Flora - Morphology, Distribution, Functional Ecology of Plants

Volume 206, Issue 11, November 2011, Pages 928934

Immediate shift towards source dynamics: The pioneer speciesConyza canadensis in an initial ecosystem
Markus Klemens Zaplataa, Susanne Wintera, 1, Detlef Biemeltb, 2, Anton Fischera, 3,
a

, ,

Technische Universitt Mnchen, Geobotany, Department of Ecology and Ecosystem Management, Hans -Carl-vonBrandenburg University of Technology, Chair of Hydrology and Water Resources Management, Konrad-Wachsmann-

Carlowitz-Platz 2, D-85354 Freising, Germany

b

Allee 6, D-03046 Cottbus, Germany http://dx.doi.org/10.1016/j.flora.2011.07.001 , How to Cite or Link Using DOI Permissions & Reprints

Abstract
The aim of this study was to detect a potential temporal shift from sink to source dynamics of pioneer plants during primary succession. The study was done in an initial ecosystem, namely an artificially constructed catchment on Pleistocene sands in eastern Germany. The import and export of seeds (including soil seed bank and seed rain), the development of vegetation cover and the population size of all pioneer species were recorded in 20052009. The interplay between the spatial distribution of seed rain and the prevailing wind direction provide evidence that seeds of Conyza

canadensis (Asteraceae) first and to a considerable amount left the developing ecosystem. A change
from a sink to source dynamics was detected already in the first year of both the existence of the site and occurrence of C. canadensis individuals within the site. Population patterns at the landscape unit scale demonstrated extensive auto-correlations in the coverage of this invasive plant at first. Our results underline the pioneer traits of C. canadensis as a prolific seeder and excellent wind disperser, which enables this species to become rapidly dominant over relatively large areas.

Keywords

Artificial catchment; Colonization potential; Ecosystem development; Initial phase; Population dynamics; Primary succession; Seed rain

Introduction
Sink and source are fundamental terms in population dynamics, describing major ecosystem variables. A source population or a source habitat provides a net export of individuals, and a sink is a net importer of individuals (Pulliam, 1988). There are relatively few empirical studies dealing with spatial sourcesink dynamics (Thomson, 2007). Particularly, the demographic rates of a given species have not been studied in different habitats which would allow assessing spatial variability in source sink dynamics. Moreover, published habitat-specific demographic data are mostly not sufficient to determine whether a habitat actually is functioning as a source or a sink (Pulliam and Danielson, 1991). Consequently, more insights into source and sink habitats are needed to understand the dynamics and potential of plant colonization, as well as to apply it to population management (Pulliam, 1988). For successful colonization of new habitats, long-distance seed transport is of great importance. Particularly in initial ecosystems wind is the predominant dispersal vector for seeds ( diaspores; Van der Pijl, 1982). The main factors influencing long-distance dispersal of anemochorous seeds are both wind speed and release height [Dauer et al., 2006, for Conyza canadensis (L.) Cronquist (horseweed)]. Additionally, settlement velocity of wind-dispersed seeds is a major component of their dispersal ability ( [Guo et al., 2008] and [Tackenberg and Stcklin, 2008]). Specific morphological structures (flying devices) control the mobility and settlement velocity of winddispersed seeds. Seeds of many Asteraceae, for example, show adaptations to long-distance wind dispersal. In a study of 19 composite species, C. canadensis exhibited the lowest settlement velocity (0.278 m/s, Andersen, 1993) which enabled the farthest dispersal. In other empirical studies it was found that 99% of the C. canadensis seeds disperse within a 100 m radius, but few up to at least 500 m. Nevertheless, they are sometimes found even up to hundreds of kilometres from the source plant ( [Dauer et al., 2007],[Dauer et al., 2009] and [Shields et al., 2006]). Hence, the dispersal distance of C. canadensis is several orders of magnitude greater than reported for most other plant species [according to Dauer et al. (2009) even greater than for any other spermatophyte].

Conyza canadensis solely reproduces generatively (Bhowmik and Bekech, 1993) and has a high selfcompatibility supporting high seed production even without neighbouring plants or pollinators ( [Mulligan and Findlay, 1970] and [Smisek, 1995]). When the population density is low (10 plants m2), the seed number per plant individual is about 200,000, which can be reduced to about 100,000 seeds

at higher density of about 200 plants m2 (Bhowmik and Bekech, 1993). Besides the high seed production, the germination rate of C. canadensis is also high (Jursik et al., 2008). Seedling emergence is highest on the soil surface, and seedlings do not emerge from seeds placed deeper than 0.5 cm belowground (Nandula et al., 2006). C. canadensis still germinates under moderate water stress conditions ( [Nandula et al., 2006] and [Neckar et al., 2008]) and therefore grows well on light and relatively dry soils (Jursik et al., 2008). The emergence of C. canadensis is highly variable depending on soil and air temperature, and rainfall (Main et al., 2006). Emerging mainly during April and September, C. canadensis is a winter or summer annual ( [Main et al., 2006] and [Shields et al., 2006]). It is native to North America and nowadays has a worldwide distribution predominantly in temperate climates (Shields et al., 2006). In its native range, it is able to be an effective ruderal plant, building up stands that approach monoculture densities up to 1500 plants m2 (Main et al., 2006). In summary, the literature provides sufficient ecological knowledge to claim that C. canadensis is a pioneer species and globally successful invader (Abhilasha and Joshi, 2009). The actual knowledge on the dispersal and establishment features of C. canadensis is mainly based on studies and experiments on a small scale and on simulations. Our paper focuses on the colonization and temporal as well as spatial distribution of C. canadensis within an artificially constructed catchment area with a size of 6 ha (landscape unit) in north-eastern Germany. After construction with loose sandy substrate, the catchment initially functioned as a sink for seeds. In the very early phase of succession C. canadensisbecame dominant. Using this field situation, we studied the temporal shift in population dynamics of this pioneer species, and focused on answering the question when the study area first became a seed source.

Materials and methods

Study system
The study site is an artificially built water catchment named Chicken Creek which is located in north east Germany in the open-cast lignite coal-mining area in the Lusatia region south of the town of Cottbus. The climate is cool-temperate with a mean annual rainfall of 563 mm and 8.9 C annual mean air temperature (January mean: 0.8 C, July mean: 18.4 C; German Meteorological Service, station Cottbus; averages from the period 19611990). The Chicken Creek catchment is a landscape unit with well-defined borders ([Schaaf and Gerwin, 2010] and [Schaaf et al., 2011]), and forms an elevated plateau within the more or less flat mining area. The sandy to loamy sand-grade material used for catchment construction originates from Pleistocene sediments which were taken from the fore-field of the adjacent active mine Welzow Sd, where they were deposited during the Saaleglacial period about 130,000300,000 years ago (Gerwin et al., 2009). The western and the eastern parts of the catchment were constructed in two different periods using almost the same material and the same construction process in 2005. Therefore, the eastern part is some weeks older than the western part. The size of the Chicken Creek catchment area is about 400 m by 150 m (about 6 ha)

with an altitudinal difference of about 15 m between the north-western (140 m a.s.l.) and the southeastern part (125 m a.s.l.) (Fig. 1; Gerwin et al., 2009).

Fig. 1. The 150 m by 400 m large water catchment Chicken Creek (grey) and the surrounding situation in 2005 and 2006: Localization and distribution of the seed rain traps (points and a) and raster points (squares and b) for sampling the soil seed bank and for recording the vegetation. The surface in November 2005 is displayed by a Digital elevation model (Schneider et al., 2011). Spacing of contour lines is 0.5 m. Figure options

In 2005 and 2006, the surroundings consisted of the active open-cast mine located west and north of the catchment. On the east side, a large reclamation area (a cultivated field with Medicago varia Martyn) was situated. Due to reclamation progress, spontaneous occurrence of C. canadensis therein was negligible. Thus, three sides (W, N, and E) surrounding the catchment area provided minor growth conditions for colonizer plant species. South of the catchment, the riverbed of the Chicken Creek stream, whose headwater region is the Chicken Creek catchment, was prepared and C. canadensis grew nearby (Fig. 1). None of these situations in the surrounding area was any closer than 30 m to the catchment interior since vehicle tracks and the catchment slopes were directly outside the fenced area. Within the catchment area, a weather station continuously stored data about the wind direction and wind speed calculating averages for every hour gained from measurements taken every minute. The measuring height was 2 m above ground (Biemelt and Nenov, 2010). Wind direction is classed to 30 degree sectors. We matched the wind data with the exposure intervals of the seed rain traps. Further meteorological parameters such as precipitation and potential evaporation were also measured or calculated continuously (Biemelt and Nenov, 2010).

There are three possibilities for plants to emerge in the catchment: (i) input by soil material (seed bank), (ii) input by wind (seed rain), animals and humans, or (iii) as descendants of Chicken Creek plants (on site reproduction). Consequently, we studied these three establishment pathways, except for animals and humans, because these vectors were regarded as having a low impact on the remote, fenced, and publicly non-accessible catchment area.

Soil seed bank and seed rain analyses

When the catchment construction was finished in October 2005, we took soil samples (10 cm 10 cm 10 cm each) of the initial catchment substrate at the grid points, where also the vegetation was monitored (Fig. 1). For the purpose of analyzing the soil seed bank and not the seeds just carried by the wind, the soil samples were collected from a depth of 2 12 cm. We analyzed the germinable number of seeds in the total sampled volume by applying the seedling emergence method from October 2005 till August 2007. In this method emerging seedlings indicate viable seeds in the analyzed soil substrate, thus this method provides the ecological significance, in contrast to filteringand-counting methods (Fischer, 1987). To overcome dormancy, the samples stored in plastic bags were exposed to field climate conditions (6 January to 28 February 2006; 23 29 January 2007; both periods included several days with temperatures below 0 C, with a minimum temperature of 13.5 C; for a detailed description see Zaplata et al., 2010). During exposure in a glasshouse, the soil material was carefully dug several times a year to impair the establishment of bryophyte carpets and to support the seeds and ensure they received enough radiation to start germination. We calculated the germination potential of C. canadensis in the initial soil seed bank (m2) as follows: total number of germinated C. canadensis seeds total number of soil samples1 100 (=correction value for an area of 1 m2) 20 (=correction value for the thickness of 0.005 m)1. The spatial distribution of C.

canadensis seeds in the catchment soil was tested for significant differences using t-tests (p < 0.05).
From October 2005 to September 2006, we equipped the catchment edge with 55 seed rain traps with a distance of about 20 m between each (Fig. 1). Traps consisted of a disposable Petri dish, 15 cm in diameter, which was completely filled by a filter paper, brushed with paraffin-based bag balm on the lower side in order to attach it to the Petri dish and on the upper side in order to trap seeds (Zaplata et al., 2010). The sticky surfaces of the seed traps were orientated almost horizontally, but slightly alternated in pointing in- and outwards of the catchment at about a height of 0.3 m. We replaced the traps 16 times during the entire one-year exposure period. The seeds were determined and counted using a binocular microscope. For analyses we grouped the seed rain traps according to their position along the catchment edge referring to the main directions north, south, west, and east; and we compared the total seed amounts and 16 separate exposure intervals between the trap groups using t-tests with p < 0.05.

Vegetation development
We recorded the vegetation in the catchment at the (semi-)terrestrial part in a 20 m 20 m regular grid (Fig. 1) using permanently marked plots with a plot size of 25 m2. Additionally, in the corners of the

plots, subplots of 1 m2 were established. Thus the monitoring consisted of a nested plot design with 119 plots of 25 m2including four 1 m2 subplots each (Fig. 1). The yearly vegetation records comprised (i) the plant species, (ii) the cover values estimated for each species according to a modified Londo scale (Londo, 1975; 0.1: 0.1%; 0.5: >0.10.5%; 1: >0.51%; 2: >12%, in 1% steps up to 10; 15: >1015%, in 5% steps up to 30; 40: >3040%, in 10% steps up to 100), and (iii) the number of individuals of each species that were present in each north-eastern 1 m2 subplot. In 2005, we recorded the vegetation immediately after the completion of the catchment in early October. Since 2006 we recorded the vegetation every year in the middle of July. We tested the C. canadensis cover distribution of the 25 m2 plots by surface pattern analysis for grid point pairs oriented in pre-specified directions, as proposed by Oden and Sokal (1986), and Legendre and Fortin (1989). Instead of a set of standard correlograms for each single direction, we used twodimensional correlograms for depicting the auto-correlations of all directions, thus it is immediately visible whether cover distribution is isotropic or anisotropic.

Estimation of seed production

The seed production of C. canadensis was estimated using the model of Bhowmik and Bekech (1993) which presents the number of seeds per plant individual depending on plant densities. The total individual number for the catchment was interpolated from the 1 m2 plots to the catchment area of 55,943 m2 (excluding the lake area).

Results
Soil seed bank and seed rain
From one third of the soil samples C. canadensis seedlings emerged, the proportional number was 3.89 seeds m2 and 217,556 in total, interpolated for the whole catchment area. In the eastern part, insignificantly more seeds were found and auto-correlation analyses showed that the C.

canadensis seeds were distributed in a random way at any spatial scale.

In the seed rain from October 2005 to July 2006 a total of 26 seeds were detected (2.44 m2), of which only four were C. canadensis (0.37 m2). However, 1996 seeds were found during the following seed shedding period in August and September 2006. Of these 1992 belonged to C. canadensis. Thus, in total 98.7% of all seeds found were of C. canadensis (151.3 m2). The second most frequent species was Chenopodium

album L.,

with

altogether

seven

captured

seeds

(0.53 m2).

Seeds

of Medicago varia were not found. The seed rain of C. canadensis was significantly different (p < 0.001) with an average of 8.9 (lower CI: 4.9, upper CI: 12.9) and 58.9 (lower CI: 41.7, upper CI 76.0) seeds per trap at the western and eastern catchment border, respectively. Thus, 6.6 times more C. canadensis seeds were caught at the eastern edge. For the low number of seeds of Ch. album, we did not detect a pattern.

Population development
Individuals of 18 different plant species, but not of C. canadensis, were present within the 1 m2 plots shortly after the completion of the catchment (Fig. 2, Zaplata et al., 2010). At that time Cirsium

arvense was the most frequent species (present at 12% of 118 plots). Ch. album was the second most
frequent species (present at 2.5% of the recorded 1 m2 plots), however individuals were mainly present as dwarf forms. ForCh. album the occurrence of plant individuals in the Chicken Creek catchment started one season earlier than for C. canadensis (but it was not detected in the initial soil seed bank analysis). The incremental growth rates of C. canadensis and Ch. album were similar ( = 20.2 and 19.7). One season after they initially occurred, both species acquired their maximum number of individuals and maximum frequency, which were 159 and 5 times higher for C.

canadensis than for Ch. album, respectively (Fig. 2). Medicago varia was not found.

Fig. 2. Conyza canadensis () and Chenopodium album (): total numbers of individuals (open symbols) and plots occupied (filled symbols). The data from the northeastern 1 m2 plots is presented; plots situated close to the lake are not considered in abundances. Standard deviations are given as error bars. Note the logarithmic scale of the primary y-axis. Figure options

C. canadensis and Ch. album were the two most frequent plant species at the early stage of
ecosystem development. In the first and second growing season, C. canadensis formed the main vegetation aspect in the initial catchment. This continued in 2008 in the eastern part of the catchment where pure sands dominate ( [Gerwin et al., 2009] and [Zaplata et al., 2010]). There were high spatial auto-correlations (25 m2 plots) for the C. canadensis cover in 2006 in the catchment (Fig. 3a), indicating that the distribution of C. canadensiswas a phenomenon at the site scale. The extent of auto-correlation was found to be almost twice as large at the 5% level of significance as at the 1% level of significance (the latter result is shown in Fig. 3b). A homogeneous zone ( symbols in Fig. 3b) was found to be at least 40 m in the eastwest direction and about 100 m in the northsouth direction. Thus, anisotropy was encountered, correlating with the main orientation of the catchment. In

fact, the C. canadensis cover was auto-correlated over an area roughly equivalent with the two catchment halves (in the case of the 5% level of significance), and in the case of the 1% level of significance with the four major sections (according to Schaaf and Gerwin, 2010). In subsequent years, autocorrelation continuously and markedly decreased.

Fig. 3. Distribution, cover, and spatial auto-correlation of Conyza canadensis in 2006 on 25 m2 plots. (a) Dots (four sizes) depict C. canadensis covers of 0.1, 0.5, 1, and 2 percent cover degree, respectively. (b) Autocorrelated neighbour plots in 2006. A neighbour of the centre grid cell (marked) corresponds to any pairs of catchment raster points with the same distance and direction. symbols indicate positive significant autocorrelations. symbols depict significant negative autocorrelations, and both significances indicate differences for alpha = 0.01. Cases of symbols indicate that no spatial autocorrelation was found for the considered distance class. Anisotropy is present: the northsouth range of C. canadensis is much larger (about 100 m) than the eastwest range (40 m). Figure options

Average density of individuals of C. canadensis was 4.0 plants m2 in 2006 (n = 115 plots, CI 1.3). This corresponds very well with the C. canadensis soil seed bank in autumn 2005 with 3.89 seeds m2. In 2007 population density reached its maximum with an average of 85.5 individuals m2 and decreased afterwards (Fig. 2). Approximately 44.8 109 C. canadensis seeds were produced in the landscape unit in 2006. This estimate is an extrapolation taken from the individual numbers of the 1 m2 plots in relation to the whole catchment and using data about seed production from Bhowmik and Bekech (1993). It corresponds to 205,714 times the starting pool (initial soil seed bank).

Effects of wind direction on seed dispersal of Conyza canadensis

Representing an elevated site in an open landscape unit, the Chicken Creek catchment area is exposed to relatively steady (8146 h of 8252 h total [99%]) and often strong winds (wind speeds of >4.4 m/s: 1997 h [24%] during the exposure period). The prevalent wind direction was from the west southwest (Fig. 4;Biemelt and Nenov, 2010).
Fig. 4. Box plots showing numbers of captured C. canadensis seeds (median, 25th and 75th percentiles, and outliers) in Chicken Creek's eastern and western trap spot line within three time intervals jointly ranging from 18 July until 20 September 2006. Below: the contemporaneous wind conditions, shown as relative wind frequencies. Therein the respective wind speeds are given (chequered hatching: 0.12.2 m/s, grey: 2.2 4.4 m/s, black: >4.4 m/s). Compared to the first interval (ranging from 18 July until 4 August; see left side), during the exposure time of the seed trap sets collected on 21 August (middle) and 20 September (right side) wind from the westsouthwest was even more frequent. Regarding wind directions, the latter interval additionally contains the largest proportion of high wind speeds from the west. Figure options

The first seed rain interval from 18 July until 4 August 2006 with numerous C. canadensis seeds comprised 404 h of traps exposure. During this period, wind was present every hour and mostly came from westsouthwest: 77 h (19.1% of the exposure time). The second interval with large numbers of seeds lasted from 4 until 21 August 2006 (404 h); 402 h with wind, only 2 h were calm. Again, the wind mostly came from the westsouthwest: 91 h (22.5% of the exposure time). The third presented seed rain interval lasted from 21 August until 20 September 2006 (716 h); calm periods did not occur. The wind mostly came from the westsouthwest: 197 h (26.1%). The numbers of seeds correlated negatively with the wind direction proportions: significantly more seeds were caught in the eastern traps the more frequent the wind came from western directions and vice versa (Fig. 4). During the seed trapping interval from 21 August to 20 September 2006, when the differences in C. canadensis seed amounts between the eastern and the western trap spot line were the highest, the proportion of wind from western directions, especially the proportion of wind with high speed, was the highest as well (Fig. 4). This pattern of seed rain suggests the following: first, the wind reaching the catchment did not contain many C. canadensis seeds, resulting in the comparatively low amounts detected in the western trap spot line. Mainly within the catchment area, seed uptake occurred due to the passing wind. A net export is given by the comparatively large amounts of C.

canadensis present at the eastern export border. Thus, the catchment had already become a large
seed source for C. canadensis in the first growing season.

Discussion
Conyza canadensis shows high seed production, high dispersal ability, high germination rates,
deferrable growth instead of dieback under dry conditions (from own observations), and a maximum population size which exceeded the second early-succession species Ch. album by 159 times in its number of individuals. These properties are the requirements for being a common ruderal and a globally successful species. Beginning with a 6 ha large sink, a vital C. canadensis population

developed, which not only built up very fast but also effectively supplied especially the eastern surroundings with numerous seeds already in the first season after it entered the initial catchment. Thus, our results indicate a change from sink to source dynamics within the landscape unit studied within just one year of succession.

Sink region and the fact of a soil seed bank in primary succession
Regarding the pathways how C. canadensis may have entered the newly constructed catchment, the soil seed bank is the apparently relevant means of advancement. Seeds of C. canadensis were already present in the soil immediately after catchment construction and their frequency nearly matched the frequency of individuals developed in the following season. However, it is guaranteed that no seeds as a substrate burden were involved as the substrate was virgin Pleistocene sediment taken from the fore-field of the lignite coal mine. Thus, all seeds originate from plants in the surroundings. We, moreover, assume that any concentrating mechanism for seeds is excluded, as the total wind-exposed substrate surfaces that existed during transportation processes (conveyor belts and loading areas for trucks) definitely will have been smaller than the 6-ha-surface of the Chicken Creek catchment. These points indicate that seed input by substrate material was of minor relevance and, in conclusion, support the view of the Chicken Creek as a very early succession site, which otherwise would require primary substrate free of viable seeds. Before mining activities, pine forests and other vegetation communities harboured C. canadensis only along disturbed trails at best. Thus it is unlikely that this species entered the initial substrate by contamination of recent topsoil. The persistent seeds of the non-forest plant and second important colonizer Ch. album were not detected in the initial soil seed bank. However, there are indications that unintentional seed entries were rather likely for forest species (Zaplata et al., 2010). It is likely that the C. canadensis seeds entered the initial substrates during transport to the construction site, and even more likely, at the site itself, when the catchment was still under construction. The latter is indicated by a trend that the first constructed part of the catchment area contained more seeds than the younger one. In conclusion, we consider it as an unintended seed rain just before initial soil sampling. From an ecological point of view the occurrence there is a new seed bank or camouflaged seed rain. Thus, the initially detected seeds in the soil substrate represent the ubiquity of airborne C.

canadensis seeds during the seed shedding.

Due to the time of the final catchment construction work during the seed shedding in 2005, seed input onto the primary substrate before sampling was most relevant: Although the top 2 cm were rejected, the soil seed bank apparently contained the seed rain which occurred shortly before sampling. As in 2005 there were no C. canadensis plants growing in the catchment, all seeds of C. canadensis in the catchment and at its edges originated from the surroundings in 2005 and before seed shedding in 2006. At that time the catchment was entirely a sink for the species.

Source region

The hallmark of the first generation of C. canadensis plants was extensive spatial auto-correlation in terms of cover referring to the 5% level of significance. This can be interpreted as the deficiency of environmental filtering (mediated by biotic interactions) at the beginning of succession, over the course of which biological filtering mechanisms will gradually gain in importance ( [Baasch et al., 2009] and [Grime, 1973]). In others words, inner ecosystem mechanisms including self-regulation will become increasingly important later on (Titlyanova, 1982). At the high 1% level of significance, the spatial structure of C. canadensis corresponded with the main parts of the Chicken Creek catchment (four major sections of the backslope area; see Gerwin et al., 2009). The reason for such anisotropy is substantiated by the directional geophysical phenomena (Legendre and Fortin, 1989): the Chicken Creek catchment environment is spatially structured by the north south alignment axis, which supported interconnections at larger distances, and of the eastern and western halves, reflecting slightly different substrates used for construction in the first and second phases of catchment construction, respectively. Thus, at the very early stage we detected already a certain pattern formation of the dominant plant species whereby the pattern matches the physical conditions. Simultaneous dominance in almost all experimental plots with nearly the same cover degrees was also reported for C. canadensis and other dynamic species (i.e. those with r-strategies) from a succession study on an abandoned field in central Germany (Schmidt, 1981). Also from the lignite mining region of Lower Lusatia, where our catchment is located, it was noted that C.

canadensis occurred in high steadiness in all early-successional vegetation types except sandy areas
with almost no vegetation cover (Felinks, 2000). In contrast to our study, in an open-cast mining area in western Germany C. canadensis reached a maximum frequency distribution only in the fifth year of vegetation development (Wolf, 1985), and this was reached during the fourth to seventh vegetation periods on cleared loess hills in South-West Germany (Fischer, 1982). Wolf (1985) reported that in some permanent plots C. canadensis was very abundant though its overall occurrence varied widely. The latter study showed at the beginning of succession only few individuals, then during the abovementioned period high steadiness and cover values, and later rapid decrease. Thus these studies show, if C. canadensis occurs in progressive succession, then at an early stage, but not as inevitably early as in our study, neither it does occur that spaciously and regularly per se. A comprehensive substantiation of the potential and relevance of C. canadensis for rapid colonization of large areas is up to the present study, where virtually neither localized dispersal nor dispersal limitation occurred and therefore in interplay with lacking biological filtering mechanisms an uniformly distributed vegetation could unfold promptly (instead of a first aggregated, within-catchment dispersal kernel). Far from being dispersal limited was also the first proven seed outflow. During the first C.

canadensis seed shed within the catchment, local patterns of seed rain occurred. Most seeds were
trapped at sites where the prevalent wind had passed over the whole catchment indicating a seed source function. Other initial colonizer plant species did not reach by far the dispersal and dispersion levels of C. canadensis. Ch. album, the second important pioneer species in the Chicken Creek catchment area, was irrelevant in seed rain and the individual number in the respective year of maximum abundance (Ch. album: year one, C. canadensisyear two of the succession). Given the dwarf forms, the reproduction capacity of Ch. album was restricted (Silvertown and Lovett-Doust,

1993). The initially frequent Cirsium arvense, known as a good disperser (Donald, 1994) and not afflicted by dwarfism, did not keep pace with C. canadensis. Medicago varia, whose neighbour abundance was very high, up till now has not at all established itself. The early dominance of C. canadensis embodies its unique propagation and dispersal characteristics combined with outstanding growth performance in the initial substrate. Given such extraordinary features, forC. canadensis and supposedly other plant species with vast dispersal potential as well, a bottle-neck situation of colonization of suitable sites during primary succession is hardly imaginable (in contrast to the general statement of Poschlod et al., 1996) as long as a source population has certain strength.

Classification of the study and outlook

This study focuses on the population dynamics of a single prominent plant species in an emerging ecosystem. It shows how a very successful r-selected species can enter a catchment area. The local development of this single taxon shows that the whole system had already changed to source function within one year after arrival of the species. Our findings supply quantitative data for such a process, whereas prior to this study most information on seed dispersal deals with qualitative descriptions of transport mechanisms (e.g. Donald, 1994). Beyond this realization in basic research, this empirical case study provides a foundation for (modelling) studies: once overall consequences on a landscape unit scale are demonstrated, henceforth the potential of a rapid change in sink source dynamics is to be derived simply from single propagation and dispersal attributes of a certain plant species.

Acknowledgments
This study is part of the Transregional Collaborative Research Centre 38 (SFB/TRR 38) which is financially supported by the Deutsche Forschungsgemeinschaft (DFG, Bonn) and the Brandenburg Ministry of Science, Research and Culture (MWFK, Potsdam). The authors thank the working group Z1 (monitoring) members of the SFB/TRR 38 who helped us to perform this study and the Vattenfall Europe Mining AG for providing the research site. Thanks to Hagen Fischer for supporting the statistical analyses, Anne Naeth, Harald Albrecht, and Johannes Kollmann for useful comments on the manuscript.

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View Record in Scopus | Cited By in Scopus (2) Corresponding author. Tel.: +49 8161 71 4732.

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