Modern pollen–vegetation relationships along transects on the Whangapoua Estuary, Great Barrier Island, northern New Zealand

Aims To quantify pollen–vegetation relationships from saline to freshwater in an estuarine gradient from surface samples of the modern pollen rain, to allow more accurate interpretations of the stratigraphic palynological record. Location Whangapoua Estuary, Great Barrier Island, northern New Zealand. Methods Six transects were laid out along a vegetation sequence running from estuarine mud to freshwater swamp. Along these transect lines, 108 plots were sampled for vegetation and surface sediments from wet sand, mud, plant litter or moss (sand and mud sites are inundated by most tides, other sites less frequently). All sediment samples were analysed for pollen. The relationships between plant species frequency and pollen representation were examined at a community scale using twinspan and ordination analyses, and for individual species using fidelity and dispersibility indices, regression and box‐plot analyses. Results The quantitative relationships between source taxon vegetation frequency and its pollen representation varied between species due to differential pollen production and dispersal. twinspan of the surface pollen samples suggests five vegetation types: (A) mangrove (Avicennia marina); (C) Leptocarpus similis salt meadow; (D) Baumea sedges; (E) Leptospermum shrubland; and (F) Typha/Cordyline swamp forest. The (B) Juncus kraussii community is not represented palynologically owing to the destruction of its delicate pollen grains during acetolysis of samples. Detrended correspondence analysis places these communities on an estuarine‐to‐freshwater gradient. However, pollen assemblages at the seaward end of the salinity gradient are less clearly representative of the associated vegetation than those at the landward end, probably because the open vegetation at the former allows the influx of wind‐ and water‐dispersed pollen from surrounding vegetation. Main conclusions The vegetation pattern (zonation) at Whangapoua is reflected in the pollen rain. When the long‐distance and over‐represented pollen types are excluded, five out of six of the broad vegetation communities can be identified by their pollen spectra. Species with high fidelity and low‐to‐moderate dispersibility indices can be used to identify the vegetation types in the sedimentary sequences. The more open vegetation types at the ‘marine end’ of the sequence tend to be ‘overwhelmed’ by regional pollen, but the nature of the sediments and the presence of discriminatory species (e.g. A. marina, Plagianthus divaricatus, Cordyline australis), even in small amounts, will allow correct identification of the local vegetation represented in sedimentary palynological sequences. A box‐plot analysis indicates that the pollen and spore types A. marina (mangroves), Sarcocornia quinqueflora (salt meadow), P. divaricatus (sedges), Gleichenia (shrubland) and C. australis (swamp forest) are highly discriminatory in relation to vegetation type. These discriminatory palynomorphs help with the interpretation of stratigraphic pollen studies. However, salt marsh vegetation communities in the sediments must be interpreted with caution as the marine sediments are easily affected by erosion, bioturbation and tidal inundation effects.     

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.     

Changes in Rotifer Species Composition and Abundance along a Trophic Gradient in Loch Lomond,Scotland, UK

Loch Lomond is the second largest body of freshwater in Great Britain. It is a long, narrow lake (36.4 km long, 8.8 km wide). The northern basin is fjord-like and surrounded by a mountainous, base-poor, rocky catchment. In contrast, the southern basin is much broader and shallower with a mainly lowland, calcareous, agricultural catchment. This causes a trophic gradient along the length of the loch that runs from the oligotrophic northern basin to the more mesotrophic southern basin. Rotifer samples were collected at monthly intervals between May and October 2002 at three locations along the length of the loch. More than 12 species were found, the commonest of which were Keratella cochlearis (Gosse) and Trichocerca stylata (Gosse). Although the species composition of the rotifer community varied very little among the sites, rotifer abundance increased markedly from north to south, apparently reflecting the trophic gradient along the length of the loch. The results suggest that rotifer abundance may be a more sensitive indicator of trophic state, and changes in trophic state, than species composition.     

Spatial variation of modern pollen in Oregon and southern Washington, USA

Surface sediments from 95 lakes provide information on the spatial variation of modern pollen spectra in Oregon and southern Washington. Percentages for 13 pollen types were compared within and between vegetation zones to characterize regional patterns of pollen spectra. The percentage data were also compared with climate variables to determine relationships between pollen percentages and regional climate gradients. The composition of modern pollen spectra corresponds well with the distribution of the pollen producers. Most pollen assemblages were generally dominated by Pinus, but those west of the Cascade Range were dominated by Alnus. Low percentages of Pseudotsuga/Larix, Tsuga mertensiana, Abies, and Picea pollen coincided with local occurrence of the trees. The distributions of the pollen data were arranged along gradients of temperature and effective moisture. West of the Cascade Range, Alnus, Tsuga heterophylla, Pseudotsuga/Larix, and Cupressaceae pollen were abundant and correlate well with moderate temperature and high effective moisture. In the shrub-steppe and woodlands east of the Cascade Range, where effective moisture is low, Artemisia, Cupressaceae, and Pinus pollen were dominant. At high elevations, Pinus, T. mertensiana, Abies, and Picea were common pollen types in areas with short growing seasons and high effective moisture. Pollen percentages collected from lake surface sediments, moss polsters, and soils were compared within a number of vegetation types to assess their similarity. The three types of sample yielded similar results for forested areas, but lake sediment samples from upper- and lower-treeline sites captured a more regional picture of the vegetation.     

Freshwater and wetland plant communities of Loch Lomond

Information on the aquatic and wetland vegetation of Loch Lomond, Scotland, is summarised for the period 1957–90. Aquatic macrophyte growth is estimated to occupy about 1% of the loch's total surface area, being limited to the 0–10 m euphotic zone, and probably excluded from much of this area by wave disturbance and unsuitable substrates. Aquatic vegetation is however abundant in sheltered bays and less-exposed shorelines, particularly in the South Basin of the loch. Although Littorella uniflora (L.) Aschers is the commonest, ubiquitous, submerged plant of the loch, three separate euhydrophyte communities have been identified within the loch. One of these is characterised by abundant stands of Elodea canadensis Michx., which appears to have invaded Loch Lomond some time between 1967–88, and has rapidly spread throughout the loch.     

中国现代花粉数据集

孢粉是重建古植被、古气候的重要基础数据。孢粉数据库对研究样点至区域和全球尺度上的古环境演变规律、古气候变化特征反演和古生物地球化学循环模拟等具有重要意义。该文收集整理了中国1960-2020年间发表和部分未发表的现代花粉数据记录, 包括样品编号、采样位置、采样地经纬度和海拔高度、样品类型、数据来源、数据类型、周边植被信息、参考文献、花粉类群及其含量等信息; 并对数据进行筛选和标准化等处理, 由此整合为中国现代花粉数据集。该数据集由4 497个现代花粉采样点的数据信息组成, 包括660个来自中国第四纪孢粉数据库数据, 1 763个前期整理发表的数据和2 074个近期收集的数据, 涵盖772个花粉类群。样品类型以土壤表层样品(3 332个)为主, 苔藓样品以及湖泊、海洋表层样品等为辅, 广泛分布于全国不同地理区域和植被类型中, 其中以温带荒漠区域(24.91%)和亚热带常绿阔叶林区域(24.02%)最丰富, 其次为温带草原区域(16.14%)和青藏高原高寒植被区域(15.83%)。数据按照来源可分为原始数据(58%)和数值化数据(42%); 按照数据类型可分为原始统计粒数的样点(59%)和以花粉百分比表达的样点(41%)。半个多世纪以来, 科研人员开展了大量的表层现代花粉取样和研究。本数据集虽然仅获取部分记录, 但样点覆盖了我国绝大多数地区, 可有效地用于古植被与古气候重建的现代孢粉与现代植被校验, 并将为中国孢粉数据库的建立与更深入的孢粉研究提供数据支撑。     

Tree line identification from pollen data: beyond the limit?

Aim The boreal tree line is a prominent biogeographic feature, the position of which reflects climatic conditions. Pollen is the key sensor used to reconstruct past tree line patterns. Our aims in this study were to investigate pollen–vegetation relationships at the boreal tree line and to assess the success of a modified version of the biomization method that incorporates pollen productivity and dispersal in distinguishing the tree line. Location Northern Canada (307 sites) and Alaska (316 sites). Methods The REVEALS method for estimating regional vegetation composition from pollen data was simplified to provide correction factors to account for differential production and dispersal of pollen among taxa. The REVEALS‐based correction factors were used to adapt the biomization method and applied as a set of experiments to pollen data from lake sediments and moss polsters from the boreal tree line. Proportions of forest and tundra predicted from modern pollen samples along two longitudinal transects were compared with those derived from a vegetation map by: (1) a tally of ‘correct’ versus ‘incorrect’ assignments using vegetation in the relevant map pixels, and (2) a comparison of the shape and position of north–south forest‐cover curves generated from all transect pixels and from pollen data. Possible causes of bias in the misclassifications were assessed. Results Correcting for pollen productivity alone gave fewest misclassifications and the closest estimate of the modern mapped tree line position (Canada, + 300 km; Alaska, + 10 km). In Canada success rates were c. 40–70% and all experiments over‐predicted forest cover. Most corrections improved results over uncorrected biomization; using only lakes improved success rates to c. 80%. In Alaska success rates were 70–80% and classification errors were more evenly distributed; there was little improvement over uncorrected biomization. Main conclusions Corrected biomization should improve broad‐scale reconstructions of spatial patterns in forest/non‐forest vegetation mosaics and across climate‐sensitive ecotones. The Canadian example shows this is particularly the case in regions affected by taxa with extremely high pollen productivity (such as Pinus). Improved representation of actual vegetation distribution is most likely if pollen data from lake sediments are used because the REVEALS algorithm is based on the pollen dynamics of lake‐based systems.     

A taphonomic study of modern pollen assemblages from dung and surface sediments in arid environments of Spain

Coastal plant communities in arid southeastern Spain are characterized by insect-pollinated scrub species, which fail to occur in Quaternary pollen sequences from valleys, marshlands and marine cores. We investigate pollen–vegetation relationships in samples from soil surfaces, animal dung, and sediments in depressions or basins that, in theory, should have pollen spectra that are comparable to those from sedimentary basins elsewhere. Pollen spectra from basins or depressions are very susceptible to long-distance wind and water transport. This can mask representation of pollen from the surrounding insect-pollinated vegetation, as can over-representation of basin-margin halophilous and hydrophilous pollen. Pollen spectra from biogenic materials of animal origin are the best analogues of local and regional vegetation, and show the best analytical potential in terms of pollen concentration and taxon diversity. Pollination properties of the species studied indicate they will rarely be found in most conventional pollen records. It cannot be stressed too strongly that insight into Quaternary vegetation of arid regions demands complementary pollen analysis of coprolites, urine-cemented deposits, and cave sediments with preserved biotic remains, in addition to water-lain sediments.     

Modern pollen representation of source vegetation in the Qaidam Basin and surrounding mountains, north-eastern Tibetan Plateau

We use a data set of 35 surface pollen samples from lake sediments, moss polsters and top soils on the north-eastern Tibetan Plateau to explore the relationship between modern pollen assemblages and contemporary vegetation patterns. The surface pollen transect spanned four vegetation zones––alpine meadow, steppe, steppe desert and desert––under different climatic/elevational conditions. Relative representation (R _rel) values and Principal Components Analysis (PCA) were used to determine the relationships between modern pollen and vegetation and regional climate gradients. The results show that the main vegetation zones along the regional and elevational transects can be distinguished by their modern pollen spectra. Relative to Poaceae, a high representation of Artemisia, Nitraria and Chenopodiaceae was found, while Cyperaceae and Gentiana showed values in the middle range, and Ranunculaceae, Asteraceae, Ephedra and Fabaceae had low relative representation values. PCA results indicate a high correlation between the biogeoclimatic zones and annual precipitation and annual temperature and July temperature. The Artemisia/Chenopodiaceae ratio and the Artemisia/Cyperaceae ratio are useful tools for qualitative and semi-quantitative palaeoenvironmental reconstruction on the north-eastern Tibetan Plateau. Surface lake sediments are found to have different palynomorph spectra from moss cushion and soil samples, reflecting the larger pollen source area in the contemporary vegetation for lakes.     

The significance of pollen signal in present-day marine terrigenous sediments: The example of the Gulf of Lions (western Mediterranean Sea)

Terrigenous marine and continental sediments constitute pollen reservoirs able to fulfil the lack of Pleistocene data extracted from autochtonous lacustrine sediments for palaeovegetation reconstructions. Nevertheless, it is essential to constrain the signification of the pollen signal in such still under-exploited sediments. This article introduces a study carried out on the shelf of the Gulf of Lions. It aims at (1) highlighting the transport of pollen grains and the signification of the record in that area as well as at (2) discussing the pollen transport in various sedimentary contexts. The present study was carried out on surface water of the Rhone River, sea water and surface sediments of the shelf of the Gulf of Lions. The pollen content of the Rhone River and of its plume is in agreement with the season of sampling. On the contrary, the other samples of sea water did not contain any pollen grain. The samples from the surface sediments of the shelf are very rich in pollen grains and evidence the absence of sorting by currents except for Pinus. Indeed, its relative abundance increases when the fluvial impact decreases. The pollen signal (except Pinus) records the phytogeographic changes linked to the different drainage basins. The riparian woodland is over-represented at the Rhone River mouth. In front of the Pyreneo-languedocian rivers, Mediterranean xerophytes pollen grains are more abundant. From these results and from others taken in the literature, a model of pollen transport is built: all thin terrigenous sediments contain pollen grains essentially transported by rivers while autochthonous sediments (i.e., found in closed lakes or marine pelagic sediments) contain pollen grains brought by wind. Small closed lakes (or peat bogs) pollen spectra document local floras while those from pelagic marine sediments document floras from several hundred square kilometres.     

The interpretation of fen carr pollen diagrams: pollen–vegetation relationships within the fen carr

Sediments deposited in fen carr environments are frequently used in pollen-based reconstructions of vegetation history, although few studies of modern pollen–vegetation relationships in fen carr have been undertaken to aid the interpretation of fossil assemblages. It is often assumed that the wetland pollen signal will reflect the overall composition of the fen carr system although, given the closed canopy context, it may actually be dominated by local vegetation elements. This paper seeks to determine whether the vegetation heterogeneity within a modern fen carr is reflected in the pollen signal of the wetland taxa. Vegetation composition was recorded around a series of moss polster sample points at Calthorpe and Wheatfen, two areas of fen carr in the Norfolk Broadland, UK. The spatial interrelationship between the wetland components of the vegetation and their palynological equivalents in the moss polster pollen assemblages is explored through trend surfaces and a correlation based statistical test (the Mantel Test). The representation of the major vegetation components in the pollen record is highly variable. Values for Alnus glutinosa, a dominant canopy species, range from 5.3% to 73% of total land pollen (TLP) at Calthorpe and 2.5–61% TLP at Wheatfen. In part, this variation reflects the enhanced input of pollen from dry land sources close to the wetland edge although the variation recorded at points equidistant from the dry land margin suggests that other processes are also influential. The strongest positive associations in the Mantel Test are for under-storey and ground flora elements that are poorly represented in the pollen record. Taxa for which there is no association include those with few vegetation occurrences and those for which a large proportion of the pollen appears to have been derived from outside the study areas. We conclude that modern pollen samples collected from fen carr broadly reflect the vegetation composition of the study areas. However, the input of pollen from fen carr species is spatially inconsistent. Differences between the sample points can be attributed to the distribution and pollination biology of the under-storey and ground flora elements and the structure of the vegetation around sites. The results caution against assuming that changes in the pollen representation of wetland taxa, in fossil sequences constructed from fen carr deposits, necessarily reflect changes in the overall composition of the community.     

An assessment,using stable isotopes,of the importance of allochthonous organic carbon sources to the pelagic food web in Loch Ness

The natural abundance of stable isotopes (δ¹³C and δ¹³15N) was determined for components of the pelagic food web in Loch Ness, a deep oligotrophic lake in northern Scotland, and compared with values from the inflow rivers and the catchment vegetation. Phytoplankton δ¹³C was low compared to values reported from other lakes, possibly reflecting a high use of ¹³C-depleted carbon dioxide from respired organic matter before further isotopic fractionation during photosynthesis. Phytoplankton δ¹³C was appreciably lower than that of dissolved and particulate organic matter (DOM and POM) in the loch. The DOM and POM were evidently overwhelmingly of allochthonous origin and ultimately derived from terrestrial plant detritus. The distinctive δ¹³C values for phytoplankton and detritus in the loch allowed the use of food sources by grazing crustacean zooplankton to be assessed, and the contributions of phytoplankton carbon and detrital carbon to zooplankton total body carbon appeared to be about equal. Comparison of δ¹³C and δ¹⁵N values for zooplankton and fish allowed assessment of trophic structure in the loch. The very high dependence of the pelagic food web in Loch Ness on allochthonous organic matter inputs from the catchment may be exceptional in a large lake, but has important implications for our understanding of lake ecosystem processes as well as for lake management.     

Notes on the Scottish flora

Background: Abernethy Forest, an area of native Caledonian Scots pine forest, includes areas of open or sparsely forested peatland, largely ombrotrophic, but locally influenced by water from mineral ground.

Aims: To relate vegetation species-composition to the limit of ombrotrophy inferred from water chemistry.

Methods: Species composition was listed in 300 sample quadrats; chemical analyses of water samples were available from 200 of these. The vegetation data were classified using TWINSPAN. Relationships between vegetation and chemical results were analysed statistically using contingency tables.

Results: Eleven ‘vegetation types’ were recognised, five (189 samples) judged as primarily ombrotrophic-bog vegetation. The others showed evidence of flushing, or transition to forest. A previous analysis of major cations in the 200 water samples indicated that a limiting Ca/Mg ratio of 1.0 divided 61 samples showing telluric influence from 139 essentially ombrotrophic samples. Neither division was sharp. Contingency tables showed that the division based on vegetation types was strongly associated with the division of the water samples based on cation ratios.

Conclusions: Large areas of bog south and east of Loch Garten are unequivocally ombrotrophic; the main telluric-influenced sites occupy openings in the forest west of the loch. The limited ‘fen’ flora probably reflects the small size and isolation of the calcium-rich sites and the dominating influence of the ombrotrophic peats.     

Tracing and interpreting fine-scale human impact in northern Fennoscandia with the aid of modern pollen analogues

A set of moss samples (n=58) was collected and analysed to obtain modern pollen analogues for both natural and human induced vegetation types in northern Fennoscandia. Vegetation types with settlement, trampling and grazing were selected from the different latitudinal forest zones (birch, pine, and mixed forest). The moss species varied between samples but the size of the sample was kept constant. Numerical analyses such as Redundancy Analysis (RDA) indicate that in northern areas human presence is mainly visible as increased values of Gramineae, Rumex acetosa/acetosella, R. obtusifolius, Solidago-type and Achillea-type pollen. Partial RDA further reveals that settlement is strongly correlated with high values of Rumex acetosa/acetosella, whereas trampled sites are significantly characterised by Gramineae and, to a smaller extent, Cyperaceae pollen. It is therefore possible to distinguish the impact of different types of human activities on vegetation on the basis of the pollen spectra. Later, these data will be used to interpret the presence, nature, and duration of human impact from fossil records from the same area. Received June 6, 2001 / Accepted December 12, 2001     

Comparing pollen spectra from modified Tauber traps and moss samples: examples from a selection of woodlands across Europe

This paper compares pollen spectra derived from modified Tauber traps and moss samples from a selection of woodland types from Bulgaria, the Czech Republic, Georgia, Greece, Poland, Switzerland and Wales. The study examines the representation of individual taxa in the two sampling media and aims to ascertain the duration of pollen deposition captured by a moss. The latter aim was pursued through the calculation of dissimilarity indexes to assess how many years of pollen deposited in a pollen trap yield percentage values that are most similar to those obtained from the moss. The results are broadly scattered; the majority of moss samples being most similar to several years of pollen deposition in the adjacent trap. For a selection of samples, a comparison of the pollen accumulation rate in pollen traps with the pollen concentration in the moss per unit surface indicates that the entrapment and/or preservation of individual pollen types in the moss differ from that in the pollen trap. A comparison of the proportion of different taxa in the moss with the pollen spectrum of 2 years of pollen deposition in the trap also revealed large differences. There is a tendency for bisaccate grains such as Pinus and Picea to have a higher representation in moss than in traps but there is considerable regional variation. The results indicate that pollen proportions from moss samples often represent the pollen deposition of one area over several years. However, bisaccate pollen grains tend to be over-represented in moss samples compared to both pollen traps and, potentially, lake sediments.     

Changes in the relative abundance of Polyarthra vulgaris and P. dolichoptera,following the elimination of fish

In the last decade palynological investigations of the sediments of large lakes in the USSR (Ladoga, Onega, Kubenskoye, Vozhe, Lacha, Beloye) and the Baltic Sea have been carried out. Limnic and marine sediments are notable for their high pollen and spore concentrations and pollen diagrams from these deposits have a regional significance. The diagrams from Lakes Onega and Ladoga belong to the Kola-Karelian type while the sediments of Lacha, Vozhe, Kubenskoye and Beloye belong to the North-Russian type (Neustadt 1957). Three types of Holocene pollen diagrams are registered for the Baltic Sea. Modern and fossil pollen spectra reflect the composition of the zonal vegetation. Late-glacial sediments are found in several profiles of lake and sea deposits. Holocene deposits are divided into eight stratigraphic zones: Late-glacial sediments (DR₂-Al-DR₃) and Post-glacial sediments (PB, BO, AT, SB, SA). The zonations of lake and sea sediments can be clearly correlated.     

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.     

Pollen distribution in the bottom sediments of the western North Atlantic Ocean

An extensive survey of pollen distribution in surficial sediments from the slope and rise between 29° and 72°N, shows patterns which reflect the distribution of vegetation in eastern North America and the effects of aeolian, fluvial, and marine sedimentation. The results of the analysis of 94 core tops show that pollen concentration ranges from 23,000 to 10 grains per gram, and decreases offshore with distance from vegetation sources and northward in relation to lower vegetation density. Maximum concentration occurs in organic-rich, fine-grained lutum deposited off rivers draining the southeastern United States. Low pollen concentrations are associated with high-energy environments, such as the Western Boundary Undercurrent and the Labrador Current. Percentages of dominant pollen types generally correspond with onshore distribution of pollen and vegetation. Oak and other deciduous pollen types are the most abundant in marine sediments off the deciduous forests of eastern United States. Hemlock pollen is essentially restricted to sediments off the hemlock-oak forests of northeastern U.S. and southeastern Canada, and spruce pollen is most abundant adjacent boreal forests and forest-tundra formations. The relative increase in pine pollen seaward reflects the selective effects of marine transport. Synthesis of pollen data by Q-mode factor analysis shows that major vegetation formations of eastern North America produce distinctive, regional marine pollen assemblages.     

The physico-Chemical limnology of Loch Lomond

Loch Lomond, the largest freshwater lake in Britain, is physically divided into a number of distinct basins by geological structures. The northern part of the loch is long, narrow and deep whilst the southern area is broad and shallow with many islands scattered throughout it.The water chemistry of the loch is dictated by the geology of catchment and the quality of the rivers flowing into it. The two major rivers, the Falloch and the Endrick Water, are both of good quality with low nutrient content. Consequently, the loch is regarded as oligotrophic, though the southern area verges towards being mesotrophic.