Seed Banks and Species Richness Potential of Coal Slurry Ponds Reclaimed as Wetlands

Seed bank experiments are described to assess the species richness potential of coal slurry ponds reclaimed as wetlands (ranging from 6 to more than 40 years old). Experimental treatments test the drawdown and flooded conditions characteristic of the vegetation dynamics of emergent wetlands in the Upper Mississippi Valley. More seedlings, primarily annuals, emerged from exposed wet sediments (freely drained) than under continuous flooded sediments in cold ponds (339 versus 136 seedlings m^?2, respectively) and in natural ponds (163 versus 47, respectively). More seeds were produced by plants established in freely drained conditions than under flooded conditions from sediments in the coal ponds (26546 versus 1842 seeds m^?2, respectively) and the natural ponds (28430 versus 4526, respectively). Similarly, more biomass was also produced by these plants in freely drained than under flooded conditions in coal ponds (118 versus 47 g m^?2, respectively) and natural ponds (118 versus 52, respectively). Fertilization (NPK) did not affect germination for the most part, but it did affect seed set and biomass production, especially for C₄ annuals such as Echinochloa crusgalli and Panicum dichotomiflorum. I propose that lime (calcium carbonate) and fertilizer be applied during the first few scheduled drawdowns for these coal slurry ponds reclaimed as wetlands to increase the number of species and to allow their more rapid development as self-sustaining systems.     

What is a suitable management for Typha latifolia control in wet meadows?

Aim

Typha latifolia causes serious problems in wet meadows by overgrowing and suppressing other native plants. To determine suitable management for T. latifolia control, we addressed the following question: What are the effects of long-term cutting at different frequencies (once or twice per year and no management) and biomass removal on cover and other characteristics of T. latifolia, and on sward productivity and plant species composition?

Location

Malá Strana nature reserve, Jizerské hory Mountains, Czechia.

Methods

A long-term experiment arranged in a randomised block design with three blocks was established in 2005. Data were collected from five treatments: unmanaged control; cutting once a year in June without biomass removal and with biomass removal; cutting twice per year in June and August without biomass removal and with biomass removal. Percentage cover of T. latifolia and other vascular plant species was visually estimated and T. latifolia characteristics (tiller density, height, dry-matter biomass [DMB] yield and litter), sward height and DMB yield were measured during 2005–2018 at the end of June.

Results and Discussion

Regular cutting once or twice per year regardless of cut biomass removal led to reductions in tiller density, height, litter and DMB yield of T. latifolia. Biomass removal had only a slight tendency to affect T. latifolia characteristics. The higher frequency of cutting significantly decreased the mean T. latifolia cover, litter and DMB yield. Cutting once or twice per year regardless of biomass removal led to successive changes in plant species composition but had no effect on the species richness and evenness.

Conclusions

Cutting at least once per year without biomass removal seems to be sufficient to achieve a decrease in DMB yield and litter of T. latifolia plants, and thereby maintain the wet-meadow vegetation without loss of species richness and also preventing the overgrowth of shrubs and trees.     

Effects of salinity and cutting on the development of Phragmites australis

The effects of increased salinity and cutting the above ground biomass on the growth of Phragmites australis were evaluated by investigating four experimental reed stands grown in outdoor tanks. Two stands were treated with 30 salinity and the other two stands with freshwater; one stand of each treatment was cut to 20 cm during the second growing season. Growth conditions were observed until all the plants were dead at the end of the second year. The number of shoots emerged from the freshwater-treated stand was about 70% higher than that of the saltwater-treated stand. The number of shoots emerged from cut plant stands were markedly lower than uncut stands. The average shoot height was negatively affected by salinity and shoots that emerged after cutting further decreased in height. The average number of leaves on a shoot was not significantly affected by salinity, but reduced by cutting in both treatments. Leaf length, width and the distance between leaves were decreased by both salinity and cutting. In the freshwater-treated uncut stand more than 50% of the shoots formed panicles, but this proportion was reduced to 6% by salinity, to 15% by cutting, and to 0% by the combination of salinity and cutting. This study showed again that salinity reduces the growth of aboveground components. The growth, however, was most severely retarded by cutting combined with salinity, which has many implications for better management of P. australis stands.     

Phragmites australis makes valuable floating mat biotopes under oligotrophic conditions

To understand the mechanism of how Phragmites australis makes valuable floating mat biotopes under oligotrophic conditions, we investigated the environmental (water chemistry) and vegetational characteristics (growth, plant species richness, and floristic composition) of a floating mat consisting of three main mat-forming species with a zonal distribution (P. australis on the land side of the floating mat, Zizania latifolia on the middle area, and Typha angustifolia on the water side). Although they showed relatively low growth in the floating mat, compared to those in land-based wetlands, P. australis grew better than other mat-forming species in terms of shoot height and biomass production. Specifically, P. australis made more below-ground parts (593?±?38 g/m²) than other mat-forming species (Z. latifolia, 100?±?10 g/m²; T. angustifolia, 167?±?8 g/m²) and more companion species were found in P. australis-dominated plots (8.5?±?1.0 species/m²) than other plots (Z. latifolia-dominated plots, 2.7?±?0.6 species/m²; T. angustifolia-dominated plots, 1.0?±?0.0 species/m²). The larger amount of below-ground P. australis parts could contribute to thicker and denser mat structures, possibly providing more favorable habitats for neighboring plant species, thus facilitating more companion species within the P. australis-dominated area of the mat.

     

Slashing Phragmites (Phragmites australis) prior to planting does not promote native vegetation establishment

Phragmites or Common Reed (Phragmites australis) is a tall rhizomatous cosmopolitan grass. While native to Australia, it can be invasive in wetlands, forming dense monocultures and reducing their ecological integrity. We assessed the potential for the cutting of Phragmites reeds prior to planting to promote the establishment of indigenous shrubs that might ultimately outcompete Phragmites. We established ten 5 m × 5 m quadrats in an area dominated by Phragmites, brush‐cut the reeds to ground level in five of them and left five uncut as controls. Within each quadrat, we planted 20 plants (~40 cm tall) of each of five indigenous shrub species, unguarded (4 plants/m²). We surveyed the plants one month after planting and annually for the following four years for survival, height and browsing damage. Browsing damage to plants was common (>50%) and unaffected by cutting. After four years, overall plant survival rates were ~25% and mean plant heights for the five shrub species ranged between 120 and 174 cm. Cutting of Phragmites had no positive effect on plant survival or height. In fact, two Melaleuca species grew taller in the uncut quadrats. Cutting of Phragmites reed beds prior to planting is unlikely to promote the establishment of woody plantings. However, planting within established Phragmites with or without prior brush‐cutting is worthy of further trialling as a potential tool for reinstating native diversity at Phragmites‐dominated sites.     

RELATIONSHIPS BETWEEN FLOODING TOLERANCE,LIFE HISTORY,AND SHORT-TERM COMPETITIVE PERFORMANCE IN THREE SPECIES OF POLYGONUM

The objective of this study was to compare the growth and short-term (single season) competitive performance of three species of Polygonum known to differ in flooding tolerance and life history. Polygonum amphibium is a perennial with low sexual reproductive effort and a relatively high degree of flooding tolerance, P. lapathifolium is an annual species with a high sexual reproductive effort and a low tolerance to flooding, and P. hydropiperoides is intermediate to the other two in terms of sexual reproductive effort and flooding tolerance. In order to determine the relative growth and competitive abilities of these species, mixtures and monocultures of plants were grown in pots and maintained under three flooding regimes: 1) flooded, 2) partially drained, and 3) well drained. Both P. hydropiperoides and P. amphibium grew best under flooded and partially drained conditions with reduced growth in the drained treatment. Polygonum lapathifolium, in contrast, grew as well in the drained treatment as in the more flooded treatments. Results from competition experiments were consistent in showing the relative competitive abilities to be P. lapathifolium > P. hydropiperoides > P. amphibium regardless of flooding regime. Thus, short-term competitive performance was found to trade off with flood tolerance rather than with sexual reproductive effort.     

Trunk cutting initiates bottom-up cascades in a tri-trophic system: sprouting increases biodiversity of herbivorous and predaceous arthropods on willows

We examined the effects of trunk cutting on the regrowth responses of two dominant willow species, Salix eriocarpa and S. gilgiana , and the subsequent effects on the community structure (abundance and species richness) of herbivorous and predaceous arthropods. We studied ten randomly selected pairs of cut and uncut (control) trees of each willow species. Field observations showed that when the trunks were cut, shoots sprouted from the base and developed rapidly. These shoots continued to grow until July, producing lateral shoots and leaves, whereas current-year shoots on uncut trees essentially stopped growing by April. In July, the upper leaves of cut trees were less tough and had a greater water and nitrogen content than leaves of uncut trees. Leaf consumption and abundance and species richness of both herbivorous and predaceous arthropods were significantly greater on cut trees than on uncut trees in both willow species. Overall, trunk cutting resulted in at least a two-fold increase in both relative abundance and species richness within the arthropod community on the willow species. We concluded that the severe physical damage caused by trunk cutting greatly increased the biodiversity of herbivorous and predaceous arthropods through bottom-up cascading effects.     

Use of horizontal subsurface flow constructed wetlands to treat reverse osmosis concentrate of rolling wastewater

According to the characteristics of the reverse osmosis concentrate (ROC) generated from iron and steel company, we used three sets of parallel horizontal subsurface flow (HSF) constructed wetlands (CWs) with different plants and substrate layouts to treat the high-salinity wastewater. The plant growth and removal efficiencies under saline condition were evaluated. The evaluation was based entirely on routinely collected water quality data and the physical and chemical characteristics of the plants (Phragmites australis, Typha latifolia, Iris wilsonii, and Scirpus planiculmis). The principal parameters of concern in the effluent were chemical oxygen demand (COD), total nitrogen (TN), and total phosphorus (TP). The results showed that the CWs were able to remove COD, TN, and TP from ROC. S. planiculmis was not suitable for the treatment of high-saline wastewater. The sequence of metals accumulated in CW plants was K>Ca>Na>Mg>Zn>Cu. More than 70% of metals were accumulated in the aboveground of P. australis. The CW filled with gravel and manganese ore and planted with P. australis and T. latifolia had the best performance of pollutant removal, with average removal of 49.96%, 39.45%, and 72.01% for COD, TN, and TP, respectively. The effluent water quality met the regulation in China. These results suggested that HSF CW planted with P. australis and T. latifolia can be applied for ROC pollutants removal.     

ANATOMICAL AND METABOLIC RESPONSES TO WATERLOGGING AND SALINITY IN SPARTINA ALTERNIFLORA AND S. PATENS (POACEAE)

The effects of waterlogging and salinity (25 or 325 mol m ³ NaCl) stressors on the anatomy and metabolism of the marsh grasses 5. alterniflora Loisel. and S. patens Aiton (Muhl.) were investigated in a V factorial greenhouse experiment over 30 d. Waterlogging and salinity in combination resulted in anatomical and metabolic responses in both species. Waterlogging reduced soil redox potential and decreased root-specific gravity significantly in both species. The inadequacy of aerenchyma development under hypoxia to support aerobic root respiration in S. patens was indicated by significant increases in root alcohol dehydrogenase (ADH) activity of 1,752% and 420%, respectively, in the low and high salinity treatments. ADH activity was not increased significantly by flooding of S. alterniflora. Proline concentrations in roots and leaves were low at low salinities and increased significantly at high salinities in both species, but only under drained conditions. Decrease in leaf elongation by high salinity occurred in drained, but not flooded treatments in both species. Under flooded conditions, leaf elongation was significantly greater in S. alterniflora than S. patens. Greatest leaf elongation occurred in flooded low salinity S. alterniflora plants that had the least proline. Although both species are adapted to waterlogging and salinity, S. alterniflora appears to be more tolerant of reducing soil conditions and less responsive to higher salinity than S. patens.     

Inhibition of <Emphasis Type="Italic">Zizania latifolia</Emphasis> growth by <Emphasis Type="Italic">Phragmites australis</Emphasis>: an experimental study

The influence of Phragmites australis on the growth of Zizania latifolia, and the morphological characteristics of both species, were investigated at two nutrient levels. The experiment was carried out over three growing seasons between May 2003 and December 2005. The experimental plot was longitudinally divided into two equal halves, one of which was planted with Zizania latifolia (Zizania zone) and the other one with Phragmites australis (Phragmites zone). Four weeks after transplantation, the plot was again divided horizontally, perpendicular to the previous division, and subject to low (LN) and high nutrient (HN) treatments. Measured growth indices of the two species were more developed in the HN than in the LN treatments. Both species were observed to invade each other’s zone, but Phragmites latifolia seemed to out-compete Zizania latifolia individuals. This was evident from the decrease in below ground biomass of Zizania latifolia in the third growing season and deterioration in above ground organs with time. It was concluded that Phragmites australis out-competes Zizania latifolia due to better developed root and rhizome system.     

Vegetation regeneration on blanket mire after mechanized peat-cutting

• 1 Blanket mire in Northern Ireland is an ecologically threatened habitat in which land use for hand peat‐cutting, forestry and agriculture has had a major influence. A recent land use change is the introduction of tractor‐powered peat‐harvesting. In this paper, the effect of machine peat‐cutting on ombrotrophic blanket mire vegetation is assessed from a regional sample of cut and uncut plots.
• 2 Principal components analysis identified water‐table depth and grazing intensity as major factors influencing the species composition of uncut mire. A key variable affecting the composition of machine‐cut mire across the drainage gradient was the number of times cut, with multiple annual cutting causing progressive decreases in acrotelm depth, catotelm bulk density and plant cover. Ericaceous species and Sphagnum spp. were particularly sensitive to cutting, with Eriophorum angustifolium and Campylopus introflexus characteristic of multiple‐cut sites.
• 3 Redundancy analysis, with number of times cut partialled out, showed that recovery time accounted for a significant amount of variance in vegetation composition. Species that significantly increased in abundance with recovery time were Sphagnum spp., Odontoschisma sphagni , Erica tetralix and Drosera rotundifolia.
• 4 Sites cut frequently, or which were grazed, recovered more slowly. Recovery from cutting was partly dependent on the post‐cutting structure of the mire surface and the species that survive cutting. The rate of recovery on sites cut once, then abandoned, is relatively rapid compared with multiple‐cut sites where species colonization is constrained by bare compacted peat.

     

生物炭添加对不同水氮条件下芦苇生长和氮素吸收的影响

生物炭能改良土壤从而促进植物生长和氮素吸收,但其作用效果是否受水氮条件的影响尚不清楚。以湿地植物芦苇为研究对象,在3种氮添加水平（无添加,30 kg hm^-2 a^-1和60 kg hm^-2 a^-1）和两种水分（淹水和非淹水）条件下分别进行生物炭添加和不添加处理,结果表明：（1）生物炭添加能促进芦苇根系生长,在非淹水条件下根系生物量增加了40.5%,在淹水条件下根系生物量增加了20.1%。（2）生物炭添加能促进非淹水条件下芦苇的氮素吸收,能提高淹水条件下芦苇的氮素生产力。（3）生物炭添加加剧了土壤氮素损失,且在非淹水高氮条件下作用最强,可能是由于生物炭促进了芦苇的氮素吸收。芦苇氮素吸收速率与土壤氮损失之间存在显著的正相关关系。因此,在添加生物炭时,需要考虑土壤水分状况和氮素富集程度以及植物的氮素吸收偏好。该研究结果可为生物炭在湿地生态系统中的应用提供参考。     

The effects of breaking or bending the stems of two rhizomatous plants, Phragmites australis and Miscanthus sacchariflorus, on their communities

To understand the effects of disturbance on vegetation, bending and cutting experiments were conducted on two rhizomatous plant species, Phragmites australis and Miscanthus sacchariflorus, in a floodplain area of the Arakawa River, Japan. The plants were damaged in the late development growth stage on 3 August 2004 (August disturbance) and in the middle development growth stage on 29 June 2005 (June disturbance). The severity of the damage was evaluated based on shoot morphology and belowground biomass. The recoveries of the two plants from the flood-like artificial disturbance were compared with undisturbed stands. The morphological response of the shoot was higher after the June disturbance than after the August disturbance in both plants. In contrast, the recovery of belowground biomass in P. australis at the end of the growth season was higher after the August disturbance (87 and 72% for bending and cutting, respectively) compared with the June disturbance (84 and 60% for the same). The recoveries in M. sacchariflorus for the two disturbances showed the opposite trend (73 and 59% for bending and cutting, respectively, after the August disturbance, and 90 and 73% after the same disturbance in June). The study demonstrated that an event like flooding, whether it is breaking or bending, will cause greater damage if it occurs at the late development growth stage in M. sacchariflorus compared with the middle development growth stage. In contrast, P. australis tolerated disturbances up to a certain magnitude; after that, the effect was more severe in the middle development growth stage compared with late development growth stage.     

Native and invasive plant interactions in wetlands and the minimal role of invasiveness

The effects of invasive plants on plants native to areas that are being invaded can be quite variable, depending on the species of the invasive plant involved as well as the physical characteristics of the location being invaded. My study focuses on the effects of Phragmites australis Linnaeus (common reed) and Lythrum salicaria L. (purple loosestrife) on the same native plant community. Uninvaded plots dominated by native plants Typha angustifolia L. (narrowleaf cattail) and Typha latifolia L. (broadleaf cattail) served as the control. I surveyed percent cover of species during early summer and midsummer for 3 years in six Hudson River freshwater tidal wetlands (sites). Differences in species richness, composition and abundance were small, but significant among invaded and uninvaded plots and among sites. However, these differences remained significant when data for dominant species (invasive and native) were removed. Differences in native plant species abundance were attributed to invasive plant species-specific characteristics and differences in species richness and composition were attributed to physical location (zonation) in these freshwater tidal marshes. “Invasive” status of a dominant plant species was less important in invasive plant–native plant interactions than species-specific characteristics and zonation. Further research into the effects of site and land-use on invasive plant impacts is recommended.     

Role of plant aeration in zonation of Zizania latifolia and Phragmites australis

The emergent plants Phragmites australis (Cav.) Trin. ex Steud and Zizania latifolia (Griseb.) Stapf. are distributed along a gradient of increasing water depth, with P. australis restricted to shallow water and Z. latifolia to deeper water. The lower limits of the two vegetational zones were discussed from the view-point of oxygen supply from aerial shoots. The oxygen influx to the underground organs from aerial shoots is proportional to the gradient between oxygen concentrations in atmosphere and inside the basal stem (C). The proportional constant (Q), called here the ventilation coefficient, expresses the efficiency of the diffusion of oxygen. An attempt was made to measure Q and C. The values of Q for the two species in their dominant niches are higher in Z. latifolia than in P. australis, corresponding to the natural distribution with increasing depths of water. Q in Z. latifolia and P. australis first increased and then decreased. Corresponding to such changes of Q, the oxygen concentration inside the basal stem (C) fell to low values for Z. latifolia at the greatest water-depth. Since in Z. latifolia a low value of C in deeper waters was accompanied by a decline in standing crop of aerial shoots, it was concluded that oxygen is the limiting factor for the distribution of Z. latifolia in the deeper waters.     

Morphological changes and resource allocation of Zizania latifolia (Griseb.) Stapf in response to different submergence depth and duration

Plants around ponds, rivers and lakes are subjected to long-term partial or complete submergence. When they are flooded, water level affects the plants simultaneously with duration of submergence. Separate and interactive effects of water level and duration on the growth of the herbaceous perennial Zizania latifolia (Poaceae) were investigated by exposing the plants in greenhouse water tanks to submergence in different water depths and for different time-spans. The plants exhibited great shoot elongation upon submergence and prolonged flood duration, and the basal tiller number of the species decreased with higher water levels. Submergence treatment advanced the flowering date and increased the inflorescence number. Plant total biomass did not differ among all the treatments, while the root:shoot ratio decreased with increased water level, prolonged duration of submergence and their interaction. The high plasticity in morphology and shifts in reproductive strategy and biomass allocation enabled the Zizania plants to survive the compound effect of flooding height and duration. This may explain the occurrence of this species in habitats subjected to long-term flooding. The results obtained in this experiment will contribute to understanding the impact of flooding dynamics on plants and the ways of adaptation responses to prolonged waterlogging.     

Observations on the colonization of a young polder area in the netherlands with special reference to the clonal expansion ofPhragmites australis

In 1968, the polder Zuidelijk Flevoland was reclaimed in the Netherlands. Observations on species succession were made during a period of seven years in an area later known as the nature reserve Oostvaardersplassen. Directly after reclamation, a large number of plant species were recruited. Here, only the dynamics of the three dominant species will be discussed, i.e. the annual mudflat speciesSenecio congestus, and the long-lived emergent macrophytesTypha latifolia andPhragmites australis. Small differences in elevation had a large impact on seedling recruitment. Overall,S. congestus dominated in the first year after reclamation, andT. latifolia prevailed during the second and third year. Thereafter,P. australis became dominant, despite fewer seedlings. In the first three years after reclamation, clones ofP. australis grew very slowly, afterwards they showed a constant expansion rate of ca. 4 m/yr. However, close to pools waterfowl grazing slowed down the expansion of clones. The observed spatial and temporal variation in the colonization of this young polder area can be explained by species differences in life history traits, resource capture, as well as in the susceptibility to stem-boring insects and waterfowl grazing.     

Oxygen demand and supply in Zizania latifolia and Phragmites australis

The oxygen demand of underground organs in tall emergent plants was assumed to be supplied with the oxygen flux through aerial shoots. Measurements were made of the respiratory activity of underground organs and oxygen flux through aerial shoots in Zizania latifolia (Griseb.) Stapf and Phragmites australis (Cav.) Trin. ex Steud.Oxygen uptake rates of Z. latifolia roots and rhizomes decreased as the supply of oxygen decreased. Phragmites australis, however, did not respond to decreases in the supply of oxygen until the oxygen supply was relatively low. The values for old rhizomes of P. australis were very low. Most of the oxygen flux for P. australis was derived from standing dead shoots. The important role of dead shoots of P. australis in plant aeration is suggested. The degree of sufficiency of oxygen supply for Z. latifolia was 179% in May and 92% in August. The values for P. australis were 45% both in May and August. Furthermore, estimation of mean oxygen concentration of inner underground organs for P. australis showed very low values, 1.6% both in May and August, even in shallow water.     

Facilitation by nurse shrubs of resprouting behavior in a post-fire shrubland in northern Patagonia,Argentina

Abstract. The nurse-plant syndrome is a widely recognized example of positive (facilitative) influences of plant species on the establishment and growth of other species. Most studies of the nurse-plant syndrome have been on species that reproduce mainly from seed rather than vegetatively. In this study, we experimentally compared the influences of two species of nurse shrubs, Schinus patagonicus and Berberis buxifolia, on the survival and growth of vegetatively reproducing herbaceous and woody plants in a post-fire shrubland in northern Patagonia, Argentina. The vegetation beneath shrubs was removed by clipping and, in a paired-sample design, one half of the canopy of each shrub was removed. We determined species richness, counted number of resprouts, and measured photon flux density and soil moisture beneath cut and uncut halves of each shrubs. Abundances of resprouts were several times greater beneath the uncut vs. the cut shrubs, as was the mean number of species. Thus, shrubs have a strong facilitating influence as measured by resprout densities and the number of species. Numbers of resprouts and of species were twice as high beneath Schinus as beneath Berberis, implying important differences in the facilitative effects of the two shrubs species. Microsites beneath Schinus were characterized by lower and more heterogeneous light levels but by greater soil moisture. Even though the reproductive mode in this post-fire shrubland is overwhelmingly vegetative, facilitation by nurse shrubs is important and differentially effective for different species of nurse shrubs.     

Morphological adaptations of emergent plants to water flow: a case study with Typha angustifolia, Zizania latifolia and Phragmites australis

1. Water velocity plays an important role in shaping plant community structure in flowing waters although few authors have yet attempted to explain the adaptation of plants to flow. 2. We aimed to test two hypotheses, that: (i) some emergent macrophytes reconfigure their shoot distribution in fast currents and form clumps, and (ii) the shape and morphology of such clumps minimises drag caused by the current. The study focuses on three emergent macrophytes that co‐occur along a gradient of water velocity. 3. The species showed a clear zonation in response to water depth and current velocity. Phragmites australis occupied shallower and more slowly flowing water than Typha angustifolia and Zizania latifolia, which had similar preferences. 4. Both T. angustifolia and Z. latifolia shoots were more clumped at high velocity, whereas they were more randomly distributed at low flow or in stagnant water. Because of the low shoot density, water flowed more easily through T. angustifolia clumps, whereas Z. latifolia clumps had a high shoot density and large amounts of trapped litter, causing stagnant water in the centre of the clump. The clumps of Z. latifolia with a high density of shoots were longer and narrower than T. angustifolia clumps. Phragmites australis was less tolerant of flow than the other two species and large amounts of litter trapped in the clumps impaired flow. 5. The shoot distribution of both T. angustifolia and Z. latifolia is reconfigured at high flow and this minimises drag on the clumps.