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.     

Optimum reproduction and dispersal strategies of a clonal plant in a metapopulation: a simulation study with Hieracium pilosella

Clonal spread is favoured in many plants at the expense of seed production in order to expand rapidly into open habitats or to occupy space by forming dense patches. However, for the dynamics of a population in a patchy landscape seed dispersal remains important even for clonal plants. We used a spatially explicit individual-based metapopulation model to examine the consequences of two trade-offs in Hieracium pilosella L: first, between vegetative and sexual reproduction, and second, between short and far-distance dispersal of seeds. Our main question was, what are the environmental conditions that cause a mixed strategy of vegetative and sexual reproduction to be optimal. The model was parameterised with field data on local population dynamics of H. pilosella. Patch dynamics were given firstly by disturbance events that opened patches in a matrix of a clonal grass that were colonisable for H. pilosella, and secondly by the gradual disappearance of H. pilosella patches due to the expanding grass. Simulations revealed opposing selection pressures on traits determined by the two trade-offs. Vegetative reproduction is favoured by local dynamics, i.e. the need for maintenance and expansion of established populations, whereas seed production is favoured by the necessity to colonise empty habitats. Similar pressures act on the proportion of seeds dispersed over short and far distances. Optimum reproductive and dispersal strategies depended on habitat quality (determined by seedling establishment probability), the fraction of dispersed seeds, and the fraction of seeds lost on unsuitable ground. Under habitat conditions supporting moderate to low seedling establishment, between 20% and 40% of reproductive effort in H. pilosella should be devoted to sexual reproduction with at least 10% of the seeds dispersed over distances suitable to attain empty patches. We conclude that in a spatially heterogeneous landscape sexual seed production in a clonal plant is advantageous even at the expense of local vegetative growth.     

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.     

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.     

Role of soil seed bank along a disturbance gradient in an alpine meadow on the Tibet plateau

We examined the role of the soil seed bank along a grazing disturbance gradient and its relationship with the vegetation of alpine meadows on the Tibet plateau, and discussed the implications for restoration. The seed bank had a high potential for restoration of species-rich vegetation; 62 species were identified in the vegetation and 87 in the seed bank, 39 species being common to both. Mean seed density was 3069–6105 viable seeds m⁻². The density of buried seeds increased significantly with increasing disturbance, indicating that restoration of disturbed areas is not seed limited. Seed density and species richness decreased with depth. The proportion of perennial species decreased with decrease in disturbance both in seed bank and in vegetation. A large portion of species with persistent seeds in the disturbed areas indicate that this seed type can be regarded a strategy of adaptation to current disturbances. Detrended correspondence analysis (DCA) showed significant differences of species composition between seed bank and vegetation, except for the seriously disturbed site. Our results suggest that the establishment of new species in severely disturbed areas is more dependent on the seed bank. By contrast, the restoration in less-disturbed and mature meadows does not rely on seed banks, and the establishment of the vegetation in these communities is more likely to rely on seed dispersal from the standing vegetation and on species with vegetative reproduction.     

Closing the gaps for animal seed dispersal: Separating the effects of habitat loss on dispersal distances and seed aggregation

Habitat loss can alter animal movements and disrupt animal seed dispersal mutualisms; however, its effects on spatial patterns of seed dispersal are not well understood. To explore the effects of habitat loss on seed dispersal distances and seed dispersion (aggregation), we created a spatially explicit, individual‐based model of an animal dispersing seeds (SEADS—Spatially Explicit Animal Dispersal of Seeds) in a theoretical landscape of 0%–90% habitat loss based on three animal traits: movement distance, gut retention time, and time between movements. Our model design had three objectives: to determine the effects of (1) animal traits and (2) habitat loss on seed dispersal distances and dispersion and (3) determine how animal traits could mitigate the negative effects of habitat loss on these variables. SEADS results revealed a complex interaction involving all animal traits and habitat loss on dispersal distances and dispersion, driven by a novel underlying mechanism of fragment entrapment. Unexpectedly, intermediate habitat loss could increase dispersal distances and dispersion relative to low and high habitat loss for some combinations of animal traits. At intermediate habitat loss, movement between patches was common, and increased dispersal distances and dispersion compared to continuous habitats because animals did not stop in spaces between fragments. However, movement between patches was reduced at higher habitat loss as animals became trapped in fragments, often near the parent plant, and dispersed seeds in aggregated patterns. As movement distance increased, low time between movements and high gut retention time combinations permitted more movement to adjacent patches than other combinations of animal traits. Because habitat loss affects movement in a nonlinear fashion under some conditions, future empirical tests would benefit from comparisons across landscapes with more than two levels of fragmentation.     

大耳姬鼠对滇西北18种植物种子的捕食

2004年9月至10月在滇西北高海拔地区对18种植物种子进行了野外被捕食研究。选取高、低两种人为干扰程度的生境;高、低两种种子密度:3粒种子/塑料盘和15粒种子/塑料盘;3种试验设置:排除啮齿动物等大型捕食者、排除蚂蚁等小型捕食者,不作任何处理作为对照。其结果表明,种子捕食者主要为夜间活动的大耳姬鼠(Apodemus latronum)。种子在高人为干扰的生境中被捕食率要显著低于低人为干扰的生境(F1,430=7.78,P<0.01);种子在高密度状况下的被捕食率要显著高于低密度状况(F1,430=13.16,P<0.001)。大耳姬鼠对于不同种类植物种子也有很强的选择性(F17,414=106.69,P<0.001),如喜好取食华山松(Pinus armandi)、中甸乌头(Aconitum chungdianensis)、豆科一种(Leguminosae sp.)和大头续断(Dipsacusasper)的种子,而不喜好取食高山柏(Sabina squamata)、黄花木(Piptanthus concolor)、子(Cotoneaster sp.)和西南鸢尾(Iris bulleyana)的种子。18种植物中,种子大小与其被捕食率高低之间无显著相关性,不同硬度等级的种子与被捕食率也无显著差异(P>0.05)。     

Leaf-cutting ant nests support less dense and impoverished seed assemblages in a human-modified Caatinga dry forest

Regenerating forests make up an increasingly large portion of tropical landscapes worldwide and regeneration dynamics may be influenced by leaf-cutting ants (LCA), which proliferate in disturbed areas and collect seeds for fungus culturing. Here, we investigate how LCA influences seed fate in human-modified areas of Caatinga dry forest. We evaluate the seed deposition and predation on Atta opaciceps nests, foraging habitat surrounding nest and control habitat away of nest influence of 15 colonies located along a forest cover gradient during the rainy and dry seasons. For each habitat, four 50-cm² plots were established and all seeds on the soil surface were collected along 1 year. We recorded 13,628 seeds distributed among 47 species and 36.57% of the total seeds did not show any sign of predation. Nest mound habitats supported low-density and species-poor seed assemblages, which were taxonomically distinct from the control habitats. These effects only occurred in the rainy season. The proportion of undamaged seeds were similar across the habitats. While forest cover did not influence seed assemblage in terms of species richness or seed predation, it did interact with habitat type via increments in seed abundance as forest cover increased across the nests. Forest cover also affected seed composition, but only in the rainy season. These results indicate that LCA decrease seed deposition in areas under their influence, particularly on the nest mounds. As LCA profit from human disturbance in the Caatinga, their role as seed ‘sinks’ should be enhanced in disturbed Caatinga patches, particularly during the rainy season, when most of the plant recruitment occurs. Our findings reinforce the importance of LCA as drivers of forest dynamics and resilience in human-modified landscapes.     

Forest Fragmentation Reduces Seed Dispersal of Duckeodendron cestroides, a Central Amazon Endemic

Fragmentation that alters mutualistic relationships between plants and frugivorous animals may reduce the seed dispersal of trees. We examined the effects of forest fragmentation on the distributions of seeds and seedlings of a Central Amazon endemic tree, Duckeodendron cestroides . In the dry seasons of 2002–2004, seeds and first-year seedlings were counted within wedge-shaped transects centered around Duckeodendron adults in fragments and nearby continuous forests at the Biological Dynamics of Forest Fragmentation Project. Analyses showed that fragmentation reduced seed dispersal quantity and quality. The percent and distance of dispersed seeds were both twice as great in continuous forest as in fragments. The distances of each tree's five furthest dispersed seeds were three times greater in continuous forest than fragments. Over the 3-yr study, 20 times more seeds were dispersed more than 10 m from parent crowns in continuous forest than fragments. A regression analysis showed more dispersed seeds at all distances in continuous forest than fragments. Dispersal differences were strong in 2002 and 2004, years of large fruit crops, but weak or absent in 2003, when fruit production was low. As distance from parent crowns increased, the number of seedlings declined more rapidly in fragments than continuous forest. Distance-dependent mortality between the seed and seedling stages appeared to be more important in continuous forest than fragments. This research provides ample, indirect evidence demonstrating that forest fragmentation can result in the breakdown of a seed dispersal mutualism, potentially jeopardizing the reproduction of a rare, tropical tree.     

Experimental studies of seed predation in old-fields

Summary In a pair of experiments conducted in old-field habitats in southwestern Michigan (USA), we examined rates of seed loss to post-dispersal predators (ants and rodents). Seeds from 4–6 species of biennial plants were tested over a range of seed densities and habitat types. We found that seed removal was significantly higher in vegetated habitats than in areas of disturbed soil (both simulated small-animal diggings and a plowed field). In the undisturbed vegetation, seed losses ranged from 1–20% of seeds removed/day.An exclosure experiment demonstrated that ants and rodents foraged selectively for seeds of the six plant species tested. Rodents (Peromyscus maniculatus) fed preferentially on species producing large seeds (predominantly Tragopogon dubius). Ants (Myrmica lobicornus) foraged on smaller seeds, although their foraging preferences were not based strictly on seed size.Seed density had only a minor effect on predation rate over the range of densities tested. Predators, instead appeared to treat each experimental group of seeds as a single prey patch. Consequently, predation intensity was quite variable over distances of <20 m within a relatively homogeneous section of habitat.These field experiments provide initial estimates of seed losses to post-dispersal predators in old-field habitats. Rates of seed loss were generally less than those reported from desert or semi-arid habitats. However, for some old-field species, seed losses averaged an appreciable 10–20% day. The selective nature of the seed predators, plusthe relative patchiness of predation intensity in space, suggest that postdispersal seed predation can play a role in determining the distribution and/or abundance of old-field herbs.     

濒危植物明党参种子散布和种子库动态研究

对伞形科濒危植物明党参（Changium smyrnioides)的种子生产、散布和种子库动态进行了研究。明党参平均每株产生完好种子132粒,86.36%来自顶生花序,不同生境中个体产生的种子千粒重有差异;其平均1.06m高的花葶在种子散布过程中易倒伏,使种子远离母株;种子散布格局为聚集型;种子库密度随时间逐渐下降。明党参种子产量和不同时期种子库密度均低于同科非濒危植物峨参（Anthriscus sylvestris),2个种的种子库均为非持续型;明党参具有低种子数、大粒种子、种子散布远离母株以减少种内竞争的K对策,在受到人类大量采挖和生境干扰后种群不容易迅速恢复;而峨参采取高种子数、小粒种子、种子近母株密集分布和产生无性系的r对策,种群受干扰后易恢复。     

Analyses of spatial patterns and population processes of clonal plants

The nonrandom spatial structure of terrestrial plants is formed by ecological interactions and reproduction with a limited dispersal range, and in turn this may strongly affect population dynamics and population genetics. The traditional method of modelling in population ecology is either to neglect spatial pattern (e.g. in transition matrix models) or to do straightforward computer simulation. We review here three analytical mothods to deal with plant populations in a lattice-structured habitat, which propagate both by seeds that scatter over the whole habitat and by vegetative reproduction (producing runners, rhizomes, etc.) to neighboring vacant sites. [1]Dynamics of global and local densities: Dynamical equations of population density considering nearest-neighbor correlation (spatial clumping) are developed as the joint dynamics of global average density and local density (comparable to mean crowding) based onpair approximation. If there is a linear trade-off between seed production and vegetative reproduction, the equilibrium abundance of the population may be maximized by engaging both means of reproduction. This result is accurately predicted by the pair approximation method, but not by mean-field approximation (neglect of spatial structure). [2]Cluster size distributions: Using global and local densities obtained by pair approximation, we predicted cluster size distribution, i.e. the number of clusters of occupied sites of various sizes. [3]Clonal identity probability decreasing with distance: Multi-locus measurement of allozymes or other neutral molecular markers tells us whether or not a given pair of individuals belong to the same clone. From the pattern of clonal identity probability decreasing with the distance between ramets, we can estimate the relative importance of two modes of reproduction: vegetative propagation and sexual seed production.     

Impact of seed abundance on seed processing and dispersal by the red ant Myrmica rubra

1. Myrmecochory sensu stricto is an ant–plant mutualism in which non‐granivorous ants disperse plant diaspores after feeding on their nutrient‐rich seed appendage, the elaiosome. Phenological traits associated with the diaspore can influence the behaviour of ants and thus their ultimate efficiency as seed dispersers. 2. This study investigated how a contrasting availability of seeds (20 vs. 200 seeds) from the diplochorous Chelidonium majus (Papaveraceae, Linnaeus) plant species influences the behaviour of Myrmica rubra (Formicidae, Linnaeus) ants, from the retrieval of seeds until their dispersal outside the ant nest. 3. Regardless of seed abundance, the ants collected the first diaspores at similar rates. Then, seed retrieval sped up over time for large seed sources until satiation took place with only one‐third of the tested colonies wholly depleting abundant seed sources. 4. No active recruitment by trail‐laying ants was triggered, even to an abundant seed source 5. In both conditions of seed abundance, the majority of the diaspores retrieved inside the nest were discarded with the elaiosome removed and were dispersed at similar distances from the nest. 6. The paper concludes with a discussion of how the quantity of seeds released by a plant with a dual mode of dispersal can potentially influence the behaviour of ant dispersers and hence the dispersal efficiency derived from myrmecochory.     

Seed dispersal in Cakile edentula var. lacustris : decoupling the fitness effects of density and distance from the home site

A factorial design of three densities of siblings at three local distances from seed parents was employed to distinguish effects of density from effects of dispersal distance on lifespan and fruit production of Cakile edentula var. lacustris, a plant with heteromorphic seeds. The segmented fruits produce two seed types: proximal and distal, with distal seeds having greater mass and greater dispersibility. Effects of longer distances (0.5 km and 30 km) on lifespan and fruit production were investigated using plants at low density. The prediction was tested that the greater seed mass of distal seeds increases fitness when seeds are dispersed into sites of unknown quality away from the home site or when seeds are dispersed to low density. High density caused earlier mortality and lower probability of reproduction. Distance from the maternal plant did not influence lifespan or reproduction at distances of 15 m or less, but lifespan was longer 0.5 km from the home site. No interaction was detected between the effects of density and distance on either lifespan or total fitness. Environmental conditions that influence fitness did not vary as a function of dispersal distance in this system, and favorable conditions at the home site did not persist between generations. Therefore, selection on dispersion patterns in natural conditions is likely to be through effects of density rather than dispersal distance. Proximal seeds had greater reproduction than distal seeds at the home site, and distal seeds had greater reproduction at the more distant sites (but not the most distant site), as expected, but these performance differences could not be attributed to differences in mass between the two seed types. Reduced seed mass was favored at the most distant site, but larger seed mass was favored most strongly at low density. Seeds that are dispersed to low density are larger, suggesting that although kin selection may limit the effectiveness of individual selection to increase seed mass under conditions of sibling competition, density-dependent individual selection on seed mass, rather than distant-dependent selection, also contributes to the observed associations among seed type, seed mass and dispersal ability. Received: 21 October 1996 / Accepted: 4 December 1996     

Effects of Local Plant Density and Incomplete Dichogamy on the Reproductive Success of the Rare Neotropical Palm Geonoma epetiolata

Density dependent processes are known to influence reproduction and establishment of plant populations. In this study, we evaluated the effects of local density and sexual expression on the reproductive success of the rare palm species Geonoma epetiolata in Costa Rica. We classified individuals in two density categories based on distances to the two nearest neighbors and recorded the occurrence of sexual overlap on each individual. Overlap between pistillate and staminate flowers in monoecious plants allows geitonogamous pollination, potentially reducing dependence on mates for reproduction. We measured plant size and light availability, and evaluated the influence of these variables on pollination success, fruit production, fruit abortion and seed mass. Pollination success significantly increased with density but there was no effect of sexual overlap. In contrast, there was no effect of density on the probability of initiated fruit, and a negative effect on fruit set. Fruit abortion was lowest in isolated plants with no sexual overlap. Plant size and light did not contribute to variation in reproductive success. In conclusion, the pollination advantage of plants in dense neighborhoods does not compensate reproductive losses incurred by fruit abortion, possibly due to mating among genetically related individuals. Geonoma epetiolata is threatened by habitat loss and poaching of seeds for the horticultural market. High fruit abortion rates associated with density in G. epetiolata suggest that seed collections from the remnant wild populations of this palm species may cause bottlenecks that further threaten population viability.     

Multiphase myrmecochory: the roles of different ant species and effects of fire

Seed dispersal by ants (myrmecochory) can be influenced by changes to ant assemblages resulting from habitat disturbance as well as by differences in disperser behaviour. We investigated the effect of habitat disturbance by fire on the dispersal of seeds of a myrmecochorous shrub, Pultenaea daphnoides. We also investigated the consequence of the seed relocation behaviours of two common dispersers (Pheidole sp. A and Rhytidoponera metallica) for the redispersal of seeds. Pheidole sp. A colonies did not relocate seeds outside their nests. In contrast, R. metallica colonies relocated 43.6 % of seeds fed to them, of which 96.9 % had residual elaiosome that remained attached. On average, R. metallica relocated seeds 78.9 and 60.7 cm from the nest entrances in burned and unburned habitat, respectively. Seeds were removed faster in burned than in unburned habitat, and seeds previously relocated by R. metallica were removed at similar rates to seeds with intact elaiosomes, but faster than seeds with detached elaiosomes. Dispersal distances were not significantly different between burned (51.3 cm) and unburned (70.9 cm) habitat or between seeds with different elaiosome conditions. Differences between habitat types in the frequency of seed removal, the shape of the seed dispersal curve, and the relative contribution of R. metallica and Pheidole sp. A to seed dispersal were largely due to the effect of recent fire on the abundance of Pheidole sp. A. Across habitat types, the number of seeds removed from depots and during dispersal trials most strongly related to the combined abundances of R. metallica and Pheidole. Our findings show that myrmecochory can involve more than one dispersal phase and that fire indirectly influences myrmecochory by altering the abundances of seed-dispersing ants.     

Fitness and survival in fragmented populations of Narthecium ossifragum at the species' range margin

Decreasing population size and increasing isolation may lead to reduced plant fitness. These effects of habitat fragmentation may especially apply to populations close to the margin of their geographical range, where populations generally are often smaller and more isolated, and the environment is less favourable than at the range centre. In this context we studied north-west German range-margin populations of Narthecium ossifragum (L.) HUDS., a clonal, perennial herb typical for acidic and nutrient-poor wet heathlands. We asked whether plant fitness and reproduction of the range-margin populations were affected by population size and habitat quality, and whether any changes in population size over the past 10–20 years were related to past population size and habitat quality.Population size varied between 60 and 100.000 individuals and was highly positively related to shoot density. Larger (and denser) populations formed more flowers and fruits, but fewer seeds. Soil water content had a strong positive effect on seed production and seed mass, and the latter also increased with increasing soil phosphorus content. Relative light intensity showed a positive impact on shoot density and, hence, population size. Compared to British range centre populations, the populations at the German range margin formed more, but smaller seeds that failed to germinate both in the field and in laboratory experiments. Despite the lack of sexual reproduction, population sizes generally had not decreased during the past two decades, most likely due to successful clonal growth. The change in population size of Narthecium, however, was negatively related to the present soil phosphorus content, indicating that an increased supply of nutrients and a decrease in light availability may have a negative effect on population dynamics.The results imply that fragmented and small range-margin populations not necessarily are doomed to extinction, provided that the habitat quality remains suitable for growth and vegetative reproduction.     

Effects of disturbance intensity on seasonal dynamics of alpine meadow soil seed banks on the Tibetan Plateau

Background and Aims

Information on soil seed bank processes is crucial for understanding vegetation dynamics. Despite the documented importance of soil seed banks in many ecosystems, their role is not fully understood in some sensitive habitats, such as the alpine meadows of the Tibetan Plateau.

Methods

We studied the seasonal dynamics of the germinable soil seed bank under four disturbance intensities in an alpine meadow on the Tibetan Plateau as well as seed size distribution relative to disturbance intensity. Composition of the seed bank was compared with that of the standing vegetation.

Results

Density of buried seeds increased with disturbance intensity, but species richness and species diversity decreased. Seed density and species richness of the seed bank varied seasonally in all layers (0–2, 2–7, 7–12 cm) and the whole (0–12 cm). The species composition of seed bank was not significantly influenced by season. There was no trend in seed size distribution as disturbance increased. Seasonal seed bank turnover rates increased with increase in disturbance. The result of the NMDS showed that species composition of seed bank and vegetation exhibited a fairly uniform pattern in each season.

Conclusions

Although as a whole the species composition of the vegetation and seed bank showed a relatively low degree of similarity in each season, similarity was highest in the most disturbed habitat. There was no alteration in species composition of seed bank regardless of disturbance intensity, but seed density decreased as disturbance increased. Disturbances in alpine plant communities might increase persistence of regeneration niches. Regeneration from the seed bank together with vegetative reproduction contributed to aboveground vegetation in highly disturbed habitats. Clonal species played an important role in regeneration of vegetation in slightly disturbed areas, where there was little contribution of ruderals from soil seed banks.     

蚂蚁对澜沧舞花姜种子散布及种苗空间分布格局的影响

 在野外系统观测了澜沧舞花姜（Globba lancangensis）18个果实共216粒种子的散布过程。共有10种蚂蚁参与了澜沧舞花姜的种子散布,距离为0.01～3.35 m,平均距离(0.47±0.03) m（平均值±SE,n=216）。其中最重要的３种蚂蚁是横纹齿猛蚁（Odontoponera transversa）、大头蚁(Pheidole sp.)和黄足厚结猛蚁(Pachycondyla luteipes),其出现频率分别为61%、50%和28%,散布的平均距离分别为(0.60±0.09) m、(0.20±0.01) m和(0.32±0.05) m。从总体上看,蚂蚁促进了种子的分散,降低了种子的聚集程度。横纹齿猛蚁对于种子上的油质体最为敏感,对人工去除了油质体的种子不搬运,对种子散布距离较远,暗示了其与澜沧舞花姜之间可能存在更紧密的互惠关系。野外样方调查结果表明,在3种舞花姜属植物中,以种子繁殖为主的澜沧舞花姜种苗之间的平均最近距离为(36.8±1.45) cm（平均值±SE,n=74）,显著大于以珠芽繁殖为主的毛舞花姜（Globba barthiri）的(29.8±2.70) m（n=34）（t73,33=2.11,p=0.037）和异果舞花姜（Globba racemosa）的(28.7±3.16) cm（n=32）（t73,31=2.33,p=0.022）;澜沧舞花姜的种苗聚集程度（Z=-1.70±0.19）显著小于毛舞花姜（Z=-2.58±0.37,t73,33=2.36,p=0.020）和异果舞花姜（Z=-3.28±0.53）（t73,31=3.54,p=0.001）。这说明相对于毛舞花姜和异果舞花姜,蚂蚁对种子的散布作用显著增加了澜沧舞花姜种苗间的平均最近距离,降低了居群的聚集度。     

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.