14. Lake Sedmo Rilsko (Bulgaria): Lateglacial vegetation history

The length of the pollen deposition period covered by an individual moss sample is a matter of discussion. Here we evaluate (1) how many years of pollen deposition are contained in individual moss samples, (2) which factors (i.e. moss type, geographical location and thickness of sample) affect the pollen content, and (3) whether the time span of the pollen deposited in the moss is related to the pollen type. The pollen deposition period covered by 20 individual moss samples adjacent to pollen traps was estimated using the accumulation method. The investigated moss samples usually contain pollen from a period of two to five years. The length of the time period covered by the moss depends on the thickness of the moss cushion and, therefore, we recommend estimating the time period over which the moss has been growing by the increment method. Compared to pollen traps, mosses tend to accumulate more Poaceae and bisaccate grains (especially Picea) but less Cyperaceae pollen. Betula and pollen tetrads tend to be equally represented in mosses and traps. This study demonstrates that moss samples may contain sufficient pollen to be a reliable modern analogue for Quaternary peat and sediment samples but cannot be used to calculate pollen accumulation rates.     

Comparisons of moss polster and pollen trap data: a pilot study

Modern pollen-vegetation relationships have been explored in recent years through analysis of either pollen trap (= absolute results) or moss polster contents (= relative results). In the past few attempts were made to compare data from these two sources. It was the aim of the present study to achieve this comparison through more exact sampling of moss polsters. A pilot study was carried at Morton Lochs, Fife, Scotland where pollen traps have been sampled continuously for ten years. Moss samples were collected from cushions adjacent to two of the pollen traps during May 1985, 1986 and 1987. Moss polsters were sampled usingan equivalent surface cross-section area to that of the traps. In comparing the pollen results from the two sources some similarities can be identified, but in general there are more inconsistencies. These may be traced to difficulties in the precision of moss sampling, variable rates of moss growth and local deposition factors. It is considered that further study, in which these difficulties are minimised, would be justified. Ultimately it is hoped that precise sampling of mosses will produce sufficiently reliable modern pollen rain results to enable pollen-vegetation studies to move away from relative (percentage) values without requiring long-term artificial pollen trapping techniques.     

Numerical analysis of surface and fossil pollen spectra from northern Fennoscandia

Aim Determination of the main directions of variance in an extensive data base of annual pollen deposition, and the relationship between pollen data from modified Tauber traps and palaeoecological data. Location Northern Finland and Norway. Methods Pollen analysis of annual samples from pollen traps and contiguous high‐resolution samples from a peat sequence. Numerical analysis (principal components analysis) of the resulting data. Results The main direction of variation in the trap data is due to the vegetation region in which each trap is located. A secondary direction of variation is due to the annual variability of pollen production of some of the tree taxa, especially Betula and Pinus. This annual variability is more conspicuous in ‘absolute’ data than it is in percentage data which, at this annual resolution, becomes more random. There are systematic differences, with respect to peat‐forming taxa, between pollen data from traps and pollen data from a peat profile collected over the same period of time. Main conclusions Annual variability in pollen production is rarely visible in fossil pollen samples because these cannot be sampled at precisely a 12‐month resolution. At near‐annual resolution sampling, it results in erratic percentage values which do not reflect changes in vegetation. Profiles sampled at near annual resolution are better analysed in terms of pollen accumulation rates with the realization that even these do not record changes in plant abundance but changes in pollen abundance. However, at the coarser temporal resolution common in most fossil samples it does not mask the origin of the pollen in terms of its vegetation region. Climate change may not be recognizable from pollen assemblages until the change has persisted in the same direction sufficiently long enough to alter the flowering (pollen production) pattern of the dominant trees.     

Do moss samples,pollen traps and modern lake sediments all collect pollen in the same way? A comparison from the forest limit area of northernmost Europe

Moss polsters, pollen traps and lake surface sediment samples are commonly used as climate calibration data or as modern analogues for reconstructing vegetation from fossil profiles, but the differences in pollen content between these media have received little attention. This study aims to analyse how the three media differ in reflecting individual vegetation types and spatial differences in vegetation. 119 modern samples (64 moss polsters, 37 lake surface sediment samples and 18 pollen traps from which a collection was made annually) were taken from northern Fennoscandia and the Kola Peninsula as a broad transect crossing the northernmost forest limits of Betula pubescens ssp. czerepanovii (mountain birch), Pinus sylvestris (Scots pine) and Picea abies (Norway spruce). The pollen assemblages from these samples were compared with the surrounding vegetation visually and via PCA (principle components analysis) and cluster analysis. Both comparisons allow a correct distinction between pollen assemblages of arctic/alpine heath, mountain birch dominated areas, and boreal coniferous forests. The differences between the vegetation zones are stronger than the differences between the sampling media. Nevertheless, lake sediment samples from the mountain birch woodland zone tend to overestimate pine and underestimate birch. Pollen traps are biased towards lower tree pollen percentages and higher values of shrubs, herbs and Cyperaceae. This bias is especially strong in traps that have missing years in the data. Irrespective of the vegetation zone, pollen traps tend to have lower Pinus pollen percentages than in the adjacent moss polsters.     

Comparative study of different methods for capturing airborne pollen,and effects of vegetation and meteorological variables

The atmosphere of Mar del Plata city (Argentina) was monitored using three different pollen traps—Burkard, Rotorod, and Tauber—during a flowering season (mid-winter to springtime) in order to compare the diversity and amounts of pollen grains collected in each trap. Sixty three percent of taxa diversity was simultaneously recorded by all three apparatuses. The five most abundant types of pollen in the three traps were Platanus, Ulmus, Quercus, Poaceae, and Fraxinus. The Tauber trap contained the most pollen from taxa with the largest grains, and exclusively trapped ten pollen types with entomophilous characteristics. The amount of pollen in each trap and local/extra-local plant abundance were compared to elucidate the effect of the method used on the pollen–vegetation relationship. The correlation coefficients for total pollen were positive and significant between all pairs of traps. The maximum peak was observed first in the Burkard trap, in agreement with an increase in temperature, but the general pattern in the three traps was similar, with a maximum in early spring. Opposite deviations from the mean value were observed for the Burkard and Tauber data; these deviations were associated negatively with wind speed and relative humidity, respectively. This paper discusses whether in some cases the pollen records could be used in an equivalent manner, although with some limitations.     

Pollen and spores transport into the Antarctic

Summary Pollen deposition in a 7 cm thick moss turf (Polytrichum juniperinum) from a site in Admiralty Bay (King George Island, maritime Antarctic zone) and in three cushions of Schistidium antarcticum (1–3 cm thick) from Bailey and Clark Peninsula (Wilkes Land, Greater Antarctica) were investigated. Twenty-four pollen and spore taxa were identified from a total of 1094 grains in the sample from Admiralty Bay. By contrast, the 3 samples from Wilkes Land contained only 10 grains (5 taxa). While the former sample was poor in lichen-mycobiont spores the latter revealed very many. Pollen deposition appears to be a good indicator of direction and frequency of immigration of cryptogamic propagules. It seems that the presence of cryptogamic species in the maritime Antarctic zone is predominantly the result of long-distance transport from neighbouring South America and a selection by climatic factors. The low frequency of pollen in Wilkes Land indicates a separation of this part of Antarctica from propagule exchange with other continents and the cryptogams of most of the Antarctic continent may represent a largely relictual flora which survived in ice-free refugia.Dedicated to Professor Dr. W. Halbsguth on the occasion of his 75th birthday     

Pollen assemblages of tauber traps and surface soil samples in steppe areas of China and their relationships with vegetation and climate

We use 86 pollen trap and surface soil pollen samples in steppe areas of China to explore the relationships between modern pollen, vegetation, and climate. The modern pollen spectra from both sources have comparable compositions with regard to the major pollen taxa. However, the number of taxa in the traps was higher than in the surface soil samples. Both pollen accumulation rates and pollen concentrations are higher in the typical steppe areas than in the desert steppe areas. Discriminant analysis indicates that pollen spectra from trap and surface soil samples roughly reflect the vegetation zones of desert steppe and typical steppe, especially in the case of the trap samples. Detrended canonical correspondence analysis suggests that pollen assemblages have a significant relationship with the temperature of the coldest month and the mean annual precipitation.     

After 8 years of annual pollen trapping across the tree line in western Norway: are the data still anomalous?

The modern pollen production of local sub-alpine and alpine vegetation has been monitored over 8 years by using pollen traps situated along an elevational transect crossing the Pinus sylvestris and Betula pubescens elevational limits and continuing into the mid- and high-alpine vegetation in western Norway. The aim of the study is to monitor annual variation in the pollen production and hence the representation of common taxa found at or near the tree-line. The results can be used to derive critical pollen values that can be used to infer the presence of these taxa in the past, and to reconstruct past changes in tree line positions. Nine modified Tauber pollen traps were critically positioned in the vegetation from 663 to 1,347 m a.s.l. Three sub-aquatic traps were located in a small lake at 800 m a.s.l. just above the present-day B. pubescens tree-line. The traps have been sampled from 2004 to 2012. The results display large variations in pollen percentages and pollen accumulation rates from year to year, as well as great differences between the traps placed in the vegetation and in the lake, suggesting that further pollen trapping is needed to get a solid long-term average. The vegetation traps follow the vegetation distribution better and, as expected, give a more local signal than the lake traps.     

A comparison between modern pollen spectra of moss cushions and Cundill pollen traps

Pollen spectra of 23 Cundill pollen traps from 23 different sampling sites in Southwest Turkey are compared with the corresponding pollen spectra of moss cushions from the same sites. The Cundill pollen traps represent the modern pollen rain data from one year whereas the moss cushions represent the pollen rain of several years. The comparative study reveals some main differences between the two pollen entrapment media. The one-year entrapment medium (pollen trap) appears to be more sensitive for local (releve area 10x10 m) and regional (100-500 m, or a few kilometres for Olea europaea) vegetation. Pollen spectra of moss cushions are dominated by high pine pollen percentage values and hardly sense fine vegetation structures. The conclusion of this comparative modern pollen study allows to interpret fossil sediment spectra from the Near East in a more critical way. It is concluded that one should preferably sample rapidly deposited sediments for palynological analyses, as the resulting highresolution pollen diagrams will be most informative about the former vegetation patterns.     

Variation in annual pollen accumulation rates of Fagus along a N–S transect in Europe based on pollen traps

Annual pollen-accumulation rates (PAR) of Fagus (beech) obtained within the framework of the Pollen Monitoring Programme (PMP) were analyzed in pollen traps along a N–S transect from the Baltic Sea to the Black Sea in different European vegetation units. The study regions are situated in the lowlands of northern Poland, the uplands of SE Poland, the Czech Krkono?e Mts, the Czech ?umava Mts, the Swiss Jura Mts, the Swiss Alps, the Bulgarian Rila Mts and the Bulgarian Strandzha Mts. Most time series are 10 or 11 years long, some are 5–16 years long. Inter-annual fluctuations in Fagus PAR were analyzed and compared with seed mast years. Years with high Fagus PAR and others with low Fagus PAR occurred most frequently in parallel within each region and often in two neighbouring regions. 2006 was exceptional as it had a very high Fagus sylvatica pollen deposition in all study regions and it was also a mast year. In Bulgaria, the trend in the 5 years of Fagus orientalis PAR in the Strandzha Mts differed from that of F. sylvatica PAR in the Rila Mts. Aiming at establishing the relationship between average Fagus PAR and tree cover, differences in Fagus PAR (averaged per pollen trap) were related in each region to the proportion of beech trees in the vegetation within 2 km of the pollen traps, the distance to the nearest pollinating Fagus tree, regional or local presence of beech forests, the degree of landscape openness, and the size of forest opening in which a trap is situated. Average Fagus PAR was found to track the regional abundance of beech trees in the vegetation, not the distance of the nearest Fagus tree. Regional occurrence of beech-dominated forests was reflected by a Fagus PAR of ca. 1,400 grains cm^?2 year^?1, local abundance very close to pollen traps by ca. 2,400 grains, small patches of forest with admixture of Fagus by ca. 170–220 grains, and scarcity or absence of Fagus by ca. 40 grains or less.     

Effect of hunger on yellow water trap catches of hoverfly (Diptera: Syrphidae) adults

1 An experiment was conducted in a winter wheat field using yellow water traps at crop height and at ground level, near to and distant from flowers, to test the hypothesis that such traps are seen as a source of food by flower‐feeding adult hoverflies and are therefore likely to selectively trap hungry individuals. Hoverflies caught in each trap were counted and identified and the amount of pollen in their guts was assessed. Ratios of numbers of hoverflies seen in the wheat crop to numbers caught in nearby traps were compared for the different treatments. 2 Most hoverflies were caught in crop‐high traps but they included a high proportion of individuals with empty guts. The taxa were: Episyrphus balteatus (De Geer), Metasyrphus corollae (F.) (these species together accounted for over 90% of the individuals trapped), with Sphaerophoria spp., Syrphus spp., Scaeva pyrastri (L.), Melanostomini and unidentified others making up the rest. 3 Significantly fewer hoverflies were captured in low than in high traps. The ratios of numbers trapped to numbers observed, in flower and in no flower treatments would be expected to be the same if the traps were not selective. This was supported for low traps. With high traps, however, there was a highly significant difference between the ratios (71.34 and 126 : 8, respectively). 4 Flies captured in high traps had less pollen in their gut than those captured in low ones. At each distance, more E. balteatus captured in high traps were in pollen category 1 (< 20 grains) than in any other category. The opposite state was seen in low trap catches, where most flies were in category 5 (> 5000 grains). Median pollen categories were 2 (21–200 grains) and 4 (1501–5000 grains) for flies caught in high and low traps, respectively. 5 The ecological selectivity of traps according to their height and the physiological condition of the targeted individuals is a problem likely to affect many trapping systems apart from the one described in this paper.     

Pollen dispersal and representation in a neotropical rain forest

Results of 3 years of pollen trapping on Barro Colorado Island, Panama, are presented. Minimum pollen dispersal distances are estimated for the most abundant pollen taxa. Dispersal distances for some taxa appear to be as low as 5 m, while for other taxa at least 50% of the trapped pollen had travelled more than 40 m. No consistent pattern of spatial variation was reflected in the pollen rain, however, samples close to large canopy gaps had exceptional abundances of 'disturbance' taxa. A preliminary analysis of the representation of canopy components indicates that as much as 19% of pollen caught in the traps was derived from large tree species.     

Pollen input to, and incorporation in, two crater lakes in tropical northeast Australia

Pollen input to the water surfaces of maar crater lakes Barrine and Eacham (1km(2) and 0.5km(2) respectively, ca. 720m a.s.l.), surrounded by rainforest in northeastern Australia, were measured in floating Tauber traps for a variety of periods spanning the years 1978 to 1985. The mean weekly total pollen catch is 75% of that estimated for the surrounding region of varied vegetation. Regardless of their sources, the influxes of pollen taxa are unevenly distributed over the lakes' surfaces. There is similar variation through time, with some indication that this might be smoothed over periods of 5years or more. There are no inflow streams and pollen input down surrounding slopes is negligible. Mean total pollen influx to seston traps close to the lakes' bottoms (1979 to 1988) is <5% of that to the Tauber traps, a discrepancy attributed to trap design. In contrast, mean annual influx measured by the Tauber traps is similar to that estimated from dated sediment samples younger than 1966 AD at Lake Barrine. The proportions of different pollen taxa are less affected by trap position or period of exposure than are their influxes; their mean percentages for Tauber traps, seston traps and sediment samples are all rather similar.The similarity between the mean contribution of exclusively rainforest pollen to the lakes' surfaces (56%) and to Oldfield traps in rainforest interiors (51%) emphasizes the importance of the rainforests, which clothe the crater walls, as sources of pollen to the lakes. The impact of pollen from other vegetation in the region, although present, is much smaller than at interfaces between rainforest and other vegetation types. All this pollen is mixed within and above the crater and delivered to the lakes' surfaces by gravity and frequent rainfall. Limnological processes redistribute it in the water body before it is incorporated into the deep-lake sediments.The value and limitations of current pollen transport and accumulation theory are noted in relation to sites with morphologies similar to these two crater lakes and to the irregular flowering and floristic inhomogeneity of tropical rainforest. The potential for the use of modern pollen input data in the interpretation of pollen analyses from the sediments of such sites is explored.     

Comparison of pollen sampling with a Burkard Spore Trap and a Tauber Trap in a warm temperate climate

Aerobiological data have been widely used by many scientists, including those that study modern flora as well as those wishing to reconstruct past vegetational associations. Burkard (Hirst-type) volumetric spore traps are widely used instruments for studying airborne pollen, while Tauber traps are typically used to analyze pollen deposition. The present study compared the pollen collected by these two methods in Tulsa, Oklahoma a warm temperate area with year-round pollen. There was a strong correlation between the pollen influx from the Tauber traps and cumulative sum of average daily airborne pollen concentrations recorded with the Burkard spore trap over the course of 12 months from 1 Feb 1997 through 1 Feb 1998. The correlation coefficient between all taxa over the 12 months was 0.914; while the correlation coefficient for the monthly totals was 0.972. The data showed that both methods reflected local anemophilous vegetation although variations occurred in the prevalence recorded by both samplers.     

Monitoring of pollen rain in moss polsters and mats in view of their suitability as controls for determining the air-borne component of pollen spectra in herbivore faeces

Pollen has been extracted from six bryophyte and two lichen taxa, sampled from three habitats: upland heath, lowland bog and mixed woodland, in both west and east Scotland. The highest number of pollen taxa was recorded in bothSphagnum andThuidium; the lowest numbers of pollen taxa were recorded in two of the mosses and both lichens. For pollen rain control purposes in herbivore faecal studies, woodland species of bryophytes are not suitable due to the sheltering effects from rain provided by the overhead canopy. Open habitats such as moorland heath and bog provide polsters/mats which normally trap pollen rain more effectively for control purposes. The aim of this study was twofold: to observe the effectiveness of selected polsters and lichens as traps for pollen rain and to establish eventually, whether or not such vegetative traps could be used as controls in pollen spectra analyses of herbivore faeces for the determination of nutritional regime and habitat. From a comparison with recent studies in north west Spain of three taxa in common with Scottish samples,Polytrichum demonstrated a consistently higher number of pollen taxa trapped.Thuidium andCladonia all yielded between one and a half and three times the number of pollen taxa as in the same three bryophyte/lichen taxa sampled in Scotland. Investigation relating the pollen rain trapped in polsters to that in faecal material are continuing, and will be reported in due course. This paper is based on one given by E.C. at the 5th International Aerobiology Conference, held at Bangalore, India, August 1994.     

Pollen analysis of coal-bearing Miocene sedimentary rocks from the Seyitömer Basin (Kütahya), Western Anatolia

The late Early-Middle Miocene sequences of the Seyitömer Basin (western Anatolia) were palynologically investigated. Fifty-five taxa belonging to seven gymnospermous and 48 angiospermous pollen genera were identified in the 19 productive samples. Two pollen zones were recognised based on the changing abundance of individual tree taxa. Zone 1 is characterized by predominance of Pinus and Cedrus. Zone 2 is characterized by predominance of deciduous Quercus and evergreen Quercus and a marked reduction in representation of Taxodiaceae. The differences in the pollen spectra between Zone 1 and Zone 2 may reflect the global Middle Miocene cooling. These results are largely comparable to pollen data derived from the neighbouring areas. The vegetation of the Seyitömer Basin was dominated by trees. This palynological analysis reveals the existence of a swamp-forest developed in a subtropical to warm-temperate humid climate.     

Evaluation of indoor passive pollen sedimentation over 1 year: a possible source of contamination?

The study aimed to evaluate the possible contamination by passive indoor pollen sedimentation in a laboratory where routine pollen analyses were conducted, but where no particular laboratory policies were adopted to limit contamination. Gravimetric pollen deposition was observed on traps (petri dishes soaked with glycerol) set in the palynology laboratory, under an extractor hood and on the bench beside it over 1 year (1995–1996), and in an air filtered room in a flow cabinet and on its roof over 1 month for comparison. Under the extractor hood, three types of airborne particles were deposited: pollen grains, spores and algae, representing 32.35, 67.28 and 0.37%, respectively, of the total sedimented particles over 1 year. The number of pollen grains deposited on the surface trap ranged from 0 (27 November to 4 December 1995) to 707 (10–18 April 1995). The highest number of taxa during a weekly collection was 23 (9–15 May 1995). The pollen flora represented by anemophilous pollen (>90% of the trapped pollen) was related to the vegetation next to the laboratory:Acer, Carpinus, Castanea, Corylus, Cupressaceae, Pinus, Quercus, Salix, Taxus for trees and shrubs andArtemisia, Brassicaceae, Plantago, Poaceae, andUrticaceae for grasses and weeds. Indoor pollen deposition corresponded to the period of the outdoor pollination (macroscopic field observation) which lasted from March to the beginning of August. However, some pollen were almost always present in the collection (Poaceae, Salix, Castanea, Betula), reflecting the occurrence of pollen grains in the atmosphere out of the pollination period. Moreover, about five times more entomophilous pollen was found under the extractor hood compared to the other area of the laboratory; even in the flow cabinet of the air filtered room, 237 particles were captured (versus only 15 on the roof). These data suggest a possible human contamination during operations under the extractor hood or in the flow cabinet. Although few airborne pollen were found, possible contamination has to be considered in investigations where even low pollen quantities are of interest.     

Airborne Pollen and Fungal Spores in Florist Shops in Worcester and in Bristol, UK: A Potential Problem for Occupational Health

Persistent allergies are common in workers in florist shops but little research has been done on the reasons for this. This paper reports an investigation of occupational exposure of florists to pollen and spores in three florist shops over a 2-week period in the autumn of 2000. In each shop three sampling methods were used: Burkard continuous volumetric samplers, deposition plates and low-tac tape for surface samples, including hands of the florists and leaves of a selection of the plants. The florists kept a record of the type of work they undertook each day and of the stock amounts of flowers in the shops. The volumetric traps collected 80 pollen and spore taxa. The average concentrations recorded through the working day were generally low but short-term peak (one hour mean) concentrations of some types were found to be relatively high or very high. For a few taxa these concentrations equalled or exceeded those typical for short-term peaks in wind dispersed pollen types in the ambient air. Fungal spore concentrations of several known allergenic types were also very high for peak periods. This was most notable for Aspergillus spp. which reached extremely high concentrations in one shop, compared with typical peak concentrations in the ambient atmosphere. Low-tac tape leaf samples demonstrated that the flowers' foliage is a major source of the fungal spores. Few pollen grains or fungal spores were found on the hands.     

阴山山脉东段花粉通量及其与表土花粉比较研究

阴山山脉东段蛮汉山和大青山山顶草甸、桦木林、虎榛子灌丛、人工油松林及杂草草原5个植被类型花粉通量与表土花粉对比研究发现,不同植被类型及同一植被类型不同样点花粉通量和花粉浓度差异明显,捕捉器样品花粉源区面积小于表土样品,能较好地反映样品点周围植被组成,但受虫媒植物和局地植物花粉影响,与表土样品花粉组合差异明显;表土样品中松属、桦属、蒿属及藜科花粉含量多高于捕捉器样品,表明这些花粉不仅易于传播,而且在表土中具有较强的保存能力;禾本科花粉具低代表性,花粉保存能力较低.地层花粉分析中应引起注意.     

Capture of algae promotes growth and propagation in aquatic Utricularia

Background and Aims Some carnivorous plants trap not only small animals but also algae and pollen grains. However, it remains unclear if these trapped particles are useless bycatch or whether they provide nutrients for the plant. The present study examines this question in Utricularia, which forms the largest and most widely spread genus of carnivorous plants, and which captures prey by means of sophisticated suction traps.Methods Utricularia plants of three different species (U. australis, U. vulgaris and U. minor) were collected in eight different water bodies including peat bogs, lakes and artificial ponds in three regions of Austria. The prey spectrum of each population was analysed qualitatively and quantitatively, and correlated with data on growth and propagation, C/N ratio and δ¹⁵N.Key Results More than 50 % of the prey of the Utricularia populations investigated consisted of algae and pollen, and U. vulgaris in particular was found to capture large amounts of gymnosperm pollen. The capture of algae and pollen grains was strongly correlated with most growth parameters, including weight, length, budding and elongation of internodes. The C/N ratio, however, was less well correlated. Other prey, such as moss leaflets, fungal hyphae and mineral particles, were negatively correlated with most growth parameters. δ¹⁵N was positively correlated with prey capture, but in situations where algae were the main prey objects it was found that the standard formula for calculation of prey-derived N was no longer applicable.Conclusions The mass capture of immotile particles confirms the ecological importance of autonomous firing of the traps. Although the C/N ratio was little influenced by algae, they clearly provide other nutrients, possibly including phosphorus and trace elements. By contrast, mosses, fungi and mineral particles appear to be useless bycatch. Correlations with chemical parameters indicate that Utricularia benefits from nutrient-rich waters by uptake of inorganic nutrients from the water, by the production of more traps per unit of shoot length, and by the capture of more prey particles per trap, as nutrient-rich waters harbour more prey organisms.