Moderation of genetic and environmental influences on diurnal preference by age in adult twins

Diurnal preference changes across the lifespan. However, the mechanisms underlying this age-related shift are poorly understood. The aim of this twin study was to determine the extent to which genetic and environmental influences on diurnal preference are moderated by age. Seven hundred and sixty-eight monozygotic and 674 dizygotic adult twin pairs participating in the University of Washington Twin Registry completed the reduced Morningness–Eveningness Questionnaire as a measure of diurnal preference. Participants ranged in age from 19 to 93 years (mean?=?36.23, SD?=?15.54) and were categorized on the basis of age into three groups: younger adulthood (19–35 years, n?=?1715 individuals), middle adulthood (36–64 years, n?=?1003 individuals) and older adulthood (65+ years, n?=?168 individuals). Increasing age was associated with an increasing tendency towards morningness (r?=?0.42, p?<?0.001). Structural equation modeling techniques parsed the variance in diurnal preference into genetic and environmental influences for the total sample as well as for each age group separately. Additive genetic influences accounted for 52%_[46–57%], and non-shared environmental influences 48%_[43–54%], of the total variance in diurnal preference. In comparing univariate genetic models between age groups, the best-fitting model was one in which the parameter estimates for younger adults and older adults were equated, in comparison with middle adulthood. For younger and older adulthood, additive genetic influences accounted for 44%_[31–49%] and non-shared environmental influences 56%_[49–64%] of variance in diurnal preference, whereas for middle adulthood these estimates were 34%_[21–45%] and 66%_[55–79%], respectively. Therefore, genetic influences on diurnal preference are attenuated in middle adulthood. Attenuation is likely driven by the increased importance of work and family responsibilities during this life stage, in comparison with younger and older adulthood when these factors may be less influential in determining sleep–wake timing. These findings have implications for studies aimed at identifying specific non-shared environmental influences, as well as molecular genetic studies aimed at identifying specific polymorphisms associated with diurnal preference.     

Habitat preference and habitat exploration in two species of satyrine butterflies

We compared the habitat preference and the mobility of different populations of primarily forest living, satyrine butterflies by observing them in a manipulated environment, containing different habitat types inside a large outdoor cage. The cage was 30 m long and divided into three parts. Each end of the cage contained a shady habitat, made to be similar to the floor of a forest with sunlit spots. In between these shady parts, there was a 14 m long open part, like a glade or a meadow. We performed two studies in the cage. In the first study, we compared a population of a species with very restricted distribution in Sweden, Lopinga achine , and a population of a widely distributed and expanding species, Pararge aegeria . We found that the two species had similar habitat preferences in the cage, but that L. achine had a lower rate of dispersal from one shady part of the cage to the other. In the second study, we compared two populations of P. aegeria , a Swedish forest living population and a Madeiran population inhabiting a fairly open habitat. We found that, although individuals of the Swedish population avoided staying in the open, middle part of the cage, they dispersed between the shady parts of the cage at a higher rate than individuals of the Madeiran population. We argue that studies of the kind we performed can serve the purpose of testing hypotheses about habitat preference and mobility that derive from other field work or from information about distribution patterns.     

Dispersal characteristics of three odonate species in a patchy habitat

Dispersal has a potentially profound effect on the dynamics of populations especially when a population occupies a patchy habitat. Ponds surrounded by terrestrial landscape are an example of patchy distribution of physical conditions and constitute "islands" for odonates. Few studies have focussed on dispersal in odonates. We have used the direct method of dispersal observing (capture-mark-recapture technique) in order to estimate the degree of linkage in three patchy populations of odonate localised on three ponds. We also examined the differences in dispersal ability within and among three species ( Coenagrion puella , Coenagrion scitulum and Libellula depressa ). The ponds were situated in southwest France on a limestone plateau. In this arid area, these ponds constitute the only surface water available and are relatively sparsely distributed. The size of the ponds ranged from 48 to 79 m² and they were 200 and 775 m apart. We demonstrated that three factors influence the dispersal ability of these odonates. The first is represented by the abiotic factors and especially weather conditions. This determines the number of days that dispersal is possible. The second is interspecific differences. We showed that sensitivity to weather conditions, species size and species behaviour influence dispersal ability. The third factor is the intraspecific characteristics. We demonstrated that there are differences in dispersal ability according to sex and age. To conclude, we discuss the importance of pond management to maintain the existing odonate populations and to facilitate introduction of new populations in this region where little exchange occurs between ponds.     

Thermal preference in the pool frog Rana lessonae: impact on the reproductive behaviour of a northern fringe population

A documented warmth preference in the pool frog Rana lessonae is also expressed in the reproductive behaviour of a northern fringe population (60°N). During the breeding period of 31 days, individually marked frogs were observed during chorus. Male choruses were formed in the warmest parts of the pond, but moved between three areas. At night, these areas maintained the highest water temperatures in the pond, especially at low temperatures prevailing in the beginning of the breeding period. Oviposition occurred in the current chorus areas, and 86% took place during the first two weeks of the breeding period. That the choruses and oviposition changed locations within the favourable breeding areas can be explained by a pronounced thermal preference in the frogs, and winds moving the warm surface water in different directions.
The water temperature required to initiate oviposition was 16°C, which is similar to those reported from related more southern populations. The thermal preference expressed in the breeding behaviour of both sexes and the tendency for early oviposition may be behavioural responses to environmental constraints on reproduction in the short season at 6O°N.     

Molecular phylogeographic evidence for multiple shifts in habitat preference in the diversification of an amphipod species

An earlier study of ours demonstrated polygenic control of habitat preference between sympatric populations of an estuarine amphipod (Stanhope et al. 1992). Knowledge of the ecological history in this estuary suggested that two new and distinct habitat types (wood debris and Fucus ) had been formed immediately adjacent to each other, in an area which was previously the habitat type common to the rest of the estuary (bank). This suggested the possibility that the ancestral population had been split into two resource specialists (disruptive selection on habitat preference). The genetic relatedness of these three populations (the proposed ancestor and the two proposed descendants) and six others occupying the same three habitat types were investigated on a regional geographic scale, using an extensive set of genomic DNA RFLPs. These data were combined with measures of habitat preference (including genetic tests) in the additional populations. A very strongly supported phylogeographic tree, unequivocally supported a shift in habitat preference in the wood-debris population of this estuary. The data did not, however, support the hypothesis of disruptive selection on the ancestor (bank) with a consequent split into two resource specialists. Instead, it clearly indicated that the occupants of the Fucus habitat type were members of a habitat specific race, and thus their presence in this estuary reflected the expansion of an old resource base. Furthermore, the combined RFLP and habitat preference data, for all nine populations, indicated that the same polygenically based shift in habitat preference that had occurred in the original estuary had occurred independently, in another estuary, 700 km removed, that had similar environmental circumstances.     

Effects of habitat complexity on trophic interactions of three congeneric fish species

Aquatic Ecology - Habitat complexity can substantially alter trophic relationships, such as competitive and predatory interactions, between fish species. This study aimed to evaluate how trophic...     

Effect of habitat structure on positional behavior and support use in three species of lemur

This study compares locomotor and postural behavior and substrate use of three species of lemur, the diademed sifaka (Propithecus diadema edwardsi), the brown lemur (Eulemur fulvus rufus), and the red-bellied lemur (Eulemur rubriventer) at two different localities within Ranomafana National Park. The object of the study is to see if there are quantitative differences in the behavior of the lemurs or their choice of substrates in forests that have different structural attributes. Analysis of the physical characteristics of the habitat demonstrates that compared to the Talatakely area, the forest at Vatoharanana has a higher proportion of larger, taller trees. The behavior of the lemurs also differs in the two areas: all species leap less and climb and move quadrupedally more at Vatoharanana. All species use small size supports less frequently at Vatoharanana, choosing insted medium size supports (all three species) or tiny supports (Propithecus andE. fulvus) found in tree crowns and terminal branches. The lemurs prefer (i.e. use more often than would be expected based on abundance) large trees at both sites. At Vatoharanana however, they are more frequently observed higher in trees and in taller trees with greater trunk breadth. The differences in locomotor behavior are in part due to the fact that at Vatoharanana, more bouts are collected during feeding and foraging than during travel. The relationship between this difference in activity pattern and the structural differerences in the two forests, however, is not clear. This study points out the need for longer term field studies of positional behavior and substrate use that incorporate the variety of forest types the subject species inhabit.     

Dynamics of antibacterial activity in three species of Caribbean marine algae as a function of habitat and life history

Antibacterial activity of lipid extracts from three species of Caribbean marine algae, Spyridia filamentosa, S. hypnoides and Wrangelia bicuspidata was evaluated monthly for one year. Activities were tested for whole plant extracts and TLC-separable zones. Whole plant extracts demonstrated monthly variability in activity with respect to both habitat and life history phase in addition to periods of similarity. No consistency was seen regarding activity against different microorganisms. TLC analyses of the extracts led to the identification of twenty-seven chromatographically distinct regions (TLC zones) each from both S. filamentosa and S. hypnoides and twenty-five from W. bicuspidata, which demonstrated antimicrobial activity. Between these species, twenty-one active TLC zones appeared to be shared based on their similar chromatographic characteristics. Individual TLC zones also demonstrated variable activity throughout the sampling period with respect to habitat and life history phase as well as periods of similarity. Algal antibiosis in these species is recognized as being highly complex, involving numerous chemical compounds, each of which is highly variable in terms of its presence and/or probable concentration.     

Synergistic effects of habitat preference and gregarious behaviour on habitat use in coral reef cardinalfish

Spatial distributions of coral reef fish species are potentially determined by habitat preferences and behavioural interactions. However, the relative importance of these factors and whether or not behavioural interactions reinforce or disrupt habitat associations are poorly understood. This paper explores the degree to which habitat and social preferences explain the association that three common coral reef cardinalfish species (Zoramia leptacanthus, Archamia zosterophora and Cheilodipterus quinquelineatus; family Apogonidae) have with coral substrata at Lizard Island, Great Barrier Reef. At diurnal resting sites, species were strongly associated with branching corals, with 80–90% of each species inhabiting one branching coral species, Porites cylindrica. Species were also highly gregarious, forming large con-specific and hetero-specific aggregations in coral heads, potentially reinforcing habitat associations. Three-way choice experiments were conducted to test fishes habitat preferences for living coral over dead substrata, for particular coral species, and the influence of gregarious behaviour on these habitat choices. The strength of habitat preferences differed among species, with Z. leptacanthus preferring live coral and P. cylindrica, A. zosterophora preferring P. cylindrica, whether live or dead and C. quinquelineatus exhibiting no preferences. All species were attracted to conspecifics, and for C. quinquelineatus and A. zosterophora, conspecific attraction resulted in stronger preferences for live corals. Gregarious behaviour also increased C. quinquelineatus associations with P. cylindrica. The relative strength of social attraction versus habitat preferences was investigated by comparing fish habitat preferences in the presence and/or absence of conspecifics. The presence of conspecifics on non-preferred rubble habitat reduced each species association with live coral. This study’s results indicate that in the field, habitat preferences and conspecific attraction combine to reinforce the association between cardinalfishes and a narrow range of coral substrata.     

Leaf age preference for oviposition by three species of whitefly on lemon

The leaf age preferences for oviposition by three species of whitefly (Dialeurodes citri [Ashmead],Aleurothrixus floccosus [Maskell], andParabemisia myricae [Kuwana]) were tested on lemon,Citrus limon (L.). Four categories of leaf age were tested (I: incompletely expanded, growing leaves; II: young leaves that are completely expanded but not yet dark green; III: completely mature leaves [dark green]; IV: completely mature leaves judged to be older than III as determined by the age of the twigs to which they were attached). Paired choice experiments were conducted in the laboratory using excised leaves and in the field using intact leaves. In both laboratory and field tests,P. myricae preferred I over all other leaf ages. It rarely oviposited on II and never oviposited on III or IV, even when they were the only two choices. The order of preference forD. citri in laboratory and field tests was II>I>III>IV. In laboratory tests,A. floccosus exhibited the same order of preferences asD. citri, but in the field, II was preferred over all others, III was slightly preferred over IV, and I varied greatly among replicates in comparisons against III or IV. In related experiments, the choice between I & III byA. floccosus andD. citri was not affected by confining the whiteflies to either I or III for three days prior to testing.

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Sélection pour la ponte des feuilles deCitrus limon en fonction de l'âge par trois espèces d'aleurodes

Résumé Des feuilles de 4 tranches d'âges appartenant àCitrus limon L.: 1) en cours de croissance et incomplètement développées; 2) développées mais pas encore vert sombre; 3) complètement développées (vert sombre) 4) plus âgées que le lot 3 d'après l'âge du rameau, ont été proposées pour pondre à 3 espèces d'aleurodes (Dialeurodes citri Ashmead,Aleurothrixus floccosus Maskell,Parabemisia myricae Kuwana). Les femelles étaient isolées dans des cages dont les ouvertures sur deux côtés opposés étaient obstruées par la face inférieure de feuilles de citronniers d'âges différents. Les feuilles utilisées au laboratoire étaient coupées, les expériences dans la nature étaient faites avec des feuilles intactes. Récoltées au bout de 2 jours, leurs ufs étaient dénombrés.Dans les 2 cas,P. myricae a préféré significativement le lot 1, les pontes sur lot 2 étaient rares, et nulles sur les lots 3 et 4 même en l'absence de choix. Les préférences significativement différentes deD. citri au laboratoire et dans la nature étaient les suivantes: 2>1>3>4.A. floccosus a présenté au laboratoire les mêmes préférences queD. citri (P0,0004), mais certains résultats étaient contradictoires. Au laboratoire et dans la nature, le lot 2 était préféré; de même le lot 3 était préféré au lot 4. Cependant de nombreuses expériences de choix dans la nature entre les lots 1 et 3 et 1 et 4 ont montré la même tendance qu'au laboratoire, mais pour d'autres, les résultats étaient inversés. Après regroupement des résultats dans la nature, le lot 3 a été préféré au lot 1 (résultats à la limite du seuil de validité); aucune différence significative n'est apparue entre les lots 1 et 4. Les choix deA. floccocus etD. citri entre les lots 1 et 3 n'ont pas été modifiés par la présence de l'un des lots pendant les 3 jours précédant l'expérience.

     

The habitat and conduit functions of roads in the spread of three invasive plant species

Nonnative plant species commonly occur along roadsides, and populations are often assumed to invade by spread along the road axis. To distinguish between the function of roadsides as movement corridors and as habitat, nonnative plant species were surveyed along roads in deciduous forest sites in southeastern Ohio, USA. The importance of road proximity was tested by comparing nonnative species abundance in 100 m transects along roads with transects in undisturbed forest. Nonnative species were most abundant and most frequently observed in roadside sites in valleys. Three common species were chosen for closer scrutiny. In a seed sowing experiment roads and open sites proved to be better locations for the germination and growth of Microstegium vimineum than non-roadside and closed-canopy sites. Tussilago farfara and Rosa multiflora occurred in a small number of disjunct patches suggesting infrequent arrival in the sampled transects. Both species were strongly clustered at scales consistent with diffusive spread by vegetative growth and short-range seed dispersal. Comparisons of distributions parallel and perpendicular to roads showed no evidence for enhanced dispersal along the road axis. Microstegium distributions were correlated with local light availability implying site saturation. Microstegium micro-distributions suggested that spread along the road axis was facilitated by movement of dormant seeds in road maintenance. Thus, roadsides appear to function as both habitat and a conduit for population expansion, with the rate of spread dependent on the life history of the individual species. These results suggest a hierarchical process of regional invasion, with different dispersal mechanisms functioning at different spatial scales.     

Spatiotemporal variability of stream habitat and movement of three species of fish

Relationships between environmental variability and movement are poorly understood, due to both their complexity and the limited ecological scope of most movement studies. We studied movements of fantail (Etheostoma flabellare), riverweed (E. podostemone), and Roanoke darters (Percina roanoka) through two stream systems during two summers. We then related movement to variability in measured habitat attributes using logistic regression and exploratory data plots. We indexed habitat conditions at both microhabitat (i.e., patches of uniform depth, velocity, and substrate) and mesohabitat (i.e., riffle and pool channel units) spatial scales, and determined how local habitat conditions were affected by landscape spatial (i.e., longitudinal position, land use) and temporal contexts. Most spatial variability in habitat conditions and fish movement was unexplained by a site’s location on the landscape. Exceptions were microhabitat diversity, which was greater in the less-disturbed watershed, and riffle isolation and predator density in pools, which were greater at more-downstream sites. Habitat conditions and movement also exhibited only minor temporal variability, but the relative influences of habitat attributes on movement were quite variable over time. During the first year, movements of fantail and riverweed darters were triggered predominantly by loss of shallow microhabitats; whereas, during the second year, microhabitat diversity was more strongly related (though in opposite directions) to movement of these two species. Roanoke darters did not move in response to microhabitat-scale variables, presumably because of the species’ preference for deeper microhabitats that changed little over time. Conversely, movement of all species appeared to be constrained by riffle isolation and predator density in pools, two mesohabitat-scale attributes. Relationships between environmental variability and movement depended on both the spatiotemporal scale of consideration and the ecology of the species. Future studies that integrate across scales, taxa, and life-histories are likely to provide greater insight into movement ecology than will traditional, single-season, single-species approaches. The Virginia Cooperative Fish and Wildlife Research Unit is jointly sponsored by the US Geological Survey, Virginia Polytechnic Institute and State University, Virginia Department of Game and Inland Fisheries, and Wildlife Management Institute.     

Experiments on habitat selection in voles: Implications for the inverse distribution of two common European species

Summary Several pairs of species of microtine rodents show inverse distributions or abundance. Here I determine if the inverse abundance relationship of the bank vole Clethrionomys glareolus (a forest species) and the field vole Microtus agrestis (an open field species) could be explained by habitat selection. Habitat selection was examined by choice experiments with combinations of soil and vegetation types, and with separate habitat factors. The effects of early experience and social behaviour were also studied, e.g. in a large outdoor enclosure. M. agrestis showed a stronger dependence on shelter (e.g. easily excavated soil or thick grassy vegetation) than C. glareolus, while the presence of preferred food did not affect habitat selection for either species. In both species, laboratory-bred specimens showed considerably fewer preferences. Habitat selection in subadult C. glareolus was influenced by social clumping, while territorial behaviour made more even distributions in M. agrestis.The less rigid habitat selection and weaker territorial behaviour observed in the forest species (Clethrionomys) compared with a species from temporary habitats (Microtus) is not consistent with present concepts of r- and K-selection.     

Chromosomal differentiation and genome size in three European mountain Lilium species

 Three related and taxonomically close species of the genus Lilium (L. pyrenaicum Gouan, L. pomponium L. and L. carniolicum Bernh.), all of them with 2n=24 chromosomes, have been studied for chromosomal differentiation, using fluorochrome banding and fluorescence in situhybridization (FISH), and for genome size and GC percentage using flow cytometry. The total DNA content of L. pomponium (2C=70.26 pg) was about 5% higher than that of L. pyrenaicum (2C=67.74) and L. carniolicum (2C=67.37 pg), while GC percentage was higher in this last species (36.60%) than in L. pomponium (35.56%) and lower than in L. pyrenaicum (37.92%). Silver staining, fluorochrome banding with chromomycin A₃ (CMA) and fluorescence in situ hybridization (FISH) clearly pointed out the number of nucleoli, the number and position of GC-rich bands and the number and location of rDNA sites thus permitting distinction of the three species at chromosomal level. Two families of ribosomal genes, 18S-5.8S-26S (18S) and 5S rRNA genes, were separated onto different pairs in chromosome complements of examined species. Chromosome regions containing both kinds of rRNA genes were also GC-rich regions. The results revealed a clear interspecific differentiation at the chromosomal level and permitted the discussion about relationships among the species. Received June 21, 2002; accepted October 4, 2002 Published online: Febraury 7, 2003     

Relationships between life-history strategies of European freshwater fish species and their habitat preferences

1. Focusing on the current environmental characteristics, the ‘habitat template’ theory proposes that life‐history strategies summarise how evolution has shaped species to cope with the temporal and spatial variability of their present environment. The hierarchical ‘landscape filters’ concept predicts that the distribution of species reflects their specific traits that allow them to pass through multiple habitat filters. Together, these theories showed the importance of identifying the functional relationships of species to selective habitat forces in order to predict the organisation and response of communities to the environment. 2. We test the relationships between life‐history traits of European freshwater fish species’ and their habitat preferences, to detect the strategies adopted by fish to cope with their current habitat. For this purpose, we use published data on species traits and habitat preferences (microhabitat hydraulics, temperature and oxygen level). We use multivariate analyses to classify fish species’ strategies and test the relationships between strategies and habitat preferences. 3. We identified a continuum of life‐history patterns between two extremes, with opportunistic and periodic species. Our study supports the idea that microhabitat hydraulics plays a more important role as a template for these species ecological strategies than temperature and oxygen level. Indeed, riffle habitats may select opportunistic species whereas weak relationships are found between species strategies and both their temperature and oxygen level preferences. In addition, the ratio between mortality and growth rate (dimensionless trait), reflecting a trade‐off between growth and survival, varied among species according to the use of their hydraulic habitat, with species using deep habitats exhibiting the highest values. 4. These general relationships between hydraulics and traits may be of importance in community ecology to develop predictive models to understand how fish communities change with the hydraulic environment.     

Local and regional influences on patterns of parasite species richness of central European fishes

Local, regional and global influences on the patterns of parasite species richness of 39 freshwater fish species from Central Europe were investigated. Host local abundance and host occurrence were considered respectively as local and regional factors, while host geographical range in longitude and latitude was considered as a global factor. Influences of size, ecology and behavior of hosts were also included in a comparative analysis using the independent contrasts method. We considered host habitat, host diet, host shoaling behavior and mobility. We found a positive relationship between local occurrence of fish and global range of their distribution. We confirmed previous findings showing the importance of host behavior and ecology on the variability of parasite species richness. Second, we showed how a global pattern, such as host geographical range, may affect the variability in parasite species richness through its effects on local abundance and distribution of hosts. A negative relationship between endoparasite species richness and host longitudinal range was found. This suggests that fish with eastern distribution live in the boundary of their distribution in Central Europe far from their center of distribution, which should also be characterized by a higher diversity of parasites.     

Diets and habitat preferences of three species of crocidurine shrews in arid southern Africa

The diets and habitat preferences of three species of crocidurine shrews, Crocidura cyanea, C. fuscomurina , and C. hirta (Insectivora: Soricidae), were studied in arid areas of Namibia in March and April, 1992. Simultaneous sampling of the prey ingested and available along foraging trails showed that, although the shrews are generalist insectivores that take most types of prey from the range available, each species selected Araneida, Chilopoda, Isoptera, and insect larvae but avoided Formicidae. The preferred prey were mostly soft-bodied with a relatively high ratio of body water to energy content, which may assist in maintenance of body water balance. The three species showed inconsistent preferences for rarer categories of prey such as Thysanura and Orthoptera, as well as for Coleoptera. Each species of shrew selected sites providing dense ground level vegetation and deep leaf litter, With the preference for cover being strongest in C. fuscomurina. Dense vegetation probably provides shelter from predators and high daytime temperatures; both C. fuscomurina and C. hirta used denser vegetation by day than by night. Each species foraged in leaf litter, perhaps increasing access to preferred types of prey. Preference for moist soil, especially in C. fuscomurina , probably also reflects increased access to preferred prey, although moist soil may also facilitate burrow construction. Because patterns of habitat and diet selection appear to be linked, future research on resource use in these shrews should be pursued most fruitfully in field experiments or captive laboratory trials.     

Brightness preference in nocturnal and diurnal South American monkeys.

In a test situation where choice was not tied to reinforcement, both squirrel monkeys (Saimiri sciureus) and owl monkeys (Aotus trivirgatus) consistently chose to view the more luminant of a pair of lights. Over a two log unit range of luminance, the degree of preference in both species for the brighter of two lights was found to depend on their difference in luminance and not on their actual luminances.     

Biomass allocation in three macrophyte species in relation to the disturbance level of their habitat

1. The hypothesis was tested that perennial plants surviving in habitats frequently disturbed by floods should demonstrate the ability to escape the frequently scoured surficial zone by using refugia located deeper in the substrate. As a consequence, they should allocate more biomass to their underground parts than when growing in rarely disturbed habitats.
2. The allocation of biomass of three aquatic macrophytes ( Berula erecta , Groenlandia densa , Myriophyllum spicatum ) to their different organs (underground and above-ground parts) was measured in former river channels organized along a gradient of scouring flood disturbances.
3. For each species, biomass allocation to underground parts varied between sites: from 10.2 to 37.5% in B. erecta , from 14.9 to 31.1% in G. densa and from 8.4 to 23.2% in M. spicatum . It was correlated to the flood disturbance level of the site for B. erecta and M. spicatum but not for G. densa. No relationship was found between biomass allocation to underground parts and sediment richness.
4. The plasticity in biomass allocation of these three macrophyte species appears to be an adaptation to the variability in environmental conditions and implies difficulties in the classification of plant primary strategies, when this type of plasticity is ignored.