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1.
Recruitment is an important process in regulating many marine benthic communities and many studies have examined factors controlling the dispersal and distribution of larval immigrants. However, benthic species also have early post-settlement life-stages that are dramatically different from adult and larval stages. Predation on these stages potentially impacts measured recruitment and the benthic populations and communities that ultimately develop.We examined the consequences of post-settlement predation on 1-day-old to 1-month-old recruits of sessile invertebrates at two field sites in southern New England. One site (Breakwater) was in a protected area with few predators and the other (Pine Island) was <1 km away in an open coast area with three different predator guilds: small and large invertebrates and fish. The Breakwater site had been dominated for >10 years by colonial and solitary ascidians. These species were absent from the Pine Island site which was dominated by bryozoans. Our goal was to examine whether post-settlement predation influenced the development and subsequent structure of the epifaunal community.Here we examine long-term changes in community development resulting from post-settlement predation, and contrast these results to those of earlier experiments examining the reductions in observed recruitment by post-settlement predation. Our first long-term experiment examined natural community development at the two sites and whether transplanted communities changed when exposed to the different levels of predation at these sites. The communities that developed at both sites were consistently different from each other and similar to resident communities at their respective sites. On panels transplanted from the Breakwater to Pine Island, solitary ascidians and the colonial ascidian, Botryllus schlosseri, suffered high mortalities on both caged and uncaged treatments, indicative of predation by small predators that could enter cages. Some solitary ascidians did survive inside cages and the colonial ascidian, Botrylloides violaceus, became dominant on all transplanted treatments. On panels transplanted from Pine Island to the Breakwater, ascidians invaded and dominated all treatments except those that were originally caged at Pine Island.In the second long-term experiment, natural communities were allowed to develop on panels exposed at the Breakwater for 1, 2, 3, and 4 weeks. Each set was transplanted to three treatments at Pine Island: open uncaged pilings, caged pilings to exclude fish and large invertebrates, and racks suspended above the bottom to exclude all predators. When 1-week-old communities were transplanted, after 2-3 weeks only bryozoans were found on the open and caged pilings, while colonial ascidians dominated the suspended rack treatment. When older 2-week-old communities were transplanted, colonial ascidians also became dominant in the caged piling treatment and when 3- and 4-week-old communities were transplanted colonial ascidians dominated all three treatments. Solitary ascidians were never abundant on open pilings exposed to fish and large benthic invertebrate predators.Post-settlement predator-prey interactions involved newly settled and juvenile life-stages of a variety of prey species and many invertebrate and vertebrate predator species. The effects of these interactions on recruitment did result in differences in the development and eventual species composition of the communities, even though predators had little if any effect on the adults of the prey species.  相似文献   

2.
We investigated the effects of temporal variability in a disturbance regime on fouling communities at two study sites in a northern-central Chilean bay. Fouling assemblages grown on artificial settlement substrata were disturbed by mechanical removal of biomass at different time intervals. Using one single disturbance frequency (10 disturbance events over 5 months) we applied 7 different temporal disturbance treatments: a constant disturbance regime (identical intervals between disturbance events), and 6 variable treatments where both variableness and sequences of intervals between disturbance events were manipulated. Two levels of temporal variableness (low and high, i.e. disturbance events were either dispersed or highly clumped in time) in the disturbance regime were applied by modifying the time intervals between subsequent disturbance events. To investigate the temporal coupling between disturbance events and other ecological processes (e.g. larval supply and recruitment intensity), three different sequences of disturbance intervals were nested in each of the two levels of temporal variableness. Species richness, evenness, total abundance, and structure of communities that experienced the various disturbance regimes were compared at the end of the experiment (15 days after the last disturbance event). Disturbance strongly influenced the community structure and led to a decrease in evenness and total abundance but not species richness. In undisturbed reference communities, the dominant competitor Pyura chilensis (Tunicata) occupied most available space while this species was suppressed in all disturbed treatments. Surprisingly, neither temporal variableness in the disturbance regime nor the sequence of intervals between disturbance events had an effect on community structure. Temporal variability in high disturbance regimes may be of minor importance for fouling communities, because they are dominated by opportunistic species that are adapted to rapidly exploit available space.  相似文献   

3.
The palatability of two solitary and three colonial species of ascidians commonly found in sub-tropical seagrass meadows was evaluated using the abundant, sympatric, omnivorous pinfish Lagodon rhomboides as a model predator. Bite-sized pieces of fresh tissues of both solitary and one of the three colonial ascidian species were unpalatable to fish. Lipophilic and hydrophilic extracts of the three unpalatable species did not cause feeding deterrence indicating that secondary metabolites are not responsible for the lack of palatability. Distaplia bermudensis, the one colonial ascidian that was unpalatable to fish, had a highly acidified outer tunic (pH = 1.5). We tested the ability of acidified agar food pellets (pH = 1.5) to deter pinfish and found that the fish readily ingested acidified pellets. The toughness of the tunic of all five ascidian species was evaluated by measuring the Force (N) required to penetrate the tunic using a penetrometer. Tunic toughness is likely to explain the lack of palatability of the solitary ascidians Styela plicata and Molgula occidentalis as their tunics required a force of > 34 N to penetrate. Tunic toughness may be a particularly effective adaptation for ascidian defense in seagrass habitats where fish with strong crushing jaws, such as those that commonly occur in coral reef systems, are rare.  相似文献   

4.
Disturbance is claimed to be one of the most important triggers of biological invasions. There is a lack of data about disturbance impacts on the youngest life stage of invasive trees and shrubs. Thus, we aimed to assess the role of disturbances in shaping responses of natural regeneration of three model invasive species—Prunus serotina Ehrh., Quercus rubra L., and Robinia pseudoacacia L.—to disturbances in forest plant communities. Our study was conducted over 3 years on 372 study plots (100 m2), across nine types of temperate forests in Wielkopolski National Park (Poland). Disturbance was assessed using ecological indicator values for disturbance severity and frequency. Our study revealed the high importance of disturbance on species composition of understory vegetation. We also found relationships between ecological success (density and biomass of natural regeneration) of invaders and disturbance indices. These models were statistically significant but their effect sizes were low. Due to the low effect sizes, we can state only limited conclusions about impact of disturbance on ecological success of invasive species natural regeneration. The results suggest that for seedlings (up to 50 cm height—threshold between understory and shrub layer) disturbance, a leading factor in biological invasions of numerous taxa, has a small role in this case. Thus, we may assume that their ecological success is connected with stochastic processes in populations of the invader’s seedlings, rather than with stochastic release from competition caused by disturbances.  相似文献   

5.
The ability of the sponge Halichondria panicea to assimilate into the fouling communities of the blue mussel Mytilus edulis and the solitary ascidian Styela rustica has been studied in a field experiment, in which sponge fragments have been introduced artificially into epibenthic communities. The growth of H. panicea was suppressed greatly in the presence of young mussels; its survival rate averaged 40%. In the communities where S. rustica dominated, the survival rate of H. panicea reached 100%, but the growth rate was lower than in the control group (without competitor species). Despite the high natural growth rate and toxicity, the settlement success of H. panicea was low on the substrate occupied by another fouling species.  相似文献   

6.
Assessing patterns of species distribution and abundance is important to understand the driving processes of, and predict future changes in, biodiversity. To this date, ecological studies have been mainly designed to investigate the effects of the mean magnitude of predictor variables, although ecological factors naturally vary in space and time. In a nine month long field experiment, we tested the effects of different temporal patterns (regular, lowly and highly irregular) in biomass removal (=disturbance event) on the diversity, species composition, and biomass accrual of macrobenthic assemblages grown on 15 × 15 cm2 PVC-panels. For each pattern of disturbance, disturbance events were timed at three sequences to control for possible confounding effects with recruitment patterns. Disturbance intensity was kept identical among treatments. Assemblages developed in the absence of disturbance for 3 months prior to a 150-day manipulation period, during which the biomass from 20% of the panel area was removed at each of ten disturbance events. Additional undisturbed settlement panels were deployed in the field to assess monthly recruitment rates and species succession over a one year period. Disturbance (i) reduced biomass and total species cover, (ii) changed species composition during the first half of the manipulation period significantly, and (iii) was without effect on species richness and evenness. Irregular disturbance regimes enhanced the abundance of the ascidian Ciona intestinalis, biomass accrual, and total species cover of assemblages relative to the regular disturbance regime, but had either no or only transient effects on diversity and species composition, respectively. Neither the degree of irregularity in disturbance nor the sequence of disturbance events affected any of the response variables significantly. Recruitment of species was strongly seasonal with almost only diatoms recruiting during winter, while recruitment was most intense during summer. Our results suggest that the temporal patterns of predictor variables might be of low explanatory power for the variance of responses in communities with seasonal recruitment patterns that are exposed to a high level of disturbance. Thus the need to include temporal patterns of predictor variables in experimental designs may depend on community dynamics and the characteristics of the process under investigation. S. Wollgast and M. Molis contributed equally to this work.  相似文献   

7.
Anthropogenic alterations of historical disturbance regimes (e.g. suppressing floods and wildfires) is a primary mechanism by which exotic species can come to dominate native communities. Unfortunately, reinstating historical disturbance regimes to restore native communities has achieved mixed success. The presence of positive frequency dependence (PFD) is commonly invoked to explain why exotic plant invasions are so difficult to eradicate. However, models examining PFD have not considered the effect of reintroducing disturbances. Using a spatially explicit individual‐based model, we consider how magnitude and direction of frequency dependence of native and exotic species affects the success of reintroducing disturbances that favour fitness of natives over exotics. Our model illustrates why restoration is difficult; there is a narrow range of parameters that allows for native species to eliminate or coexist with exotics once they have established. Dominance by exotic invaders occurs with moderate initial frequencies of exotic individuals, aggregation of these individuals, or an exotic propagule production advantage. Reintroducing disturbances allows native dominance only when PFD of the exotic is weaker than that of the native species, disturbance intervals are short, and/or exotics are not initially frequent. Our framework provides guidelines for conditions in which the reintroduction of disturbances will effectively restore invaded habitats.  相似文献   

8.
This paper reports the findings of a short-term natural invasibility field study in constructed Mediterranean herbaceous communities of varying diversities, under a fire treatment. Three components of invasibility, i.e. species richness, density and biomass of invaders, have been monitored in burnt and unburnt experimental plots with resident diversity ranging from monocultures to 18-species mixtures. In general, species richness, density and biomass of invaders decreased significantly with the increase of resident species richness. Furthermore, the density and biomass of invading species were significantly influenced by the species composition of resident communities. Although aboveground biomass, leaf area index, canopy height and percent bare ground of the resident communities explained a significant part of the variation in the success of invading species, these covariates did not fully explain the effects of resident species richness. Fire mainly influenced invasibility via soil nutrient levels. The effect of fire on observed invasibility patterns seems to be less important than the effects of resident species richness. Our results demonstrate the importance of species richness and composition in controlling the initial stages of plant invasions in Mediterranean grasslands but that there was a lack of interaction with the effects of fire disturbance.  相似文献   

9.
Biological invasions are a major threat to natural biodiversity; hence, understanding the mechanisms underlying invasibility (i.e., the susceptibility of a community to invasions by new species) is crucial. Invasibility of a resident community may be affected by a complex but hitherto hardly understood interplay of (1) productivity of the habitat, (2) diversity, (3) herbivory, and (4) the characteristics of both invasive and resident species. Using experimental phytoplankton microcosms, we investigated the effect of nutrient supply and species diversity on the invasibility of resident communities for two functionally different invaders in the presence or absence of an herbivore. With increasing nutrient supply, increased herbivore abundance indicated enhanced phytoplankton biomass production, and the invasion success of both invaders showed a unimodal pattern. At low nutrient supply (i.e., low influence of herbivory), the invasibility depended mainly on the competitive abilities of the invaders, whereas at high nutrient supply, the susceptibility to herbivory dominated. This resulted in different optimum nutrient levels for invasion success of the two species due to their individual functional traits. To test the effect of diversity on invasibility, a species richness gradient was generated by random selection from a resident species pool at an intermediate nutrient level. Invasibility was not affected by species richness; instead, it was driven by the functional traits of the resident and/or invasive species mediated by herbivore density. Overall, herbivory was the driving factor for invasibility of phytoplankton communities, which implies that other factors affecting the intensity of herbivory (e.g., productivity or edibility of primary producers) indirectly influence invasions.  相似文献   

10.
We aim to develop a simple model to explore how disturbance and propagule pressure determine conditions for successful invasion in systems where recruitment occurs only in disturbed sites. Disturbance is often thought to favour invaders as it allows recruitment; however, the effects of disturbance are more complicated when it results in mortality of the invader. When disturbance rates in both invader occupied and unoccupied sites are the same, recruitment and mortality effects are exactly balanced, and successful invasion is independent of the disturbance regime. Differences in the disturbance rates between invader occupied and unoccupied sites can occur through invader modification or management of disturbance. Under these conditions, we found a novel mechanism for the generation of an Allee effect, which occurs when the invader promotes disturbance in sites it already occupies. When Allee effects occur one-off, large-scale disturbances can result in permanent, dramatic shifts in invader abundance; and conversely, reducing the population below a critical threshold can cause extinction.  相似文献   

11.
Biological invasions have become major players in the current biodiversity crisis, but realistic tools to predict which species will establish successful populations are still unavailable. Here we present a novel approach that requires only a morphometric characterisation of the species. Using fish invasions of the Mediterranean, we show that the abundance of non‐indigenous fishes correlates with the location and relative size of occupied morphological space within the receiving pool of species. Those invaders that established abundant populations tended to be added outside or at the margins of the receiving morphospace, whereas non‐indigenous species morphologically similar to resident ones failed to develop large populations or even to establish themselves, probably because the available ecological niches were already occupied. Accepting that morphology is a proxy for a species' ecological position in a community, our findings are consistent with ideas advanced since Darwin's naturalisation hypothesis and provide a new warning signal to identify invaders and to recognise vulnerable communities.  相似文献   

12.
For species recruiting into established sessile communities, the adult colonies and individuals already present form a significant part of the environment and have the potential to alter both larval settlement rates and post-settlement mortality. Settlement rates can be reduced by predation on larvae, by the removal or addition of substratum space, or by stimulation or prohibition of larvae from settling on adjacent substratum. Once attached, the recruiting individual can still be influenced by predation or overgrowth by residents, by the added physical structure for firmer attachment, or by being camouflaged from motile predators. To examine those processes by which residents affect recruitment we exposed experimental substrata with three densities of adults of a single species at a site in eastern Long Island Sound, USA for a 1-wk period. Seven different species of common invertebrates were used in nine separate experiments. The major effect of most resident species was the usurpation of space and the restricting of recruitment to adjacent unoccupied areas. This was particularly true for resident ascidians and bryozoans, but less so for barnacles and oysters. In fact several species recruited in higher densities on or next to oysters and barnacles. Comparison to 1-day settlement experiments indicated that the encrusting ascidian species Diplosoma and possibly Botryllus reduced recruitment relative to settlement, probably by overgrowing newly-settled individuals. However, in the presence of most resident species, recruitment patterns were not greatly different from settlement patterns, indicating that the effects of the attached community on recruitment may result from influences on settlement.  相似文献   

13.
The solitary ascidian Halocynthia papillosa (Linnaeus, 1767) is proposed as a good indicator of the deleterious effect of SCUBA diving on the Mediterranean coralligenous communities. A comparative survey of H. papillosa populations at frequented and unfrequented localities was carried out over a two-year period (during 2006 and 2007), before and after a peak diving season in the Sierra Helada Marine Park (SW Western Mediterranean Sea). We observed bigger and more abundant individuals of H. papillosa at undived sites than at frequented dived sites during the period of study. Furthermore, individuals of H. papillosa in the most frequented localities occupied more cryptic positions than in the undived localities. H. papillosa was shown to be very sensitive to the adverse effects of SCUBA diving. This species could represent a reliable bioindicator of diving activity and as such constitute a useful tool for the quick and easy monitoring of impact on coralligenous communities before this damage becomes unmitigatable.  相似文献   

14.
The role of disturbance in community ecology has been studied extensively and is thought to free resources and reset successional sequences at the local scale and create heterogeneity at the regional scale. Most studies have investigated effects on either the disturbed patch or on the entire community, but have generally ignored any effect of or on the community surrounding disturbed patches. We used marine fouling communities to examine the effect of a surrounding community on species abundance within a disturbed patch and the effect of a disturbance on species abundance in the surrounding community. We varied both the magnitude and pattern of disturbance on experimental settlement plates. Settlement plates were dominated by a non-native bryozoan, which may have established because of the large amount of initial space available on plates. Percent covers of species within the patch were affected by the surrounding community, confirming previous studies' predictions about edge effects from the surrounding community on dynamics within a patch. Disturbance resulted in lower percent cover in the surrounding community, but there were no differences between magnitudes or spatial patterns of disturbance. Disturbance lowered population growth rates in the surrounding community, possibly by altering the abiotic environment or species interactions. Following disturbance, the recovery of species within a patch may be affected by species in the surrounding community, but the effects of a disturbance can extend beyond the patch and alter abundances in the surrounding community. The dependence of patch dynamics on the surrounding community and the extended effects of disturbance on the surrounding community, suggest an important feedback of disturbance on patch dynamics indirectly via the surrounding community.  相似文献   

15.
Disturbance can affect both the diversity and invasibility of communities. Many field studies have found correlations between diversity and susceptibility to invasion, but if both factors independently respond to disturbance then spurious non-causal relationships may be observed. Here, we show that disturbance can cause a temporal shift in the diversity-invasibility relationship. In a field experiment using sessile marine communities, disturbance strongly affected both diversity and invasion such that they were highly correlated. Disturbance facilitated initial invasion, creating a negative diversity-invasibility relationship when the invader first arrived. Over time, disturbance hindered the persistence of invaders, creating a positive diversity-invasibility relationship. We suggest that temporal changes in the diversity-invasibility relationship may have contributed to the 'invasion paradox', a term for the contrasting patterns of experimental and observational studies of the diversity-invasibility relationship.  相似文献   

16.
Past disturbance events shape future community trajectories if their ecological legacies interact with contemporary disturbance regimes to affect recruitment and reproduction. In pastures, large-scale disturbance events can become undetectable in aboveground community components but persist as augmented weed seedbanks, suggesting disturbance history may be a predictor of weed invasion magnitude following subsequent disturbance. Using a randomized block factorial design, we investigated how soil disturbance intensity, timing and history interact to affect weed recruitment, diversity and reproduction by implementing a range of disturbance intensities in April and June 2004 in pastures of differing disturbance history. Response variables were followed for two years. Disturbance timing interacted with disturbance intensity and history to affect recruitment, with some June treatment combinations having ≥48% higher weed abundance than those in April. Relative to undisturbed controls, low to intermediate disturbance intensity facilitated recruitment by ≥117% but high intensity disturbance had highest recruitment, particularly in previously disturbed pastures (≥543% increase). In both years, weed species richness was highest in intense disturbances in previously disturbed pastures. Importantly, weed reproduction was nearly one order of magnitude higher in intense disturbance patches, especially in previously disturbed pastures, and increased through time by ≥243%. These findings indicate weed recruitment, diversity and reproductive output are seed- and microsite-limited, and that intense soil disturbance may result in high long-term weed abundance. Although moderate disturbance facilitated recruitment, reproduction here was low, suggesting modest disturbance will not appreciably increase weed abundance. The most common species were Taraxacum officinale and Poa pratensis, both of which can benefit forage-livestock production, but most establishing species were of low palatability. We recommend grassland managers explicitly integrate disturbance history into dynamic management planning, consider augmenting seedbanks with ‘desirable’ weed species to help communities recover following inevitable disturbance events and do not rely exclusively on aboveground characters to evaluate their system's vulnerability to undesirable plant establishment and persistence.  相似文献   

17.
Evolution of solitary or colonial life histories in tunicates is accompanied by dramatic developmental changes that affect morphology and reproduction. We compared vasa expression in a solitary ascidian and a closely related colonial ascidian, in an effort to uncover developmental mechanisms important during the evolution of these contrasting life histories, including the ability to reproduce by budding. In this study, we explored the origin of germ cells in new buds developing by asexual reproduction in a colonial ascidian, Botrylloides violaceus and compared it to the source of germ cells in a solitary ascidian Boltenia villosa. We studied expression by in situ hybridization of vasa, a DEAD box RNA helicase gene found in germ cells across the metazoans. In B. villosa, bv-vasa mRNA was expressed in putative germ cells and oocytes of adult gonads, and was sequestered into a posterior lineage during embryogenesis. In mature colonies of the ascidian B. violaceus, bot-vasa mRNA was expressed in putative spermatogonia, in oocytes of zooids, and in some circulating cells in the zooids and differentiating buds. We propose that expression of vasa in cells other than gonadal germ cells of zooids in a colonial ascidian may serve as a source of germ-line stem cells in the colony.  相似文献   

18.
Much uncertainty remains about traits linked with successful invasion – the establishment and spread of non‐resident species into existing communities. Using a 20‐year experiment, where 50 non‐resident (but mostly native) grassland plant species were sown into savannah plots, we ask how traits linked with invasion depend on invasion stage (establishment, spread), indicator of invasion success (occupancy, relative abundance), time, environmental conditions, propagule rain, and traits of invaders and invaded communities. Trait data for 164 taxa showed that invader occupancy was primarily associated with traits of invaders, traits of recipient communities, and invader‐community interactions. Invader abundance was more strongly associated with community traits (e.g. proportion legume) and trait differences between invaders and the most similar resident species. Annuals and invaders with high‐specific leaf area were only successful early in stand development, whereas invaders with conservative carbon capture strategies persisted long‐term. Our results indicate that invasion is context‐dependent and long‐term experiments are required to comprehensively understand invasions.  相似文献   

19.
Despite the progressive accumulation of exotic species in natural communities, little effort has been devoted to elucidating the mechanisms underpinning the coexistence of invaders in environmentally and biologically heterogeneous systems. The exotic seaweeds, Asparagopsis taxiformis and Caulerpa racemosa, exhibit a segregated distribution on Mediterranean rocky reefs. A. taxiformis dominates assemblages in topographically complex habitats, but is virtually absent on homogenous platforms. In contrast, C. racemosa achieves extensive cover in both types of habitat. We assessed whether differences in their distribution were generated by biotic interactions (between invaders and/or between invaders and natives) or by environmental constraints. Three models were proposed to explain seaweed distribution patterns: (1) invaders inhibit one another; (2) native assemblages, differing between complex and simple habitats, prevent the establishment/spread of one invader, but not that of the other; and (3) environmental conditions regulate the establishment/persistence of the seaweeds in different habitats. We removed the dominant invader and resident assemblages in each type of habitat. Moreover, A. taxiformis thalli were transplanted into the habitat dominated by C. racemosa to establish whether its failure to colonize the simple habitat was due to the lack of propagules or post-recruitment mortality. C. racemosa spread in the complex habitat was not influenced by the removal of resident assemblages, but it was slightly enhanced by A. taxiformis removal. Neither C. racemosa removal nor that of resident assemblages promoted A. taxiformis colonization and survival in simple habitats. Our results suggest that heterogeneity in environmental conditions can promote invader coexistence by mitigating the effects of negative biotic interactions. Therefore, the accumulation of introduced species in native communities does not necessarily imply established invaders fostering further invasion.  相似文献   

20.
Many shallow water subtidal habitats in Massachusetts, USA have recently been invaded by five non-indigenous ascidian species: Ascidiella aspersa, Botrylloides violaceus, Didemnum sp., Diplosoma listerianum and Styela clava. This study examined the effects of seawater temperature, as a proxy for climate change, on B. violaceus and D. listerianum and the impact these ascidians have on native sessile fouling communities. Field experiments were conducted over a four month period at two locations (Lynn and Woods Hole, MA) to examine growth dynamics over regional thermal and geographic ranges. Invasive ascidians occupied as much as 80% of the primary substratum and accounted for the majority of species richness. B. violaceus and D. listerianum growth were similar at both study sites, but initial colony growth of D. listerianum was positively affected by temperature. B. violaceus and D. listerianum exhibited rapid two-week growth rates during the summer months with more rapid growth at the warmer Woods Hole site. Competition for space between B. violaceus and D. listerianum typically resulted in neutral borders between colonies. Overgrowth occurred if the colony of one species was disproportionably larger than the colony of the other species. Recruitment and growth of native species influenced the long-term composition of experimental communities more than the pre-seeding with B. violaceus or D. listerianum colonies. Elevated temperatures, however, increased initial growth of B. violaceus and D. listerianum and may have facilitated the species success to invade the communities during crucial periods of introduction. With projected global climate change, a rise in sea surface temperatures may exacerbate the cumulative impacts of invasions on benthic communities and facilitate the invasion of other non-native ascidian species.  相似文献   

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