Phytosociology and ecology of old hay meadows in Hordaland,western Norway in relation to management

Well-drained mown hay meadows in Hordaland, western Norway, were investigated. The hay meadows are either managed in a traditional or in a semi-traditional way. Traditional management in the area involves little or no use of manure or fertilizer, mowing once or twice a year with a first cut later than June 23, clearing in spring and intensive grazing for short periods in spring and autumn. Semi-traditional management involves use of small to medium quantities of commercial fertilizer, often in addition to manure. Grazing may be irregular, light or lacking.Eight units of hay meadow vegetation are described, all assigned to the class Molinio-Arrhenatheretea, order Arrhenatheretalia; four units to the Atlantic alliance Cardaminion pratensis, association Cardamino pratensis-Conopodietum majoris (with three subassociations); one unit to the alliance Arrhenatherion elatioris, three units are provisionally grouped as Galium uliginosum-Knautia arvensis meadows.Continuous management of these meadows has resulted in characteristic species compositions, which vary along a west-east climatic gradient, and are related to the amount of commercial fertilizer used and the natural mineral-content of the soil.These hay meadows seem to be best maintained by continuing the traditional management regime. Grazing in spring and in autumn reduces fast growing dominants and creates openings for new seedlings of annuals and biennials. A late first cut allows for ripening of the seeds and creates openings for light for the species of the lower field layer. A list of species characteristic of traditionally managed permanent hay meadows is presented, and possible effects of present changes in management on the species composition of the meadows are discussed.The amount of organic matter in the topsoil is small in most stands. The pH values are between 4.0 and 5.6. The highest values were recorded in soil where little or no commercial fertilizer is used, and where the subsoil is naturally rich in minerals. A tendency to Mg impoverishment in the best fertilized stands is demonstrated. Species-rich stands on Ca-rich soils are poor in P.The expected changes in vegetation, future use, and conservation of permanent hay meadows in Hordaland are discussed.     

The ecology of ruffe,Gymnocephalus cernuus (Pisces: Percidae) introduced to Mildevatn,western Norway

Synopsis Diet, habitat use, diel and seasonal activity and a number of population parameters were studied on ruffe,Gymnocephalus cernuus, introduced to Mildevatn, western Norway. This lake is sited outside the natural range of the ruffe and has a lower fish diversity and a different fish species composition than within its native range. From June through September the ruffe was planktivorous and mainly caught at 4 to 6 m depth in the benthic zone. At other times of year ruffe was feeding on zoobenthos and caught deeper in the benthic zone. Ruffe was mainly day active. Zooplankton feeding during summer is the clearest difference compared to ruffe populations living within its natural range. Presence of large zooplankton organisms available for ruffe is suggested as the main reason for the difference found in food choice. The availability of large zooplankton is probably due to community structure caused by a predator and lack of interspecific competition for zooplankton in the deeper parts of the lake. Piscivorous brown trout.Salmo trutta, restrict the habitat of threespined stickleback,Gasterosteus aculeatus, to the zone of littoral vegetation, allowing high densities of larger zooplankton species likeBythotrephes longimanus to be present in the lake. Brown trout is present only in the upper light and well oxygenated parts of the lake, leaving a refuge for the ruffe, where they can feed on the rich zooplankton community.     

Plant communities in broad-leaved deciduous forests in Hordaland county, Western Norway

The vegetation of broad-leaved deciduous forests in Hordaland, Western Norway is described with regards to species composition and structure. The investigation is based on phytosociological analyses of native forest stands dominated by Corylus avellana, Fraxinus excelsior, Ulmus glabra , or Tilia cordata. Two-way-indicator analysis (TWINSPAN) and correspondence analysis (CA) are used to distinguish different vegetation types and assess possible gradients in the vegetation data. The vegetation types are discussed in relation to equivalent forest communities and to syntaxonomical units. Two forest types are described at the first hierarchical TWINSPAN level: 1) A hygrophilous and slightly thermophilous Fraxinus excelsior-Cirriphyllum piliferum forest characterised by an open tree canopy, a dense field layer of tall herbs and ferns, and a high cover of bryophytes. 2) A thermophilous and less hygrophilous Corylus avellana-Brachypodium sylvaticum forest characterised by a dense tree layer, a more open field layer with larger elements of small herbs, and a somewhat lower bryophyte cover. The CA analysis clearly separates the samples from the first TWINSPAN division along the first ordination axis. Five forest types have been described at the second hierarchical level, mainly associated with differences in mesotrophic and eutrophic species, but there is clearly a gradient structure in the species composition between these plant communities. In relation to syntaxonomy, the first TWINSPAN division supports the separation of the west Norwegian broad-leaved deciduous forests into a hygrophilous plant community, the Eurhynchio-Fraxinetum (Blom 1982) Øvstedal 1985 and a drier and more thermophilous community, the Primulo-Ulmetum (Blom 1982) Øvstedal 1985.     

Phenology of Dryopteris affinis ssp. affinis and Polystichum aculeatum: modeling relationships to the climatic variables in a Mediterranean area

The phenology of leaves and stages from immature sori to spore release were studied in natural populations of two perennial herbaceous ferns, Dryopteris affinis ssp. affinis and Polystichum aculeatum, for 2 years in Italy. The Generalized Additive Model for Location, Scale and Shape (GAMLSS) was used to predict a particular phenological event from the climatic variables. Both fern species are evergreen with leaf lifespan of 13–14 months in D. affinis ssp. affinis and 15–24 months in P. aculeatum. Their leaf production is seasonal with most flushed in spring. In D. affinis ssp. affinis the decaying of old leaves is sudden whereas in P. aculeatum it is gradual, and in both types the decaying is accelerated just as the new leaves emerge. Temperature is the best predictor for the development of sori and spore release. The end of spore release and leaf emergence are positively affected by temperature and negatively affected by rainfall and snow cover for P. aculeatum. An almost similar response to climatic factors of emergent and senescent leaves and their phenology supportsthe hypothesis that old leaves serve as nutrient storage organs for new leaf growth. Comparison the phenological patterns between the 2 years indicated that the time lapses between each phenological event were the same within seasons for D. affinis ssp. affinis but show some differences for P. aculeatum. We also hypothesize that endogenous factors may play an important role in the phenology of P. aculeatum.     

Air humidity as key determinant of morphogenesis and productivity of the rare temperate woodland fern Polystichum braunii

(1) Most ferns are restricted to moist and shady habitats, but it is not known whether soil moisture or atmospheric water status are decisive limiting factors, or if both are equally important. (2) Using the rare temperate woodland fern Polystichum braunii, we conducted a three‐factorial climate chamber experiment (soil moisture (SM) × air humidity (RH) × air temperature (T)) to test the hypotheses that: (i) atmospheric water status (RH) exerts a similarly large influence on the fern's biology as soil moisture, and (ii) both a reduction in RH and an increase in air temperature reduce vigour and growth. (3) Nine of 11 morphological, physiological and growth‐related traits were significantly influenced by an increase in RH from 65% to 95%, leading to higher leaf conductance, increased above‐ and belowground productivity, higher fertility, more epidermal trichomes and fewer leaf deformities under high air humidity. In contrast, soil moisture variation (from 66% to 70% in the moist to ca. 42% in the dry treatment) influenced only one trait (specific leaf area), and temperature variation (15 °C versus 19 °C during daytime) only three traits (leaf conductance, root/shoot ratio, specific leaf area); RH was the only factor affecting productivity. (4) This study is the first experimental proof for a soil moisture‐independent air humidity effect on the growth of terrestrial woodland ferns. P. braunii appears to be an air humidity hygrophyte that, whithin the range of realistic environmental conditions set in this study, suffers more from a reduction in RH than in soil moisture. A climate warming‐related increase in summer temperatures, however, seems not to directly threaten this endangered species.     

The drift of zooplankton and microzoobenthos in the river Strandaelva,western Norway

The drift of zooplankton (rotifers, cladocerans, cyclopoid copepods) and microscopical zoobenthos (mainly bdelloid rotifers and small chironomid larvae) was investigated by filtering samples of river water. The number of drifting benthic rotifers varied between 1 000 and 6 000 ind. m^–3 in the lake inlet, and between 30 and 500 ind. m^–3 in the lake outlet, without any seasonal trend. The number of drifting insect larvae was approx. equal in the lake inlet and outlet, with a maximum in summer (250–300 ind. m^–3) and minimum in winter (ca. 10 ind. m^–3). Increasing water flow resulted in an increasing number of drifting zoobenthos. Downstream from the lake, the number of drifting benthic rotifers was increasing from approx. 300 ind. m^–3 in the outlet to 6 500 ind. m^–3 3.4 km downstream, while the number of insect larvae was ca. 100 ind. m^–3 in the outlet and leveled off at approx. 300 ind. m^–3 after 200 m. The number of drifting zooplankton in the lake outlet varied between 20 and 2 000 ind. m^–3 for crustaceans, and between 300 and 20 000 ind. m^–3 for rotifers, both with a maximum in late summer/autumn and a minimum in winter. The number of drifting zooplankton decreased by some 45% in the first 200 m from the lake outlet, but some zooplankton was still found in the drift 3.4 km downstream. The largest species was removed first from the drift. The diurnal variation in the number of drifting zooplankton in lake outlets appear to be related to the vertical migration in the lake, i.e. the largest number drifting when most animals are in the upper water layers.Contribution from the Voss Project, University of OsloContribution from the Voss Project, University of Oslo     

The comparative ecology and biogeography of parasites

Comparative ecology uses interspecific relationships among traits, while accounting for the phylogenetic non-independence of species, to uncover general evolutionary processes. Applied to biogeographic questions, it can be a powerful tool to explain the spatial distribution of organisms. Here, we review how comparative methods can elucidate biogeographic patterns and processes, using analyses of distributional data on parasites (fleas and helminths) as case studies. Methods exist to detect phylogenetic signals, i.e. the degree of phylogenetic dependence of a given character, and either to control for these signals in statistical analyses of interspecific data, or to measure their contribution to variance. Parasite–host interactions present a special case, as a given trait may be a parasite trait, a host trait or a property of the coevolved association rather than of one participant only. For some analyses, it is therefore necessary to correct simultaneously for both parasite phylogeny and host phylogeny, or to evaluate which has the greatest influence on trait expression. Using comparative approaches, we show that two fundamental properties of parasites, their niche breadth, i.e. host specificity, and the nature of their life cycle, can explain interspecific and latitudinal variation in the sizes of their geographical ranges, or rates of distance decay in the similarity of parasite communities. These findings illustrate the ways in which phylogenetically based comparative methods can contribute to biogeographic research.     

The altitudinal gradient of vascular plant richness in Aurland, western Norway

Differences in vascular plant species richness; along the altitudinal gradient in the Aurland area of western Norway have been investigated. Based on field surveys, as complete lists as possible of all vascular plants have been compiled for each 100 m altitudinal band, from sea level to the highest mountain (1764 m). For each of the 18 altitudinal bands, climatic data have been estimated. A total of 444 vascular plant species were recorded. Highest species richness (263 species) occurred in the 600–700 in band, whereas the uppermost band had only 10 species. There are minor differences in species number between the altitudinal bands < 1000 m. Partial least squares regression shows that species richness for the overall altitudinal gradient is well predicted by mean July and January temperatures and mean annual precipitation. Species turnover is highest in the 100–200 m. 600–700 m. and 1400–1500 m altitudinal bands. In terms of the gradient in summer temperature, the study supports the generally assumed linear relationship between July temperature and the number of vascular plant species between 700 and 1500 m corresponding with a mean July temperature range of 7–11°C. The study shows a decrease of ca 30 vascular plant species with a 1°C decrease in mean July temperature, and that the “climatic vascular plant limit” is here estimated to occur at a mean July temperature of 2.4°C. Above 1500 and below 700 m. species number is lower than expected based on summer iemperature conditions alone. The 700–800 m band represents the highest floristic difference compared to the other bands. Ordination and classification analyses of the floristic compositional data of all the bands highlight the 600–800 and 1500–1600 m altitudinal bands as the major biotic boundaries along the gradient. No major discontinuity in species richness, composition, or turnover was consistently found, however, at the 1100–1200 m band representing the forest-limit ecotone in Aurland.     

Polystichum lilianae sp. nov. (Dryopteridaceae) and its relationships to P. fournieri and P. turrialbae

Morphological, isozyme, and cytological analyses of the small, pale-scaled polystichums from oakdominated montane rain forests in Costa Rica and Mexico reveal the presence of a separable undescribed species endemic to the Cordillera de Talamanca of Costa Rica. The new taxon,Polystichum lilianae, is an allotetraploid hypothesized to have the sympatricP. turrialbae as one diploid progenitor based on isozyme characters. The isozyme and morphometric data also support the inclusion ofP. smithii, described from southern Mexico, inP. turrialbae, described from Costa Rica. The namePolystichum fournieri, formerly used for all of these plants, applies to species endemic to Oaxaca and Chiapas, Mexico. It is not a progenitor ofP. lilianae.     

The ecology of ostracodes (Ostracoda,Crustacea) in western Mongolia

Aquatic biota in Central Asia witnesses and faces a changing environment. Because ostracodes contribute to both extant and fossil lacustrine diversity, they can be used to track evolution in water quality. Living ostracode communities in a variety of aquatic habitats of western Mongolia were analyzed in relation to environmental and hydrochemical variables of those habitats, based on presence/absence data from net samples. The sampled water bodies represent broad gradients in ionic concentration and composition. Ostracode community composition of springs differed from all other sampled habitats, which was also reflected in CCA analyses. Our data indicated that the ostracode fauna of western Mongolia shows a high response to calcium content, alkalinity, salinity, temperature, nutrients, and altitude. Species composition and diversity seem to be determined by solute evolution dominated by Ca-depletion. In general, this seems to be an important regulator of lacustrine ostracode species composition in several semi-arid regions. Binary logistic regression was used to predict the occurrence of the most common species with a minimal set of environmental variables. The resulting models showed on average low performance, and mainly demonstrated the potential of such modeling to predict the distribution of typical bio-indicator species.     

Population ecology of polar bears at Svalbard, Norway

The population ecology of polar bears at Svalbard, Norway, was examined from 1988 to 2002 using live-captured animals. The mean age of both females and males increased over the study, litter production rate and natality declined and body length of adults decreased. Dynamics of body mass were suggestive of cyclical changes over time and variation in body mass of both adult females and adult males was related to the Arctic Oscillation index. Similarly, litter production rate and natality correlated with the Arctic Oscillation index. The changes in age-structure, reproductive rates and body length suggest that recovery from over-harvest continued for almost 30 years after harvest ended in 1973 and that density-dependent changes are perhaps being expressed in the population. However, the variation in reproduction and body mass in the population show a relationship between large-scale climatic variation and the upper trophic level in an Arctic marine ecosystem. Similar change in other polar bear populations has been attributed to climate change, and further research is needed to establish linkages between climate and the population ecology of polar bears.     

Life-history evolution in guppies. VII. The comparative ecology of high- and low-predation environments

Prior research has demonstrated a strong association between the species of predators that co-occur with guppies and the evolution of guppy life histories. The evolution of these differences in life histories has been attributed to the higher mortality rates experienced by guppies in high-predation environments. Here, we evaluate whether there might be indirect effects of predation on the evolution of life-history patterns and whether there are environmental differences that are correlated with predation. To do so, we quantified features of the physical and chemical environment and the population biology of guppies from seven high- and low-predation localities. We found that high-predation environments tend to be larger streams with higher light levels and higher primary productivity, which should enhance food availability for guppies. We also found that guppy populations from high-predation environments have many more small individuals and fewer large individuals than those from low-predation environments, which is caused by their higher birth rates and death rates. Because of these differences in size distribution, guppies from high-predation environments have only one-fourth of the biomass per unit area, which should also enhance food availability for guppies in these localities. Guppies from high-predation sites allocate more resources to reproduction, grow faster, and attain larger asymptotic sizes, all of which are consistent with higher levels of resource availability. We conclude that guppies from high-predation environments experience higher levels of resource availability in part because of correlated differences in the environment (light levels, primary productivity) and in part as an indirect consequence of predation (death rates and biomass density). These differences in resource availability can, in turn, augment the effect of predator-induced mortality as factors that shape the evolution of guppy life-history patterns. We found no differences in the invertebrate communities from high- and low-predation localities, so we conclude that there do not appear to be multitrophic, indirect effects associated with these differences in predation.     

The role of sugars in western spruce budworm nutritional ecology

Abstract.

• 1 The western spruce budworm, Choristoneura occidentalis Freeman, and Douglas-fir, Pseudotsuga menziesii (Mirb.) Franco, have been used to test the hypothesis that variation in levels of foliar sugars form part of the basis for plant resistance to herbivore attack.
• 2 Budworm population growth was evaluated on artificial diets with 2–45% sucrose using a three generation bioassay. Diets with 1.2% and 3.9% N were tested to determine if responses to sugar were dependent on levels of protein. The 3.9% N diets were supplemented with a mineral salt mixture, so they had high levels of N and minerals.
• 3 The response of budworm population growth to sucrose concentrations ≤20% was convex at 1.2% N and flat for 3.9% N. Population growth on the 1.2% N diet, which had levels of N and minerals similar to host foliage, was good with only 2% sugar, but optimal at the 6% sucrose level; the number of F₁, F₂ and F₃ larvae produced declined substantially when sugar was increased to 11% and 20%. On the 3.9% N diets, population growth was equivalent for diets with 2% and 11% sucrose. Sucrose concentrations ≥29% were detrimental to the budworm at both N levels.
• 4 Sugar concentrations in Douglas-fir foliage varied between 5.7% and 18.4%. Thus, results from the 1.2% N experiment indicated that budworm performance was best on diets with sugar concentrations near the lower limit observed for host foliage. This implies that plants with higher foliar sugar may be inferior hosts for the budworm. Field observations supported this conclusion, as putatively resistant Douglas-fir trees had significantly higher levels of sugars in their foliage than nearby paired susceptible trees. Variation in foliar sugars among individual trees may be part of the mechanism in Douglas-fir resistance to C.occidentalis damage.

     

The comparative germination ecology of nine Rumex species

The germination requirements, dormancy cycle and longevity of nine Rumexspecies were studied in field conditions and laboratory experiments to show theadaptations of the related species to their specific habitat. Within one genus,rather striking differences were observed in germination ecology. However, theclosely related species, R. acetosa and R.scutatus, are very similar: they fruit in early summer; theirseeds can germinate immediately after dispersal, and they are nondormant andshort-lived. R. acetosella also has fruits insummer, but the seeds do not germinate the first season after dispersal. Theyare long-lived, but buried seeds do not show a dormancy cycle; they mightgerminate in different seasons after exposure to light. Seeds of four species (R. conglomeratus,R. maritimus, R. sanguineus andR. crispus) are long-lived and undergo aseasonal dormancy cycle, with a low level of dormancy in winter and early springand a deep dormancy in summer as was already known for R.obtusifolius. These seeds are shed in the autumn, and they germinatemainly in the spring in consecutive years. R. maritimusalso germinates in summer and autumn on drying muddy soils. The seeds of R. hydrolapathum only germinate onwaterlogged soils, which explains its growth at the edge of streams and ponds.Its seeds are rather short-lived. The seeds of the species on very wetplaces require a higher temperature for germination.     

On the ecology of Thelypteris limbosperma in W Norway. The distribution in relation to climatic factors

In 31 selected districts in W Norway the vertical distribution of Thelypteris limbosperma has been recorded. Special attention has been paid to the area within which T. limbosperma occurs as a dominant in the vegetation. Both horizontal and vertical distribution areas are related to climatic factors, namely the mean temperature of the wannest and coldest months, precipitation and humidity conditions expressed by Martonnes aridity index. Correlations between the distribution of T. limbosperma and these parameters have been elucidated. In oceanic districts correlations exist between the vertical distribution limits and the mean temperature of the warmest month, t. max = 14.2°C and 10.3°C. In suboceanic districts correlations are with t. min = -5.3°C, P = 121 mm; and H = 87.0. In districts where H < 60 the fern is sparce or missing.     

Phylogeny, ecology, and the coupling of comparative and experimental approaches

Recent progress in the development of phylogenetic methods and access to molecular phylogenies has made comparative biology more popular than ever before. However, determining cause and effect in phylogenetic comparative studies is inherently difficult without experimentation and evolutionary replication. Here, we provide a roadmap for linking comparative phylogenetic patterns with ecological experiments to test causal hypotheses across ecological and evolutionary scales. As examples, we consider five cornerstones of ecological and evolutionary research: tests of adaptation, tradeoffs and synergisms among traits, coevolution due to species interactions, trait influences on lineage diversification, and community assembly and composition. Although several scenarios can result in a lack of concordance between historical patterns and contemporary experiments, we argue that the coupling of phylogenetic and experimental methods is an increasingly revealing approach to hypothesis testing in evolutionary ecology.     

A comparative cytogenetic analysis of five pine species from North America, Pinus banksiana, P. contorta, P. monticola, P. resinosa, and P. strobus

The genus Pinus comprises more than 100 species, which are widely distributed in the Northern hemisphere. Cytogenetic information on North American pines is very limited despite their economic importance. In the present study, a detailed comparative cytogenetic analysis is presented for five pine species from North America, P. resinosa, P. monticola, P. contorta, P. banksiana, and P. strobus. Morphometric analysis and physical mapping of rDNA probes were performed. The karyotype of P. monticola was considered ancestral with small difference in relative chromosome lengths. P. banksiana, P. contorta, and P. strobus karyotypes were considered semi-asymmetrical and less ancestral type. P. banksiana showed five secondary constrictions, P. strobus six, and P. contorta eight. P. resinosa karyotype was semi-asymmetrical and derived with 14 secondary constrictions identified on eight different chromosomes. Karyological data were consistent with molecular cytogenetic information. A significant association was observed between the number and locations of secondary constrictions and the 18S-5.8S-26S rDNA sites detected using fluorescence in situ hybridization. The two methods were used to establish a reliable comparative karyotype of the selected pines. In general, karyotype and chromosome evolution were not related to genetic relationships among pine species studied.     

Ecological studies on the phytoplankton of Boknafjorden, western Norway. II. Environmental control of photosynthesis

Both predicted (incubator) and measured (in situ) ¹⁴C-assimilationrates were studied from February to November 1981 at three stationsin Boknafjorden, a deep silled fjord of western Norway. Sampleswere taken from different light depths within the euphotic zone.A high degree of conformity was found between the two approaches.Daily values of carbon assimilation integrated over the euphoticzone varied between 0.05 and 1.4 g C m^–2. Yearly primaryproduction varied between stations from 82 to 112 g C m^–2(120–148 g C m^–2 when based on average light conditions).The light-saturation curve parameters ^B and P^B_max ranged from0.0056 to 0.0537 mg C mg Chla^–1 h^–1 µE^–1m² and from 0.7 to 8.5 mg C mg Chla^–1 h^–2 (in situassimilation numbers ranged from 0.9 to 9.3 mg C mg Chla^–1h^–1) respectively, which compare well with those publishedfrom the northwestern side of the Atlantic. The overall importanceof light in controlling photosynthesis throughout the year wasrevealed by the light utilization index , estimated to be 0.43mg C mg Chla^–1 E^–1 m². The maximum quantum yieldwas encountered on August 17, with 0.089 mol CE^–1. Chla/Cratios above and below 0.010 were found to be typical for shade-and light-adapted cells respectively. Assimilation numbers andgrowth rates were linearly related only when considering light-adaptedcells. Consistent with the findings of this study, the applicabilityof I_K, ^B and P^B_max as indicators of light-shade adaptation propertiesshould be questioned. Maximum growth rates were encounteredduring an autumn bloom of the dinoflagellate Gyrodinium aureolum(1.0 doublings day^–1), while 0.7–0.8 doublings day^–1were found for a winter bloom (water temperature of 2°C)of the diatom Skeletonema costatum. No unambiguous temperatureeffect on assimilation number and growth of phytoplankton couldbe recognized in Boknafjorden. A tendency towards increasedassimilation numbers coinciding with increased water columnstability was revealed. The highest P^B_max values were oftenencountered at almost undetectable nutrient concentrations.At least during summer this could be attributed to recyclingof nutrients by macro- and/or microzooplankton, responsiblefor a greater part of the primary production now being grazeddown. This study supports the convention that the depth of theeuphotic zone may extend considerably below the 1% light depth.     

Vegetation history of Molinia heaths in Tysvær, Rogaland, western Norway

Investigations were undertaken in an area of western Norway, approximately 30 km from the coast. The area is dominated by Molinia caerulea heaths on gentle slopes of silt-dominated glacial deposits. Pollen analyses were made on 3 soil profiles. Each profile could be divided into 2 local pollen assemblage zones (PAZs) corresponding with a change from dry to damp soils. From a comparison of the fossil pollen assemblages with pollen assemblages from surface samples collected from present-day Molinia heaths, and a radiocarbon date, the Molinia heaths can be shown to have originated about 1900 years BP. The persistence of the Molinia heaths is probably related to the high water-holding capacity of the sediments and the influence of human activity.