Behavioural responses of the lizard Pedioplanis l. lineoocellata to overgrazing

We studied the effects of overgrazing on the foraging behaviour of the lizard Pedioplanis l. lineoocellata (Spotted Sand Lizard), a sit-and-wait forager, in habitats of differing vegetation states to determine the effects of habitat degradation on this species. At high grazing intensity where vegetation cover and diversity is low, the lizard P. lineoocellata moves more frequently, spends more time moving and covers larger distances than in habitats where vegetation cover and diversity is high. These behavioural changes in movement patterns can be explained by less abundant prey in habitats with low vegetation cover and diversity. Although morphology, phylogeny and physiology of P. lineoocellata should constrain the change in foraging behaviour, the species has modified its foraging strategy from sit-and-wait to actively foraging. We assume that this behavioural flexibility of P. lineoocellata is a buffer mechanism enabling the species to use and survive in degraded (unfavourable) habitats.     

Strategies for Savanna Restoration in the Southern Great Plains: Effects of Fire and Herbicides

Woody plant encroachment has degraded grassland and savanna ecosystems worldwide by decreasing herbaceous production and diversity, and altering these physiognomies toward woodlands. This study evaluated the long-term efficacy of fire and herbicide restoration strategies used in the southern Great Plains to reduce Honey mesquite ( Prosopis glandulosa ) dominance, restore a grassland/savanna physiognomy, and increase herbaceous production and diversity. Three treatments were evaluated: high-intensity winter fire, aerial spray of clopyralid + triclopyr (C + T), and aerial spray of clopyralid and were compared to untreated mesquite woodland (control). Post-treatment mesquite stand physiognomy was different between fire (low mortality, high basal sprouting), C + T (high mortality, high basal sprouting of surviving plants), and clopyralid (moderate mortality, low basal sprouting of surviving plants) treatments. From 6 to 8 years post-treatment, herbaceous production was increased in C + T and clopyralid treatments but not in the fire treatment. Mesquite regrowth in the fire treatment exerted a competitive influence that limited herbaceous production. Herbaceous functional group diversity was increased in fire and C + T treatments due to a decrease in C₃ perennial grass dominance and an increase in C₄ perennial grasses and/or C₃ forbs. Treatments that maintained mesquite overstory (control and clopyralid) had lower herbaceous diversity due to C₃ perennial grass dominance and lower C₄ perennial grass cover. The clopyralid treatment demonstrated greatest potential for long-term restoration of southern Great Plains savanna by reducing mesquite canopy cover to historic levels, limiting mesquite basal regrowth and increasing grass production.     

Severely degraded dunes of the southern Kalahari: local extinction,persistence and natural re‐establishment of plants

This study aimed to quantify and understand the impact of severe land degradation on plant attributes and diversity on dunes of the southern Kalahari. Heavy grazing pressure in particular resulted in a significant decline of canopy cover and species number in annual and perennial life forms; forb and graminoid growth forms; erect and prostrate habits; and leafy stem, tussock and stoloniferous architectures. However, no significant change was found in shrub and tree forms which persisted without apparent new recruitment. Under these conditions, species diversity dropped sharply and a number of species, mainly graminoids, became apparently locally extinct. The perennial shrub, Crotalaria cf. spartioides, showed the converse with a high frequency of establishing seedlings on the degraded dunes. Changes in relative dominance show that the extreme treatment favours perennial over annual, woody over graminoid and forb, erect over prostrate and leafy stem over stoloniferous and tussock. Some of these results and certain species and soil responses differ from those reported from grazing studies elsewhere, and are possibly no longer directly related to the impact of the primary grazing pressure but to the secondary effect of subsequent instability of the dunes.     

Land use affects rodent communities in Kalahari savannah rangelands

Shrub encroachment caused by overgrazing has led to dramatic changes of savannah landscapes and is considered one of the most threatening forms of rangeland degradation leading to habitat fragmentation. Although changes to plant assemblages are becoming better known, however, our understanding of how shrub encroachment affects rodent communities is low. In this study, we investigated relative abundance of five rodent species in sixteen southern Kalahari rangelands where shrub cover ranged from low (<5%) to high (>25%). Rodent abundance was determined on three trapping grids (40 × 100 m) for each site. Our results show that increasing shrub cover affected rodent species differently. The relative abundance of hairy‐footed gerbil, short‐tailed gerbil and bushveld gerbil declined with increasing shrub cover, whereas highveld gerbil and striped mouse exhibited hump‐shaped relationships with shrub cover. Overall, species richness decreased with increasing shrub cover and a negative impact of high shrub cover above 15% on rodent abundance was congruent for all species. We conclude that our results support the hypothesis that long‐term heavy grazing that results in area wide shrub encroachment, threatens the diversity of arid environments.     

Swimming activity of seabass: comparing patterns obtained in natural environment and in re-circulating tanks under high density

Seabass ( Dicentrarchus labrax ) swimming activity was compared between natural environments and aquaculture facilities. Behaviour under natural conditions was assessed in a saltmarsh pond (250 m², 18 × 14 × 0·8 m) using acoustic telemetry. From several surveys, we documented the diel activity rhythm and demonstrated group effects on swimming patterns and amplitudes by comparing activity of solitary fish with that of a fish living in a group of 60. Consequences of weather variability were also analysed and revealed a high sensitivity of fish to atmospheric conditions for both swimming and demand‐feeding behaviour. Behaviour in fish tanks was also studied using acoustic telemetry, as part of the EUREKA EU1 960 'Aqua‐Maki 2' project investigating aspects of fish culture in re-circulating tanks under high density. A re-circulating hexagonal tank (5·4 × 5·4 m, 1·8 m depth, 48 m³) was equipped with positioning and demand‐feeding systems, oxygen and temperature probes. Initial density was 50 kg m³ in March and rose to 90 kg m³ at the end of the experiment in May. During this period, the movements of nine fish were continuously recorded for 24 h each, reaching a total of six 24 h episode at eight days interval. Swimming activity was analysed in terms of activity rhythms and space occupation specially around feeding events. The two data set and main results will be presented and compared to assess seabass behavioural plasticity and sensitivity to husbandry conditions.     

Responses of mammalian carnivores to land use in arid savanna rangelands

Shrub encroachment due to heavy grazing has led to dramatic changes in arid savanna landscape structure worldwide. It is considered to be one of the most threatening forms of rangeland degradation altering plant diversity. However, possible impacts of shrub encroachment on species diversity at higher trophic levels remain poorly understood. Additionally, indirect effects, such as changes of trophic interactions, are often ignored when trying to understand changes in biodiversity patterns. In this study, conducted in the southern Kalahari, we explored how the diversity of small carnivores and their prey is affected by shrub encroachment. We analysed the relationships between abundance and diversity of small carnivores, the availability of their main prey groups (coleopterans, termites, grasshoppers, rodents) and the structural diversity of the landscape (shrub, grass and herb cover). Eight hundred track surveys were conducted to determine carnivore abundance on 20 rangeland habitats, which represented a gradient of grazing intensity. Prey availability was surveyed for each study site and related to vegetation cover.

Results show a significant impact of shrub cover on abundance and diversity of carnivores and their prey. The diversity of both, carnivores and their prey, showed a hump-shaped response to increasing shrub cover whereas relative carnivore abundance decreased. Availability of prey groups was affected differently by shrub cover increase. Diversity of carnivores was best predicted by shrub cover (R²>0.7, p<0.001) indicating the overriding role of habitat structure as compared to prey availability. We conclude that intermediate shrub cover values enrich structural diversity of savanna landscape and in consequence sustain diversity of small carnivores and their prey in arid and semiarid ecosystems.     

Demography and habitat selection of small mammals on Zomba Plateau, Malawi

Small mammals were studied in tow montane grassland-forest areas on Zomba Plateau (1900m), Malawi, in 1984–85. Seasonal changes in numbers, age-structure and reproduction are given for the three commonest species. Mus triton (36%), Praomys delectorum (29%), and Lophuromys flavopunctatus (19%). Total population numbers for L. flavopunctatus and P. delectorum were lowest at the end of the dry season, and highest at the end of the wet season; reproduction is seasonal, and most young are born in the wet season. Mus triton did not exhibit such large fluctuations in numbers, and reproduced in both wet and early dry seasons. Each species was found in several habitats, but L. flavopunctatus showed a strong preference for unburnt grasslands, M. triton for burnt grasslands, and P. delectorum for forest and tangle habitats. Each species had a different population survival rate, and few individuals survived for more than 5–6 months. Trapped-revealed home ranges were about 3100 m² for L. flavopunctatus , and 2700 m² for M. triton . Monthly changes in grass biomass, cover and height were recorded and related to the numbers of small mammals. General features of the small mammals of African mountains, based on data from the Drakensberg, Ethiopia and this study. are presented.     

Changes in arthropod diversity along a land use driven gradient of shrub cover in savanna rangelands: identification of suitable indicators

Shrub encroachment linked to heavy grazing has dramatically changed savanna landscapes, and is a major form of rangeland degradation. Our understanding of how shrub encroachment affects arthropod communities is poor, however. Here, we investigate the effects of shrub encroachment on abundance and diversity of ground-dwelling (wingless) arthropods at varying levels of shrub cover in the southern Kalahari. We also ascertain if invertebrate assemblage composition changes with habitat structure and identify which aspects of habitat structure (e.g., grass cover, herbaceous plant cover, shrub density) correlate most strongly with these changes. Ant, scorpion and dung beetle abundance increased with shrub cover, whereas grasshoppers and solifuges declined. Spider and beetle abundance exhibited hump-shaped relationships with shrub cover. RTU richness within orders either mirrored abundances, or exhibited no trend. Shrub density was the habitat component most correlated with similarities between invertebrate assemblages. Ground-dwelling arthropods showed clear shifts in species assemblage composition at a similarity level of 65% according to shrub density. Changes in indicator species showed that within the Tenebrionidae (darkling beetles), certain species respond positively to shrub thickening, replacing other species within the Family. Small-bodied, wingless Scarabaeidae (dung beetles) tended to increase with increased shrub density and three species emerged as significant indicators of more thickened habitats, although this might be a response to greater dung availability, rather than habitat structure itself. We conclude that because ground-dwelling invertebrates showed such clear responses in species assemblage composition, they present excellent candidates for use as indicator species in further studies into bush encroachment.     

Effect of grazing on plant patterns in arid ecosystems of Patagonian Monte

Our objective was to assess the relationship between the spatial patterning of perennial grasses (total, grazed, and non‐grazed) and shrub patches in rangelands under different grazing pressures of the Patagonian Monte. We selected three grazed paddocks with the usual stocking rate for the area, where previous studies showed that a piosphere formation is common. At each paddock, we analysed the grain of heterogeneity at sites located at two distances from the single watering point (near, far), using high‐resolution aerial photographs. At these sites, we also assessed in the field the density, size, cover, and spatial patterning of grazed and non‐grazed perennial grasses and shrub patches. The grain of heterogeneity of shrub patches was coarser in sites near the watering point than in those distant from it, as a consequence of the increase in size of both, bare soil and shrub patches. Field sampling showed that a coarse grain of heterogeneity relative to fine‐grained sites resulted from changes in species composition, increased bare soil areas and reduced perennial grass cover. In coarse‐grained sites, lower perennial grass cover resulted from lower density and/or smaller size of grass bunches than in fine‐grained sites. We did not find significant differences among sites in the proportion of perennial grazed grasses. Since the density and cover of perennial grasses was higher in fine‐ than in coarse‐grained sites, we suggested that fine‐grained sites are more important as feeding stations than coarse‐grained sites. The consequences of this differential use could lead to degradation of fine‐grained sites and to higher homogeneity in spatial plant structure and floristic composition within paddocks with respect to the condition observed at present, increasing the size of the highly degraded zone within the piosphere. At the patch level, we found that at about one third of the sampled transects, both total and non‐grazed perennial grasses were spatially aggregated with shrub patches. However, in most transects grazed perennial grasses were indifferently distributed in relation with shrub patches, showing that grazers display high selectivity of foraging sites at macro level (i.e. high and low grazing pressure sites at the paddock level), but random occupancy of vegetation units (randomness in the distribution of grazed perennial grasses at the patch level). The intensity of the positive association between non‐grazed grasses and shrub patches was higher in fine‐grained than in coarse‐grained sites and may be attributed to higher protection against herbivores associated to denser shrub patches in fine‐ relative to coarse‐grained sites. We concluded that a feedback exists between the spatial distribution of species preferred by grazers and the spatial patterning of use of these species.     

Movement patterns of Cephalopholis cruentata in a marine reserve in St Lucia, W.I., obtained from ultrasonic telemetry

Home range size and shape, and within home range movement patterns were determined from 10 adult graysby Cephalopholis cruentata implanted with acoustic transmitters on two reefs within the Soufriere Marine Management Area, St Lucia. The mean home range area was 2120 m²(range 1200 to 4000 m²) and non‐circular in shape. All fish demonstrated a clear preference for a specific area in their home range where, on average, 72·9% of an individuals position were located. Time spent in this favoured area was correlated with reef complexity. Fish with greater activity and displacement possessed a larger home range. Activity and displacement were higher by night than by day, and consistent with this, fish spent less time in the preferred area of their home range by night than by day. Tagged individuals demonstrated a strong tendency to avoid sand habitat present at reserve boundaries.     

Consequences of anthropogenic disturbances on soil seed bank diversity and nurse shrub effect in a semiarid rangeland

What is the effect of common land use histories on the diversity, richness, spatial distribution and abundance of the soil seed bank (SSB)? Does the effect change between different microsites under shrub canopy? To address these questions we selected seven sites with different grazing and fire histories in the Patagonian Monte desert. We took soil samples in seven microsites at different distances of the trunk along a windward/leeward transect through shrub patches to estimate the SSB of perennial grasses, shrubs and annuals. Shrubs SSB was scarce. The nurse shrub effect on perennial grass SSB was evident at sites with a low disturbance intensity (as sites without continuous grazing), and higher on the leeward side under the shrub canopy. High disturbance intensity (such as permanent grazing) promoted a decrease in perennial grass SSB and an increase in annuals SBB (especially non-native). Land use histories related to fire showed a moderated disturbance response (medium values for both groups). Differences between land use histories varied depending on the windward/leeward microsite from which the SSB was analysed. Our results suggest a nurse effect of the shrub patches on the spatial distribution and abundance of the SSB, but this effect decrease under continuous grazing. Fire seems to have a positive effect on perennial grass SSB. Nevertheless, fire characteristics must be controlled. Moreover, perennial grass SSB was almost depleted under continuous grazing, driving the grazed system towards a vulnerable state; and annual non-native species take advantage of this disturbance and dominate the SSB.     

Testing the grass-fire cycle: alien grass invasion in the tropical savannas of northern Australia

Abstract. Invasive alien grasses can increase fuel loads, leading to changes in fire regimes of invaded ecosystems by increasing the frequency, intensity and spatial extent of fires. Andropogon gayanus Kunth. (Gamba grass), a tall perennial grass from Africa, is invading ecosystems in the Top End of northern Australia. To determine whether A. gayanus alters savanna fire regimes, we compared fuel loads and fire intensities at invaded sites with those from native grass savannas. Savanna invaded by A. gayanus had fuel loads up to seven times higher than those dominated by native grasses. This higher fuel load supported a fire that was on average eight times more intense than those recorded in native grass savannas at the same time of year (means 15700 ± 6200 and 2100 ± 290 kW m⁻¹, respectively), and was the highest early dry season fire intensities ever recorded in the Northern Territory. These results suggest that A. gayanus is a serious threat to northern Australia's savannas, with the potential to alter vegetation structure and initiate a grass-fire cycle.     

Movement Patterns and Home Range Of The Bongo (Tragelaphus Eurycerus) In The Rain Forest Of The Dzanga National Park, Central African Republic

Bongos ( Tragelaphus eurycerus Ogilby) were studied for 8 months in the Dzanga National Park, Central African Republic. Tracks were followed and mapped with a compass and a pedometer to study movement patterns and home range. Natural licks were shown to be central points in the home range of the bongos: they visited the licks recurrently to consume soil, but also to forage on grass and herb species, and for social reasons. Forest areas far from licks were used much less than forest areas close to licks. When a lick was visited, distances between two resting sites were longer than in the forest without lick visits, caused by a direct and straight movement from the denser forest areas toward a lick. The study area of about 150 km² was presumably occupied by two groups of bongos. One of them seemed to split temporarily into two subgroups. Groups were not larger than 10–20 individuals. Estimated home ranges were at least 49 km² and 19 km² for the two groups, respectively. Estimated density in the Dzanga National Park was 0.25 bongos per km^2. This study shows the importance of natural licks for the largest social forest antelope, the bongo, and provides information which is important for its future conservation.     

Plant palatability and disturbance level in aquatic habitats: an experimental approach using the snail Lymnaea stagnalis (L.)

1. The palatability of aquatic macrophytes to the snail Lymnaea stagnalis was investigated in the laboratory. Eight species of macrophyte were selected from habitats that differed in either flood disturbance regime or nutrient status.
2. In a non-choice test, single macrophyte species were offered to individual snails. The average amount of plant dry mass consumed per Lymnaea dry mass ranged from 3.6 ± 1.4 (±SE) to 63.6 ± 13.9 mg g^–1 day^–1 across plant species. In a choice test, all eight plant species were presented simultaneously to sets of five snails. The average total consumption was 66.1 ± 3.8 mg g^–1 day^–1 and the maximum average consumption for a single plant was 26.2 ± 3.6 mg g^–1 day^–1.
3. In both tests, the amount consumed by snails differed significantly between the plant species. The species growing in undisturbed habitats were the least consumed. Habitat nutrient status was unrelated to plant palatability.
4. These results suggest that macrophyte species growing in habitats that are rarely disturbed by floods allocate a greater proportion of their resources to resisting herbivory.     

Pattern of movements of adult Barbus haasi in a small Mediterranean stream

Most adult Barbus haasi , at a 1950 m-long site in Vallvidrera creek, were highly sedentary and resided within a home range <20 m (32 m²), while a small group were more mobile. On successive sampling occasions, between 52·3 and 64·9% of fish were recaptured in the same 10-m-long section in which they had previously been captured. Movements over long distances were infrequent, and only 5·6% of the fish moved >100 m. The movement pattern of the population was seasonally stable, although the mean distances were slightly greater in summer because of the medium-range movements of a few individuals. Overall, upstream and downstream movements were equally common but a significant downstream movement occurred in spring. The size of the fish did not influence the movement rate. Fish inhabiting those sections of the stream with greater depth, slower current and more cover had a lower movement rate than fish occupying shallower, exposed sections. The restricted movement of B. haasi could increase the survival of fish by increasing the probability of staying in the remaining pools during summer dry-out, and may therefore be of adaptive significance.     

Inbreeding depression and family variation in a social insect, Bombus terrestris (Hymenoptera: Apidae)

Arid and semi-arid ecosystems often exhibit diverse plant growth forms in water-limited environments, but it is unclear whether resource competition (interference) is actually important in structuring communities. We chose a diverse Chihuahuan desert shrubland to examine the response of the plant community to experimental removals of selected perennial plant species or groups of species. Four treatments involved the removal of all individuals of all species of a single functional group (functional group removals: shrub removal, succulent removal, subshrub removal, perennial grass removal). Three other treatments involved removing species within functional groups. These seven treatments plus a control (no plants removed) were replicated six times each in 25×25 m experimental plots, in summer 1995. Permanent belt transects were surveyed for number and sizes of all vascular plants in spring and fall in 1997, 1999, 2000, and 2001. Those plots from which the dominant shrub, Larrea tridentata , was removed had not recovered in total plant cover or volume by 2001, but cover and volume in all other treatments were similar to those in control plots. Relatively few species demonstrated a positive response to the removal of other species or functional groups. The perennial grass group and forbs were the most responsive; perennial grass cover increased in the shrub removal treatment relative to the control but treatment differences diminished after dry growing seasons in 2000 and 2001. Results over the first five years suggest that either environmental conditions or intrinsic biological characteristics limit the ability of most plant species to respond to the removal of substantial fractions of community biomass and composition in the short term. Such slow response by both dominant and less abundant components of the community has implications for the recovery of semi-arid systems after human disturbance or other events leading to the reduction of biological diversity.     

Land use,rangeland degradation and ecological changes in the southern Kalahari,Botswana

Dual‐scale analyses assessing farm‐scale patterns of ecological change and landscape‐scale patterns of change in vegetation cover and animal distribution are presented from ecological transect studies away from waterpoints, regional remotely sensed analysis of vegetation cover and animal numbers across the southern Kalahari, Botswana. Bush encroachment is prevalent in semi‐arid sites where Acacia mellifera Benth. is widespread in communal areas and private ranches, showing that land tenure changes over the last 40 years have not avoided rangeland degradation. Herbaceous cover is dominated in intensively grazed areas by the annual grass Schmidtia kalahariensis Stent and in moderately grazed areas by the perennial grass Eragrostis lehmanniana Nees. Nutritious perennial grass species including Eragrostis pallens Hack. Ex Schinz remain prevalent in Wildlife Management Areas. Other ecological changes include the invasion of the exotic Prosopis glandulosa Torr. and dense stands of Rhigozum trichotomum Kuntze. in the arid southwest. Regional patterns of wildlife species show that the expansion of cattleposts and fenced ranches has led to large areas of low wildlife conservation value even in areas where cattle production is not practiced. Findings show the need for integrated landscape‐scale planning of land use if the ecological value and biodiversity of the southern Kalahari is to be retained.     

Aeolian process effects on vegetation communities in an arid grassland ecosystem

Many arid grassland communities are changing from grass dominance to shrub dominance, but the mechanisms involved in this conversion process are not completely understood. Aeolian processes likely contribute to this conversion from grassland to shrubland. The purpose of this research is to provide information regarding how vegetation changes occur in an arid grassland as a result of aeolian sediment transport. The experimental design included three treatment blocks, each with a 25 × 50 m area where all grasses, semi-shrubs, and perennial forbs were hand removed, a 25 × 50 m control area with no manipulation of vegetation cover, and two 10 × 25 m plots immediately downwind of the grass-removal and control areas in the prevailing wind direction, 19° north of east, for measuring vegetation cover. Aeolian sediment flux, soil nutrients, and soil seed bank were monitored on each treatment area and downwind plot. Grass and shrub cover were measured on each grass-removal, control, and downwind plot along continuous line transects as well as on 5 × 10 m subplots within each downwind area over four years following grass removal. On grass-removal areas, sediment flux increased significantly, soil nutrients and seed bank were depleted, and Prosopis glandulosa shrub cover increased compared to controls. Additionally, differential changes for grass and shrub cover were observed for plots downwind of vegetation-removal and control areas. Grass cover on plots downwind of vegetation-removal areas decreased over time (2004-2007) despite above average rainfall throughout the period of observation, while grass cover increased downwind of control areas; P. glandulosa cover increased on plots downwind of vegetation-removal areas, while decreasing on plots downwind of control areas. The relationships between vegetation changes and aeolian sediment flux were significant and were best described by a logarithmic function, with decreases in grass cover and increases in shrub cover occurring with small increases in aeolian sediment flux.     

Structure and composition of benthic macroinvertebrates of a tropical forest stream, River Nyamweru, western Uganda

Benthic invertebrates from River Nyamweru, a tropical forest stream in western Uganda were sampled bimonthly between April and December 1997 using a modified Hess Sampler. A total of 3708 benthic fauna from thirteen taxonomic orders were collected. Benthic samples were dominated by Diptera (mainly Chironomidae) representing over 60% of all the organisms, followed by Ephemeroptera. Benthic invertebrate densities ranged from 63 ± 9.03 organisms/m² to 300 ± 33.36 organisms/m², with higher densities occurring during the dry season and lower densities during the wet season. The benthic community structure in River Nyamweru reflected mainly collectors and scrapers as the most important groups (83%), while predators were very rare (3.1%). River discharge influenced benthic abundance, with more invertebrates at lower discharge and fewer invertebrates at higher discharge. The applicability of the River Continuum Concept to tropical forest stream situation is discussed.     

Effects of regulated discharge on burbot migration

Burbot Lota lota movement and river discharge were studied in the Kootenai River, Idaho, U.S.A. and British Columbia, Canada, downstream of Libby Dam, Montana, U.S.A. A total of 24 adult burbot with transmitters were tracked from 1994 to 2000, for analysis of a travel distance of ≥5 km in ≤10 days termed 'stepwise movement'. Of 44 'stepwise movements', significantly greater movements during pre‐spawning and spawning were observed when average daily discharges from Libby Dam were <300 m³ s⁻¹, with a mean of 176 m³ s⁻¹, similar to pre‐dam conditions. Burbot travelled at a greater rate during all seasons (3·36 km day⁻¹) at discharges >300 m³ s⁻¹(mean = 1·84 km day⁻¹) than at discharges >300 m³ s⁻¹ but no difference was found for the pre‐spawning and spawning period. Burbot that started 'stepwise movements' in low discharge conditions frequently stopped during low discharges.