Can we detect a west Norwegian tree line from modern samples of plant remains and pollen? Results from the DOORMAT project

In the DOORMAT (Direct Observation of Recent Macrofossils Across Treeline) project, the modern representation of local vegetation by pollen and plant remains (plant macrofossils) across a west Norwegian tree line, composed of Betula pubescens and Pinus sylvestris, has been studied over 2 years. The aim was to discover if the modern tree line could be detected and therefore how precisely past tree-line movements could be reconstructed and related to Holocene climate changes by using one proxy or a combination of both. Traps were placed in the vegetation from 663 to 1,120 m a.s.l., spanning the pine altitudinal species limit, the birch tree-line ecotone, and the vegetation zones up to the mid-alpine zone. Three traps were also set in the small lake Trettetjørn close to the modern tree line at 800 m a.s.l. Traps were emptied twice a year to sample both summer and winter seasons. Macrofossils represent their local vegetation well. However, tree Betula remains were trapped above the tree line and Pinus and Picea remains were recorded 1.0–1.5 km away from their sources, demonstrating considerable dispersal capacity. This shows that rare macrofossil remains do not necessarily represent the local presence of these trees. Aerial tree pollen deposition in traps at the upper limit of pine woodland and in the subalpine birch woodland was unexpectedly low, whereas pollen accumulation rates (PAR) were orders of magnitude higher in the lake traps. We hypothesise that the lake receives regional pollen rain washed in from its catchment by snow meltwater and that high values in traps are due to continuous suspension of pollen in the lake water during summer. The interpretation of tree-line changes from existing Holocene pollen and plant macrofossil data from Trettetjørn was supported and refined by the DOORMAT macrofossil data, but the modern pollen data were anomalous.     

Archaeobotany of the Old Prague Town defence system, Czech Republic: archaeology, macro-remains, pollen, and diatoms

The stone fortification of Prague Old Town was constructed shortly after A.D. 1230. Large-scale archaeological and archaeobotanical research has been carried out in this area in recent years. The deep moat and the drainage ditch have been excavated at several places in front of a line of former ramparts. During the initial period of their existence, the moat and the ditch contained only minimal waste. A sudden change in the management of these features occurred at the beginning of the 14th cent., when they started to be filled with large amounts of rubbish. This article concentrates on the evaluation of these rubbish strata at three profiles through the drainage ditch and one profile through the dry moat. The results of plant macro-remains (including wood), pollen, and diatom analyses clearly demonstrate the development of the local (aquatic) environment from being relatively clean to highly polluted. An actualistic (phytosociological) approach was adopted to reconstruct the local environment of the town periphery in detail, and to distinguish a variety of habitats that were situated in the economic hinterland of the city. Some species from the rich assemblages of plant remains are interpreted in terms of their use in High Medieval households. Diatom analysis completes the picture by providing the proxy data on water quality history in the drainage ditch. Received September 15, 2001 / Accepted May 21, 2002     

Old World and New World Cupressus pollen: morphological and cytological remarks

The aim of this work was to collect new information about pollen morphology and pollen wall structure comparing Cupressus species from the Old World and New World. Cupressus is a nonmonophyletic genus that includes species that appear to be divided into Old World and New World clades. Observations in this study on cypress pollen indicate that grain size and composition of intine β-glucans are different between the considered Old World and New World species. Different from all the other American cypress species, pollen of C. macrocarpa reacted to dying in a similar manner to Old World species. Rehydrated pollen grains collected from 20 Asian, Afro-Mediterranean and American cypress species were measured under a light microscope. The size of the pollen grains and the percentage of intine in relation to the pollen grain diameter were significantly different between Old World and New World species. Pollen wall composition was tested after addition of different dyes to the hydration solution, and subsequent observations were carried out by light and fluorescence microscopy. Lugol and calcofluor staining showed differences in composition of the middle and inner intine layers between New World and Old World species.     

丽江城市不同区域景观美学

我国正处在快速城市化阶段,古城或古镇正面临城市化和旅游业的双重压力,古城风貌和景观美学受到影响。以世界文化遗产丽江古城所在地——丽江盆地为研究对象,按照大研古镇-丽江新城-郊区的梯度设置评价样点,采用评分法(SBE-Scenic beauty estimation procedure)进行景观美学评价。结果显示:参加评价的男性与女性显示出性别间的细小差异,两个曲线的趋势基本一致,其中最大值和最小值均由男性评价者产生。丽江古城的三大古镇中,束河古镇的景观美学评价值最高,为6.79,白沙古镇的景观美学评价值为5.82,大研古镇景观美学评价值为5.68。大研古镇与丽江新城交错带的景观美学评价均值为4.9,丽江新城区的平均值均值为4.03,玉龙县城的均值为3.88,评价分值最低点在玉龙新县城,为2.36。在世界文化遗产核心区大研古镇到郊区的梯度上,整个评价曲线呈V型,中间低,两边高的趋势,大研古镇与城市新区交错带的景观美学评价值较低。景观美学评分较高的景观要素包括:自然景观(山体、水体)、农田、古镇、有历史文化特色和地方民族特色的景观等,现代城市景观的评价值较低。研究结果对保护丽江古城具有重要意义,可以用来完善丽江城市景观规划、优化城市空间格局,在城市建设发展的同时,保护好丽江的古城风貌和田园风光,以维系景观美学价值。     

Holocene forest development along the Setesdal valley, southern Norway, reconstructed from macrofossil and pollen evidence

The Setesdal valley in South Norway runs north to south for 200 km, from alpine vegetation at 1200 m, passing the tree-line at around 1000 m, through Boreal forests, to Nemoral forest at sea level. The Holocene vegetation history and its altitudinal differentiation were reconstructed using pollen percentages and influx and plant macrofossil concentration records from four lakes along an altitudinal transect. During the early Holocene (c. 10500–8000 cal b.p.) Betula pubescens, Pinus sylvestris, Alnus, and Corylus expanded in the lowlands. Only Pinus and B. pubescens reached 1000 m asl (Lille Kjelavatn). Only B. pubescens reached Holebudalen (1144 m asl) at about the same time as it arrived in the lowlands. Between c. 8000–3000 cal b.p. mixed deciduous forest developed around Dalane (40 m asl) and to a lesser extent around Grostjørna (180 m asl), birch woodland with pine surrounded Lille Kjelavatn and birch woodland occurred at Holebudalen. From c. 3000 cal b.p. to present, the vegetation at Dalane hardly changed except for slight human impact and the immigration of Picea abies. At Grostjørna Pinus expanded. At Lille Kjelavatn Pinus disappeared and Betula became sparse as at the tree-line today. Betula retreated from Holebudalen thus leaving it above the tree-line in low-alpine vegetation. The strengths and weaknesses of pollen and plant macrofossil data were assessed for forest reconstructions. Where local pollen production is low, as near the tree-line, percentages of long-distance tree pollen can be misleadingly high. Pollen influxes of Betula and Pinus were much smaller near their altitudinal limits than at lower altitudes, although their macrofossils were equally abundant. The limited dispersal capacity of macrofossils documents the local presence of species and the character of the local vegetation, although macrofossils of some tree taxa are rarely found. Pollen and plant macrofossil evidence complement each other to provide a more complete reconstruction of Holocene tree-limits and tree-lines and hence climate changes, than either form of evidence alone.     

长江三峡库区玉溪遗址的环境与人类活动的孢粉记录

对玉溪遗址剖面53个地层样品和附近山地的9个现代表土样品进行了孢粉分析,结果表明:玉溪文化时期(6 000a.BP—7 200a.BP)孢粉组合以乔灌木为主,草本次之,蕨类孢子含量较少。乔灌木中以松属、桦木属、栎属占绝对优势;草本中藜科、蒿属含量较高;蕨类孢子以凤尾蕨为主。孢粉谱反映了较为湿润的气候特征。表土孢粉以超代表性的松属占绝对优势,可能是植被群落演替过程中松属植物作为先锋种类表现在演替初级阶段具有明显生长优势的结果,表土孢粉组合基本与当地植被状况相吻合。本研究在14 C测年基础上结合其它代用指标发现在剖面上部第4—8层和下部淤泥层中Rb/Sr值较高且乔灌木花粉百分含量明显增加,可以解释为该段淤泥层为古洪水遗迹,洪水可能有利于对孢粉的富集。地层剖面中文化层与淤泥层呈现交互沉积,表明周期性的洪水事件影响到了人类活动的连续性。     

Vegetation history and lake-level changes from the Younger Dryas to the present in Eastern Pyrenees (France): pollen, plant macrofossils and lithostratigraphy from Lake Racou (2000 m a.s.l.)

A multi-proxy palaeoecological investigation including pollen, plant macrofossil, radiocarbon and sedimentological analyses, was performed on a small mountain lake in the Eastern Pyrenees. This has allowed the reconstruction of: (1) the vegetation history of the area based on five pollen diagrams and eight AMS¹⁴C dates and (2) the past lake-level changes, based on plant macrofossil, lithological and pollen analysis of two stratigraphical transects correlated by pollen analysis. The palaeolake may have appeared before the Younger Dryas; the lake-level was low and the vegetation dominated by cold steppic grasslands. The lake-level rose to its highest level during the Holocene in the Middle Atlantic (at ca. 5060±45 b.p.). Postglacial forests (Quercetum mixtum and Abieto-Fagetum) developed progressively in the lower part of the valley, while dense Pinus uncinata forests rapidly invaded the surroundings of the mire and remained the dominant local vegetation until present. The observed lowering of the lake levels during the Late Atlantic and the Subboreal (from 5060 ± B.P. to 3590±40 b.p.) was related to the overgrowth of the mire. The first obvious indications of anthropogenic disturbances of the vegetation are recorded at the Atlantic/Subboreal boundary as a reduction in the forest component, which has accelerated during the last two millennia.     

The spread of<Emphasis Type="Italic"> Abies</Emphasis> throughout Europe since the last glacial period: combined macrofossil and pollen data

In this paper we reconstruct the location of the last glacial refugia and postglacial spread of Abies throughout Europe based on combined pollen and macrofossil data. More than 208 pollen sequences available in the European Pollen Database (EPD) and 38 macrofossil sites are used to produce distribution maps encompassing a time span between 38000 and 5500 years B.P. The investigation excludes more recent periods, because these could be strongly influenced by human impact. The pollen data presented here confirm long-lasting refugial areas such as southern Italy and Greece already described in previous studies. The combined pollen/macrofossil dataset identifies the Pyrenees as a further important refugium. In addition the pollen data indicate potential refugia in south-east France and north-west Italy. Possible migration tracks of Abies are discussed by comparing the palaeobotanical evidence with isozyme studies on gene markers of recent fir populations.Electronic Supplementary Material Supplementary material is available in the online version of this article at http://dx.doi.org/10.1007/s00334-004-0049-4.     

Plant remains from the early modern garden of the manor of Kumpula,Helsinki, Finland: an alternative sampling method for macrofossil analysis

Garden history can be investigated through archaeobotanical research. This paper discusses the plant remains which were obtained from the soil of the historical garden of the manor of Kumpula in Helsinki, Finland. This study was an experiment to enable macrofossil analysis without archaeological excavations. The aim was to develop an alternative method for sampling for macrofossils also including radiocarbon dates, and to evaluate the usability, cost and functionality of this method. The character of the garden was also considered. The soil samples for macrofossil analysis were collected from the garden from three to eight different levels using an end-filling open-ended sampler. A total of 38 one litre soil samples from eight different pits yielded 2,036 identified macrofossils, mostly seeds. These comprised 63 different taxa, of which 26 were identified to species level with certainty. Taxa with more than 25 seeds found were Chelidonium majus, Chenopodium spp., Juncus spp., Rubus idaeus, Sambucus racemosa and Urtica dioica. Important species were Secale cereale and Hordeum vulgare. Nine AMS radiocarbon dates were obtained from macrofossil material from four different pits, giving results ranging from 1120–920 cal bc to cal ad 1680–1930 for charred wood, and from cal ad 1450–1640 to 1640–1930 for charred grains of Secale cereale and seeds of Chenopodium album. The sampling method proved to work reasonably well, considering the limitations of the sample size.     

Late Glacial and early Holocene environment in the middle Lahn river valley (Hessen,central-west Germany) and the local impact of early Mesolithic people—pollen and macrofossil evidence

The Late Glacial to early Holocene river valley landscape of the middle Lahntal in Hessen, central-west Germany, is reconstructed by means of pollen and macrofossil analyses. AMS ¹⁴C dating combined with pollen, macrofossil and geomorphological mapping provide a detailed chronology of the floodplain sediments of the river Lahn. Archaeological evidence for early Mesolithic settlements in the middle Lahntal is backed up by pollen and macrofossil evidence, which indicates an increase in light demanding plants, together with ones indicating nutrient-rich and disturbed environments and more macroscopic charcoal and charred pieces of pine. Different phases of human impact were dated; a first phase was dated at ca. 9,270 uncal b.p. and a second phase at ca. 9,120 uncal b.p. Comparison of the palynological data from different fluvial channel fills demonstrates that during this second phase, between ca. 9,120–8,700 uncal b.p., there were several cycles of woodland clearance. The outcome of combined archaeological, palynological and macrofossil data is discussed in terms of the impact of early Mesolithic people using fire on the vegetation during the early Holocene in the middle Lahntal.     

Molecular‐ and pollen‐based vegetation analysis in lake sediments from central Scandinavia

Plant and animal biodiversity can be studied by obtaining DNA directly from the environment. This new approach in combination with the use of generic barcoding primers (metabarcoding) has been suggested as complementary or alternative to traditional biodiversity monitoring in ancient soil sediments. However, the extent to which metabarcoding truly reflects plant composition remains unclear, as does its power to identify species with no pollen or macrofossil evidence. Here, we compared pollen‐based and metabarcoding approaches to explore the Holocene plant composition around two lakes in central Scandinavia. At one site, we also compared barcoding results with those obtained in earlier studies with species‐specific primers. The pollen analyses revealed a larger number of taxa (46), of which the majority (78%) was not identified by metabarcoding. The metabarcoding identified 14 taxa (MTUs), but allowed identification to a lower taxonomical level. The combined analyses identified 52 taxa. The barcoding primers may favour amplification of certain taxa, as they did not detect taxa previously identified with species‐specific primers. Taphonomy and selectiveness of the primers are likely the major factors influencing these results. We conclude that metabarcoding from lake sediments provides a complementary, but not an alternative, tool to pollen analysis for investigating past flora. In the absence of other fossil evidence, metabarcoding gives a local and important signal from the vegetation, but the resulting assemblages show limited capacity to detect all taxa, regardless of their abundance around the lake. We suggest that metabarcoding is followed by pollen analysis and the use of species‐specific primers to provide the most comprehensive signal from the environment.     

Long-term vegetation dynamics and the infilling process of a former lake (Švarcenberk, Czech Republic)

Natural lakes are a rare phenomena within extraglacial areas of Central Europe. Almost all have been completely terrestrialized during the Holocene. This paper deals with one such former lake, located in southern Bohemia. Its extensive lacustrine and peat deposits were subjected to a multidisciplinary study that resulted in high-resolution pollen, macrofossil, algal and sediment-chemistry data interpreted in terms of past climate, geomorphology, soil, and regional vegetation development over the last 16,000 years. Against the background of these large-scale processes, local development took place, comprising the lake’s ontogeny from an arctic-type ecosystem hosting pioneer aquatic communities, through a highly diversified mosaic of eutrophic hydrosere habitats (shallow pools,Phragmites andCarex fen, alder carr), towards an oligotrophic mire that started to dome over the terrestrialized lake. At every individual development stage, specific processes characterized ecosystem function and composition: during the Late-Glacial with its rapid climatic changes, external forces induced the major stresses; while during the Holocene, autogenic changes of the wetland ecosystem played the most important role.     

An evaluation of the airborne pollen–precipitation relationship with the superposed epoch method

A statistical analysis was performed on a sample of 4 days sequences in which rainfall occurred during the second day, in order to establish airborne pollen–precipitation relationships. Drizzle (precipitation ≤ 1 mm) and rain (precipitation > 1 mm) events were considered as well as arboreal or non-arboreal pollen. Results showed that arboreal pollen decreases synchronically with the occurrence of a drizzle, registering the minimum value the following day of the precipitation occurrence. The high frequencies of fog and wet soil associated with the following day of drizzle occurrence support the hypothesis that besides the wash out, airborne pollen concentration could decrease by nucleation and impaction processes. As well, diurnal variation suggests that high humidity conditions could also affect pollen emission as already demonstrated by other authors.     

A comparative study of ancient sedimentary DNA, pollen and macrofossils from permafrost sediments of northern Siberia reveals long-term vegetational stability

Although ancient DNA from sediments (sedaDNA) has been used to investigate past ecosystems, the approach has never been directly compared with the traditional methods of pollen and macrofossil analysis. We conducted a comparative survey of 18 ancient permafrost samples spanning the Late Pleistocene (46-12.5 thousand years ago), from the Taymyr Peninsula in northern Siberia. The results show that pollen, macrofossils and sedaDNA are complementary rather than overlapping and, in combination, reveal more detailed information on plant palaeocommunities than can be achieved by each individual approach. SedaDNA and macrofossils share greater overlap in plant identifications than with pollen, suggesting that sedaDNA is local in origin. These two proxies also permit identification to lower taxonomic levels than pollen, enabling investigation into temporal changes in species composition and the determination of indicator species to describe environmental changes. Combining data from all three proxies reveals an area continually dominated by a mosaic vegetation of tundra-steppe, pioneer and wet-indicator plants. Such vegetational stability is unexpected, given the severe climate changes taking place in the Northern Hemisphere during this time, with changes in average annual temperatures of >22 °C. This may explain the abundance of ice-age mammals such as horse and bison in Taymyr Peninsula during the Pleistocene and why it acted as a refugium for the last mainland woolly mammoth. Our finding reveals the benefits of combining sedaDNA, pollen and macrofossil for palaeovegetational reconstruction and adds to the increasing evidence suggesting large areas of the Northern Hemisphere remained ecologically stable during the Late Pleistocene.     

Palaeoecological evidence for the middle and late Holocene vegetation, climate and land use in the upper Don River basin (Russia)

New pollen and plant macrofossil data, backed by radiocarbon dates, from the Kulikovo battlefield area in the forest-steppe region of the Upper Don River basin (central part of European Russia) indicate that the area was covered by mosaic vegetation in the second half of the Holocene. Steppe communities dominated during the mid—late Atlantic (7.2–5.7?cal.?kyr b.p.) and early Subatlantic (2.7–2.4?cal.?kyr b.p.), while forest-steppe dominated during the Subboreal (2.7–5.7?cal.?kyr b.p.), middle and late Subatlantic (2.4?cal.?kyr b.p. – present). Climatic reconstructions based on these data show that landscape dynamics in the region were most probably driven by changes in effective moisture: an excess of precipitation over evaporation. Even small reductions in annual precipitation, accompanied by a rise in summer temperatures by 1–3°C above present values, were sufficient to increase the proportion of steppe communities within this landscape complex, and also probably resulted in higher frequencies of wildfires. Signals of anthropogenic disturbance of vegetation are clearly pronounced in the pollen and plant macrofossil records since the middle Atlantic. However, human-induced changes in the vegetation remain subtle until the medieval period.     

Andromonoecy in an Old World Papilionoid legume,Erophaca baetica

Andromonoecy (i.e. the occurrence on individual plants of hermaphroditic and male flowers) is a rare sexual system among the angiosperms, regarded by some authors as a transitional stage from hermaphroditism to monoecy. Having discovered the occurrence of andromonoecy in Erophaca baetica (a Mediterranean shrubby legume with two subspecies), a novelty for Old World papilionoid legumes, we investigated the morpho‐functional correlates and the geographical distribution of this phenomenon in the species. The relative frequencies of hermaphrodite and male flowers were determined in two field and 111 herbarium populations. Biomass allocation within flowers, pollen production and viability, pollen tube growth, nectar production and the temporal pattern of male flower production were also studied in two nearby southern Spanish populations. Virtually all of the studied populations were andromonoecious. Male flowers tended to appear at apical positions within the inflorescence, and became more abundant by the end of the flowering season. Male flowers were externally similar to hermaphroditic flowers (although with less biomass and smaller parts) and released equivalent amounts of pollen and nectar; however, their pollen germinated significantly better. Erophaca is the first example of an andromonecious Papilionoid in the Old World. Since the main difference among floral morphs in this species is functional (i.e. pollen germination rate) rather than morphological, andromonoecy is not readily noticeable, and very careful inspection may be required to reveal it. The potential effect of andromonoecy in enhancing outcrossing rate in this species is discussed.     

The long-term development of urban vegetation in Helsinki, Finland: A pollen diagram from Töölönlahti

A core from Töölönlahti, a bay close to the central business district of Helsinki, the capital of Finland, was analysed for its pollen stratigraphy. The extrapolated age-depth curve based on¹³⁷Cs dating, sphaeroidal carbonaceous particle analysis and historical data suggests that the core goes back to the 17th or early 18th century. Four local pollen assemblage zones (LPAZ) were identified. Detrended correspondence analysis was used to detect the main phases of change represented by these zones. The basal LPAZ indicates that the vegetation was characterized by forest dominated byPinus sylvestris andPicea abies during the early history of the city of Helsinki and that human impact remained relatively minor until the end of the 19th century. A drastic change in vegetation took place at the turn of the 19/20th century when bothPinus andPicea declined rapidly while pollen values of several anthropogenic indicators rose. These changes were synchronous with the rapid expansion of population and urban area in Helsinki. Rarefaction analysis shows that palynological richness was highest during the early phase of increased human impact but then started to fall. During the 20th century the main vegetational trend has been the proliferation of introduced park trees. Palynological richness has decreased.     

Air and surface soil samples – two different pairs of shoes? Comparing the pollen spectrum on different days of the pollen season

The pollen spectra of air and surface soil samples from a rooftop (at 14 m) and from ground level (at 1.6 m) in the suburbs of Vienna (Austria) were compared. Two soil samples and two air samples were taken on four different days to account for possible differences: in winter when no pollination occurred (reference day), in spring during the main flowering of Betula (birch day), in spring/summer during the main flowering of Poaceae (grass day), and in autumn during the main flowering of Ambrosia (ragweed day). Thirty-five different pollen types were used to describe the pollen spectra. Frequencies of certain pollen types reflect a seasonal impact on both the surface soil and air samples and show a similarity between air and soil samples on most of the days. However, the seasonal impact is higher in the air samples and shows a high consistency for ground and rooftop level. Kendall’s tau correlation coefficients further substantiate the similarities of the samples especially for the pollen season days. Exceptions include the winter day when pollination was low and the air samples recorded nearly no pollen at all, and the ragweed day when Ambrosia pollen was abundant in three of four samples but not in the ground surface soil sample. Thus, (1) air and surface pollen samples record similar signals during the pollen season but not during the ragweed and winter season and (2) air and surface pollen samples show the impact of local vegetation also in pollen traps located at different heights.     

Biome reconstruction from pollen and plant macrofossil data for Africa and the Arabian peninsula at 0 and 6000 years

Biome reconstruction from pollen and plant macrofossil data provides an objective method to reconstruct past vegetation. Biomes for Africa and the Arabian peninsula have been mapped for 6000 years bp and provide a new standard for the evaluation of simulated palaeovegetation distributions. A test using modern pollen data shows the robustness of the biomization method, which is able to predict the major vegetation types with a high confidence level. The application of the procedure to the 6000 years data set (pollen and plant macrofossil analyses) shows systematic differences from the present that are consistent with the numerous previous regional and continental interpretations, while providing a more extensive and more objective basis for such interpretations. Madagascar, eastern, southern and central Africa show only minor changes in terms of biomes, compared to present. Major changes in biome distributions occur north of 15°N, with steppe in many low-elevation sites that are now desert, and temperate xerophytic woods/scrub and warm mixed forest in the Saharan mountains. These shifts in biome distributions imply significant changes in climate, especially precipitation, between 6000 years and present, reflecting a change in monsoon extent combined with a southward expansion of Mediterranean influence.     

The biogeographical status of <Emphasis Type="Italic">Alnus crispa</Emphasis> (Ait.) Pursch in sub-Arctic southern Greenland: Do pollen records indicate local populations during the past 1500 years?

Phytogeographical studies of south-western Greenland suggest that Alnus crispa is not native to the far south of the island. Palynological investigations dating to the 1970s concluded that this was the case throughout the Holocene, with the regular occurrences of Alnus seen in pollen diagrams from this region explained as the result of long-distance transport of alder pollen from Canada. Recently, macrofossil evidence from an archaeological site in southern Greenland has emerged that indicates that alder was amongst the fuel resources available to the Norse settlers around AD 1000–1400. In light of this discovery, we present data from 13 pollen diagrams produced since 2008 to re-examine the past status of Alnus within southernmost Greenland over the last 1500 years. Only at one site with a very large pollen source area do Alnus pollen frequencies regularly exceed a threshold which may be interpreted as indicating a regional presence for the plant. This pattern is argued to be consistent with the presence of a small but variable regional population of the plant, perhaps restricted in its distribution to the inland district of Vatnahverfi.