators resulting in a reduced outcrossing frequency and genetic diversity in plants growing in fragments. However, the response to fragmentation was considerably delayed. This finding strengthens the claim for long-term field experiments with proper replications and controls to assess delayed effects of habitat fragmentation.     

Abundance and Movement of Understory Birds in a Maulino Forest Fragmented by Pine Plantations

Understory birds are especially vulnerable to habitat fragmentation because of the reduction in habitat quality and bird movement. We study the separate effects of understory, overstory and landscape on four understory birds (tapaculos), in Central Chile, comprising a landscape mosaic of pine (Pinus radiata D. Don) plantations and native Maulino forest fragments. We also determined whether habitats with poor understory could be barrier to tapaculos movements. Abundance was measured using stationary playbacks and habitat barrier through playbacks. Understory structure was the main factor that predicted tapaculos presence and abundance. Two species, the Andean Tapaculo (Scytalopus magellanicus fuscus Gmelin) and the Ochre-flanked Tapaculo (Eugralla paradoxa Kittlitz), were positively associated with dead pine branches and negatively to forest fragment size. Tapaculos were less abundant in mature native forest, but appeared willing to cross between different habitat types. However, the Chestnut-throated Huet-huet (Pteroptochos castaneus Philippi and Landbeck), did not move from forest fragments to pine with poor understory. Overall, tapaculos species varied in their response to fragmentation depending on their habitat selection and movement capacities.     

Surrogate Habitats Demonstrate the Invasion Potential of the African Pugnacious Ant

Many ant species are highly invasive and are a significant component of disturbed ecosystems. They can have a major suppressive effect upon indigenous invertebrates, including other ants. Despite overwhelming circumstantial evidence for the ecological resourcefulness of many ants, there appears to be no experimental evidence illustrating the habitat breadth of a potentially invasive ant species. We demonstrate here that a particularly opportunistic and locally dominant ant Anoplolepis custodiens, which is a major indigenous African pest, overrides habitat structure to maintain its population level. We compared A. custodiens activity, morphology, foraging behaviour and ant species diversity in artificially established surrogate habitats (cover crops) in a vineyard containing an ample food resource in the form of the honeydew-producing mealybug Planococcus ficus. These cover crops were chosen so as to create highly altered habitats. The ant's ability to overcome these potentially suppressive habitat conditions hinged on its tight mutualism with the mealybug, and on its chasing away mealybug parasitoids. This ant species is predicted to be a latent invasive beyond Africa. It is unlikely to be impeded once it has established a foothold in a variety of novel habitats. It could locally invade to obtain food resources in a wide range of habitat types. Furthermore, in agricultural systems, cover crops are unlikely to control such an ant. Potential invasives such as this ant should be flagged as important quarantine suspects.     

Potential positive effect of the ant species Lasius niger on linyphiid spiders

Ants are highly abundant generalist predators and important ecosystem engineers which can strongly affect the composition of animal communities. We manipulated the density of the ant species Lasius niger with baits in a small‐scale field experiment to study the role of intraguild predation, top‐down control and bottom‐up effects of ants in a dry grassland surrounded by agricultural fields. Two different kinds of baits (honey and tuna) were presented near to the nests and at a distance of 2 m from six L. niger colonies in a dry grassland habitat, where L. niger was a highly abundant, omnipresent species. The experiments were performed for 1 month in spring. Additionally, the natural abundance of L. niger varying with the distance to their nests was used to study the effects on spiders and potential prey groups. The activity of L. niger was significantly higher at tuna baits compared with that at honey baits and empty control dishes. We found no effects of higher activity of L. niger on the arthropod community. However, there is evidence for a facilitation effect of ants on Collembola near to their colonies, probably due to habitat modification, which also influenced the density of Linyphiidae. Both groups had up to four times higher denisities next to L. niger colonies than at a distance of 2 m. Furthermore, δ¹³C values demonstrated that linyphiid spiders and L. niger predominantly feed on Collembola. We conclude that there is no evidence of top‐down effects of L. niger in a grassland in spring, but we found a facilitation of linyphiid spiders and their prey by the ants, which acted as ecosystem engineers.     

Interspecific interference for nest sites betweenLeptothorax congruus andMonomorium intrudens

Summary Interference competition for nest sites was studied in ant communities dominated byLeptothorax congruus in the suburbs of Tokyo, central Japan. At the two study plots located in a deciduous wood and in grassland,L. congruus constructed nests in dead branches or stems of dead grasses. Approximately 50% of the nests were physically broken within a year, suggesting that they were very fragile. Of totals of 67 and 91 nests ofL. congruus marked at the two sites, 12 (17.5%) and 53 (58.2%) nests, respectively, were replaced by other ant species (Monomorium intrudens,Crematogaster brunnea teranishii,Camponotus itoi andLasius sp.) which were common in both habitats. Field observation suggested that, among these ants,M. intrudens was a major competitor usurping the nests ofL. congruus by aggressive invasion.     

Effect of three mound-building ant species on the formation of soil seed bank in mountain grassland

The effect of three ant species (Lasius flavus, Formica spp., Tetramorium caespitum) on soil seed bank formation was studied in temperate mountain grassland. Seed removal experiments, analysis of soil seed content and seed survival experiments were carried out to evaluate the influence of ground ants on the seed fate. In the seed removal experiment seeds of 16 species, including 5 species with elaiosome-bearing seeds (myrmecochores), were exposed and their removal followed for 39 h. On average, ants removed 63.8% of myrmecochorous seeds and 10.9% of seeds without adaptation to ant dispersal. Analysis of soil seed content revealed that myrmecochores, in spite of expectations that they would accumulate in nests of seed dispersing ants, were most abundant in the soil of control plots. Evidence on seed relocation to the ant nests was obtained from a comparison of mounds of seed dispersing and seed non-dispersing ant species, as more seeds were found in the mounds of Formica spp. and Tetramorium caespitum (seed dispersers) in comparison with the mounds of Lasius favus (non-disperser).The soil seed bank of the compared microhabitats (control plots and mounds of 3 ant species) differed in their species composition, seed abundance and vertical distribution. The most distinct qualitative differences were between seed flora of control plots and mounds of Tetramorium caespitum. Control plots had approximately 30,000 propagules per m², which was double the number of seeds found in the ant mounds. In control plots, abundance and diversity of seeds steeply declined with depth; this trend was not observed in the mounds probably due to bioturbation. In the seed survival experiment, more seeds (2 out of 3 species) survived in control plots, which may also contribute to the higher seed abundance in this microhabitat.This study showed that seed relocation by ants does not contribute significantly to seed bank build-up at this study site. Ants may, however, increase the regeneration success of myrmecochores, mainly by dispersal for distance and placement in a larger spectrum of microsites, in contrast to species not adapted for myrmecochory.     

Plant diversity in sacred forest fragments of Western Ghats: a comparative study of four life forms

The effect of fragmentation on different life forms within tropical forest plant communities is poorly understood. We studied the effect of degree of fragmentation and surrounding matrix on trees, lianas, shrubs and epiphytes in tropical forest fragments of Kodagu, Western Ghats, India. These fragments exist as sacred groves amidst a highly modified agricultural landscape, and have been preserved by the religious sentiments of local communities. Plants were sampled at two sites in continuous forests and 11 forest fragments. A total of 122 species of trees, 29 species of lianas, 60 species of shrubs and 66 species of epiphytes were recorded. Trees exhibited a significant species–area relationship (R ² = 0.74). Richness estimates after controlling for stem density (rarefaction) revealed that observed species–area relationship was not an artefact of passive sampling. Variation in species richness of the other three groups was explained by stem density and structural diversity. Linear distance from the reserve forest did not explain any variation in species richness. All life forms exhibited significant nested pattern. Trees were nested along the area gradient while nestedness in the other three groups showed evidence in support of habitat nestedness. The four life forms thus responded variably to degree of fragmentation. Our study revealed that 74% of the regional diversity for trees was contributed by diversity among plots, highlighting the importance of inter-patch habitat diversity in maintaining the total regional species pool. We conclude that trees alone cannot serve as good indicator for taking appropriate conservation measures to mitigate species loss resulting from habitat fragmentation.