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Examining native and exotic species diversity in European riparian forests

Annik Schnitzlera, Brack W. Haleb,*, Esther M. Alsumc
a

Paul Verlaine, Metz, LBFE, Rue du Ge ne ral Delestraint, 57 070 Metz, France Universite Franklin College Switzerland, Via Ponte Tresa 29, 6924 Sorengo, Switzerland c Botany Department, 430 Lincoln Drive, University of Wisconsin-Madison, Madison, WI 53706, USA
b

A R T I C L E I N F O

A B S T R A C T

Article history: Received 6 February 2006 Received in revised form 10 March 2007 Accepted 6 April 2007

This study performs a meta-analysis of existing studies of European riparian forests to investigate which exotic species have successfully established in intact riparian forests and which characteristics of these forests correspond with successful establishment. We used analysis of covariance models to investigate the relationship between community species richness, percent exotic species, and several environmental variables. We found a total richness of 1380 species, of which 45 (3.3%) were exotic. Species-rich communities generally had the higher percentage of exotics, but were not signicantly related to latitude

Keywords: Riparian forests Exotic species Invasive species Disturbance Europe River oodplains

or environmental variables. Exotics, in contrast, were at generally higher levels at lower latitudes and were more abundant in large river plains and communities with intermediate levels of disturbance. These results suggest that future climate change and human actions that mimic intermediate levels of disturbance may further enhance the spread of exotic species. 2007 Published by Elsevier Ltd.

1.

Introduction

Riparian forests are complex and dynamic ecosystems that strongly depend on ood pulses for primary productivity and biodiversity (Carbiener, 1970; Ward et al., 1999). River systems naturally experience frequent, periodic ood disturbances that are responsible for high variations in texture and drainage at local scales. Floods also transport vegetative matter and seeds from upland and upstream sources, as well as across the oodplain, thereby encouraging propagation. The abundant supply of water and the high nutrient content of freshly deposited sediments combine with the typically plentiful light availability to promote successful germination and establishment (Walter, 1979; Sanchez-Perez et al., 1993, 1999; Schnitzler et al., 2003). These conditions also promote certain types of species, such as lianas, which are often less

competitive in upland forests (Schnitzler, 1995a; Allen et al., 2005). Moreover, these factors help explain why river systems provide opportunities for the establishment of exotic speciesnon-native species introduced intentionally or unintentionally by human action (Tickner et al., 2001). The expansion of exotics in river-oodplain systems characterized the 20th century as human alterations to these systems, such as channel modications, ow regulation, and oodplain drainage, intensied (Dynesius and Nilsson, 1994; the mont, Knutson and Klaas, 1998; Nolet and Rosell, 1998; Be 2000). Increasing organic pollution also favored certain types of exotics with high nutrient demand. The abandonment of traditional hay and grazing lands allowed the establishment and growth of dense stands of exotic clones (Pysek and Prach, 1993). Human activity further enhanced the expansion rates by increasing the dispersal of plant propagules into anthropo-

* Corresponding author: Tel.: +41 91 986 3650; fax: +41 91 994 4117. E-mail addresses: schnitz@univ-metz.fr (A. Schnitzler), bhale@fc.edu (B.W. Hale), emalsum@yahoo.com (E.M. Alsum). 0006-3207/$ - see front matter 2007 Published by Elsevier Ltd. doi:10.1016/j.biocon.2007.04.010

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genically disturbed riparian and oodplain areas. As many exotics are typically wetland species, they are generally well adapted to the riverine environment (Zedler and Kercher, 2004). Some of these exotics can then expand from river banks to other types of habitats, such as along roads and paths and on plantations. One well-known example is giant Japanese knotweed (Fallopia japonica), which, along with its hybrids, is threatening Western European river systems and adjacent areas (Tickner et al., 2001; Bailey and Schniztler, 2003). Exotics represent a particular class of species in the ora, which sometimes develop new morphologies and reproductive behaviors (unusual in their native habitats) to compete successfully in new habitats (Alpert et al., 2000). A result of phenotypic plasticity or competitive superior genotypes, exotics often exhibit larger growth forms: 34 m for some species such as Impatiens glandulifera or Fallopia spp. in Europe; Arundo donax has been reported to achieve heights greater than 9 m in coastal rivers in California (Dudley, 2005). They can occur at high vegetative densities (clonal perennials), which make them impenetrable for other vegetation. Other competitive characteristics include mycorrhizal transfer of carbon from native plants (Callaway et al., 2001), large seed sets typical of many annuals (Zedler and Kercher, 2004), hybridization, which can improve competitiveness (Abbot, 1992; Bailey, 1999), and resistance to local predators or pathogens (Zedler and Kercher, 2004). Many exotic species can also integrate into riparian forests through very active dispersal of rhizomes or stems deposited during oods. Exotic species do not affect all habitats equally. Some exotics achieve extensive, self-reproducing populations that can be classied as invasive. Invasive species are typically considered species that have become naturalized in the new habitat, that produce abundant offspring and that have the ability to spread to large areas (Richardson et al., 2000; Pysek et al., 2004). The potential of exotics to become invasive is generally high in altered rivers; the inuence of exotic plants on biodiversity within altered oodplains is well documented for open, disturbed areas, such as deforested riverbanks, and abandoned alluvial meadows (Planty-Tabacchi et al., 1996; Zavagno and DAuria, 2001). However, their effects on intact riparian forests have not been well documented. This paper performs a meta-analysis of existing studies of European riparian forests to investigate (1) which exotic species have successfully established in intact riparian forests and the abundance and range of these species, (2) which characteristics of these forests correspond with establishment, and (3) which exotic species are the most widespread. In so doing, we investigate how plant diversity varies across different riparian forest communities, which environmental factors are associated with diversity, and which are associated with exotic species.

data encompass 177 communities across numerous river systems in Europe from studies published between 1965 and 2006. The communities lie in one of four different climatic zones: temperate, meridional, submeridional, or boreal (Bohn et al., 2003). We classied each community into one of seven main habitat-types (Table 1), according to the Braun-Blanquet classication system commonly used in European ecology (Braun-Blanquet, 1965). The Braun-Blan ) that quet system uses a standardized sampling unit (releve allows for species richness comparisons across multiple sites. We included data if the forests were located in areas with intact riparian forests and excluded data from rare communities. In addition to community type and species composition, we recorded data on several environmental characteristics that could potentially inuence species diversity for all sites where data had been collected. These variables included: A. surface topography (sensu Sutley, 1982) 1 = level plains and at relief; 2 = irregular plains with relief 3090 m; 3 = plateaus with relief > 90 m; 4 = hill-studded plains, relief 90300 m; 5 = mountain-studded plains, relief > 300 m; 6 = hill lands, with steeper slopes, relief < 300 m; 7 = mountains, with steeper slope, relief 300500 m; 8 = mountains of great relief, relief > 1500 m, B. latitude, C. mean annual temperature (Sutley, 1982), D. mean annual rainfall (Sutley, 1982), and E. annual productivity. Primary productivity (g m2 year1) was calculated from the above measure of temperature using the equation in Lieth (1975).

2.2.

Analysis

2.
2.1.

Materials and methods

Data sources

We obtained data on European riparian forest communities from the phytosociological literature (see Appendix). The

For each community type, we calculated the average number of species, genera, and families present. We also calculated the average number of species present in three broad categories: woody, vine, and exotic species (as dened in Section 1). These three broad types of species can be used as indicators of the general features of riparian forests (cf. Section 1). Lastly, we summarized the site characteristics such as latitude, annual average temperature, precipitation, and primary productivity by community type. We assessed the distribution of the exotic species using the following schema. We considered the species to have a local range if it was located in 12 communities; it has an intermediate range if it was located in 34 communities, and can be considered widespread if it is found in 4+ communities. We also assessed the frequency of occurrence. An exotic species was considered scarce if it was present in less than 5% of all records; moderate if it was present in 620% of the records; and abundant if it was present in greater than 20% of the records. We analyzed the drivers of species richness and exotic species richness in the various communities using analysis of covariance (ANCOVA) models, which allowed us to incorporate both categorical and continuous variables. For the categorical variables, we used Willow Scrub as the reference for

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Table 1 Riparian forest types used in the analysis Community

Willow Scrub Willow-Poplar

Dominant tree species

Salix purpurea Salix alba Populus nigra Populus alba Quercus robur Fraxinus excelsior Ulmus laevis/minor Fraxinus excelsior Alnus glutinosa Alnus glutinosa Alnus incana

Successional stage
Pioneer Intermediate

Characteristics
Unstable river banks Silty-gravelly river banks

N of studies
29 24

White Poplar OakAshElm

Intermediate Later

Large rivers of the Mediterranean basin Large rivers

12 29

Ash-Alder

Later

Moist, heavy texture soils

41

Marsh Alder White Alder

Later Later

Soils waterlogged much of the year Boreal or alpine

24 18

community type and category 8 (mountainous terrain of great relief) as the reference for topography. Several variables did not possess normal distributions; consequently, we used a natural log transformation (or ln(x + 1), if variable included zero values) before running the analysis. Ln-transformed variables included exotic species richness and percent woody, vine, and exotic species. We performed stepwise variable testing to determine which variables to include in the nal models. We performed all statistical analyses in STATA 9.0, using an alpha value of 0.05 for statistical signicance.

With respect to climate and productivity, the White Poplar communities were generally the warmest and most productive, with an average annual temperature of 13.9 1.0 C and an annual production of 1745 89.8 g m2 year1. White Alder communities were the coldest and least productive, with an average annual temperature of 7.6 0.8 C and annual production of 1207 67.8 g m2 year1. The community averaging the greatest annual precipitation was OakElm Ash (733 32.1 mm), while Willow-Poplar and White Alder were the driest with an average annual precipitation of 643 40.4 and 644 20.3 mm, respectively.

3.
3.1.

Results
Average characteristics by community

3.2.

Community richness

The communities in this study ranged in latitude from 36.0 N to 66.4 N; most centered around the mid-latitudes (Table 2). The northernmost type was the White Alder community, which was generally limited to more boreal regions (mean = 53.7 1.6 N). In contrast, the White Poplar community was more typical of the southern areas in this study, averaging 42.4 1.2 N. Most community types were also generally found in more mountainous regions (modal topographic value = 7). The two exceptions were the OakElmAsh communities, which were most frequent on level plains (value = 1) and the Willow Scrub communities, which were most typically found on the irregular plains (value = 2).

Fig. 1 shows the mean number of species and geometric mean number exotic species per community type. The three latersuccessional communities, White Alder, OakElmAsh, and Ash-Alder, along with the intermediate Willow-Poplar typically had the much greater numbers of overall species than the early successional communities. Marsh Alder communities, albeit typically later-successional, tended to have lower values for all species richness akin to the earlier-successional types. Willow-Poplar communities had the highest mean number of exotics (2.7 0.2); White Poplar, OakElmAsh, and AshAlder also averaged 1+ exotic species. In contrast, Marsh Alder and White Alder communities averaged less than 0.30 exotic species per site.

Table 2 Mean characteristics by community (standard deviation in parentheses) Community

Willow Scrub Willow-Poplar OakAshElm Ash-Alder Marsh Alder White Alder White Poplar

Topographya
2 7 1 7 7 7 7

Latitude
50.0 49.0 47.5 46.4 48.5 53.7 42.4 (7.7) (7.9) (4.5) (5.9) (7.5) (6.7) (4.2)

Temperature (C)
9.5 9.3 11.1 11.8 10.8 7.6 13.9 (4.9) (5.0) (3.6) (3.1) (4.2) (3.3) (3.6)

Productivity (g m2 year1)
1373 1359 1499 1560 1480 1207 1745 (401) (402) (300) (275) (367) (288) (311)

Precipitation (mm)
696 643 733 728 717 644 726 (210) (198) (177) (139) (154) (86) (235)

a Value represents the mode of the data.

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100 90 80

Species Exotic species

10 9 8 7 6 5 4 3 2 1

Number of species

70 60 50 40 30 20 10 0 Willow Willow - White Scrub Poplar Poplar Marsh Alder Whit e Alder Ash- Oak-ElmAlder Ash

Fig. 1 Mean species and exotic species richness per community (NB: exotic is geometric mean). Bars represent 1 SE. Assessing the general composition of these communities, we found the White Poplar communities tended to have the greatest proportion of both woody and vine species, 36.9% and 10.6%, respectively (Fig. 2). The Marsh Alder community had the lowest proportion of woody species (20.7%), while the White Alder communities tended to have the smallest proportion of vine species (2.2%). Communities of higher woody species richness, such as the later-successional communities, also exhibited higher proportions of vines (4.97.5%). With respect to exotic species, Willow-Poplar and White Poplar communities had the highest mean percentage of exotic species (3.4 0.2 and 2.5 0.3 respectively), while Marsh Alder and White Alder showed the lowest mean percent exotics (0.27 0.1 and 0.32 0.1, respectively).

shrub and liana life-forms. Most of the exotics are thermophilous and light-demandingpioneer species from warmtemperate oodplains. Thirty-two percent are from the Asteraceae family. Many exotics found in this study were introduced intentionally either from North America (51%) or Asia (38%). The introductions generally took place between the 17th and the 19th century for various uses, including agriculture (e.g. Robinia pseudoacacia, Vitis riparia, Morus spp.), timber (e.g., Acer negundo), and horticulture (e.g. Solidago spp., F. japonica, Helianthus tuberosus, Amorpha fruticosa). The exotics include so-called archeophytes, such as Ficus carica, Morus nigra, M. alba (all of Asian origin), Vinca minor, which was introduced during the Roman times, and Juglans regia, which was spread by Neo zel and Me dail, 2003). lithic peoples (Que The distribution of exotics in the 177 communities recorded is highly unequal (Table 3). Twenty-six are present at low levels in very few communities; seven have a intermediate distribution; and twelve (27%) are abundant in a large range of habitats.

Number of exotic species

3.4.

Biodiversity models

3.3.

Exotic species richness and distribution

The studies summarized for this paper recorded 1380 species. Of these, 45 (3.3%) were exotic species (Table 3). The exotics belong to various life-forms: approximately 50% are grasses (polycarpic perennials, summer and autumn annuals), while the rest are phanerophytes, equally distributed among trees,

Our analysis of species richness revealed several factors associated with species richness (Fig. 3). The model was statistically signicant, but only explained approximately 24% of the variance within the data (Table 4). Signicant variables in this model included: community type, ln-transformed percent woody species, ln-transformed percent vine species, and ln-transformed percent exotic species. As Fig. 3 shows, within the variable community type, three showed a signicant positive relationship with species richness: Ash-Alder (t = 3.64, p = 0.00), OakElmAsh (t = 4.04, p = 0.00), and White Alder (t = 3.30, p = 0.00). The variable percent exotic species also has a signicant positive relationship with species richness (t = 3.78, p = 0.00), whereas the variables percent woody species (t = 4.47, p = 0.00) and percent vine species (t = 2.29, p = 0.02) showed signicant negative relationships with species richness. None of the environmental or geographic variables proved signicant. Our analysis of exotic species richness revealed several factors associated with species richness (Fig. 4). The model

40 35 30 % number of spp 25 20 15 10 5 0 White Poplar Oak-ElmAsh Ash-Alder

Woody Vine Exotic

WillowPoplar

Willow Scrub

White Alder

Marsh Alder

Fig. 2 Mean percent woody, vine, and exotic species per community type. Bars represent 1 SE.

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Table 3 Range and frequency of exotic species Species

Acer negundo Ailanthus altissima Amaranthus blittoides Amaranthus retroexus Ambrosia artemisifolia Amorpha fruticosa Artemisia verlotorum Aster novo-belgii Aster squamosus Aster tradescenti Bidens frondosa Chondrilla juncea Cotula coronopifolia Diospyros lotus Echynocystis lobata Erigeron annus Erigeron canadensis Fallopia japonica Ficus carica Fragaria indica Fraxinus pennsylvanica Galinsoga parviora

Growth type
Phanerophyte Phanerophyte Other Other Other Phanerophyte Other Other Other Other Other Other Phanerophyte Phanerophyte Other Other Other Other Phanerophyte Other Phanerophyte Other

Range
Local Local Local Intermediate Intermediate Intermediate Intermediate Local Local Intermediate Intermediate Local Local Local Local Local Widespread Local Widespread Local Local Local

Frequency
Abundant Moderate Moderate Moderate Moderate Abundant Moderate Scarce Moderate Moderate Abundant Moderate Moderate Scarce Scarce Scarce Abundant Scarce Abundant Scarce Moderate Scarce

Species
Gleditschia triacanthos Helianthus tuberosus Impatiens capensis I. glandulifera Impatiens parviora Juglans regia Lonicera japonica Morus alba Morus nigra Oenothera biennis Parthenocissus quinquefolia Paspalum distichum Phytolacca americana Prunus laurocerasus Robinia pseudoacacia Rudbeckia lasciniata Solanum lypopersicum Solidago gigantea Solidago graminifolia Trachycarpus fortunei Vinca minor

Growth type
Phanerophyte Other Other Other Other Phanerophyte Phanerophyte Phanerophyte Phanerophyte Other Phanerophyte Other Phanerophyte Phanerophyte Phanerophyte Other Other Other Other Phanerophyte Other

Range
Local Intermediate Local Intermediate Widespread Intermediate Local Intermediate Local Intermediate Intermediate Local Local Local Widespread Local Local Widespread Local Local Intermediate

Frequency
Scarce Abundant Scarce Abundant Abundant Moderate Scarce Moderate Scarce Moderate Abundant Scarce Scarce Scarce Abundant Scarce Scarce Abundant Scarce Moderate Abundant

Range (local: in 12 communities; intermediate: in 34 communities; widespread: in 4+ communities); frequency (scarce: present over < 5% of all records; moderate: 620%; abundant: >20%).

200 150 Coefficient 100 50 0 -50 -100

W hi te po W pl ill ar ow -p op M la ar r sh al de A r sh -a O ld ak er -e lm a W sh hi te al de r st an ic * y* * t W oo d ne Vi C on Ex ot
Coefficient

2 1.5 1 0.5 0 -0.5 -1

W h W ite ill p ow op M -po lar ar p s h la A a r O sh lde ak -a r - ld W elm er hi -a te sh al de r

Topography

Community type

Community Type

Fig. 3 Coefcient values for variables in the nal species richness model. Starred (*) variables were ln-transformed; bars represent 95% condence interval.

Fig. 4 Coefcients for variables in the nal exotic species model. Starred (*) variables were ln-transformed; bars represent 95% condence interval.

Table 4 Model statistics Species richness model

Independent variable N df F Statistic (p-value) r2 (Adjusted r2) Number of species 177 9 7.18 (0.00) 0.279 (0.240)

Exotic species model

Ln(number of exotic species + 1) 177 16 12.61 (0.00) 0.558 (0.513)

was statistically signicant and explained approximately 51% of the variance within the data (Table 4). Signicant variables in this model included: community type, topography, latitude, species richness, and ln-transformed percent woody species. As Fig. 4 shows, within the variable community type, three showed signicant relationships with exotic species richness: Willow-Poplar (t = 3.13, p = 0.00) with a positive relationship; Marsh Alder (t = 2.36, p = 0.02), and White Alder (t = 2.68, p = 0.01) with a negative relationship. Of the topography categories, only category 1 (at plains) showed a signicant (positive) relationship (t = 2.58, p = 0.01) with exotic species richness. The variable latitude has a signicant negative relationship

7 Sp La p titu r % ich de W ne o ss C ody on * st an t

Fl at

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with exotic species richness (t = 5.12, p = 0.00), while the variables species richness (t = 8.44, p = 0.00) and percent woody species (t = 2.04, p = 0.04) showed signicant positive relationships with exotic species richness. None of the other environmental or community variables proved signicant.

species coefcients in our species richness model. The benets of warmer temperatures and higher primary productivity may also inuence richness in other aspects of oodplain biota, such as birds (Buckton and Ormerod, 2002).

4.2.

Exotic species trends

4.
4.1.

Discussion and conclusions

Biodiversity trends
The importance of the dynamic nature of the river system in shaping the riparian community presents challenges and opportunities to potential exotic species. On one hand, the hydrologic regime altering from times of water stress to times of water (over)abundance requires a certain level of adaptation for any resident plant species, native or exotic (Naiman camps, 1997; Kozlowski, 2002). Nonetheless, the natuand De ral disturbances also create niche opportunities that exotics can exploit (Shea and Chesson, 2002). We discuss these two general inuences in the following paragraphs. The examination of the data showed that the distribution of exotics varies greatly in Europe: generally lower levels of exotics were found in areas with longer summer droughts, such as communities along intermittent rivers in the Mediterranean region, and in areas with colder (boreal) climates, such as the White Alder communities. Exotics are also nearly absent on the eastern edge of the temperate zone (up to the Ob River, east of the Urals), likely in part due to harsh winters (Taran, 1994). In order to establish and ourish, exotics require the availability of essential resources (Davis et al., 2000; Shea and Chesson, 2002), such as water and an appropriately long growing season. Thus, it should not come as a surprise that exotic species are less abundant in these areas. Nonetheless, harsher environmental conditions by no means preclude the invasion of exotic species, as suggested by data from North America. Dynesius et al. (2004) found a greater number of exotics in the boreal zone in North America compared to the boreal zone of Scandinavia. They attributed this to accidental and intentional introductions from human settlers as well as to the shorter history of planthuman co-evolution of North America. Species intentionally introduced by settlers beneted from an adaptation to anthropogenic disturbances. Thus, although harsher conditions certainly pose a challenge to the competitiveness of any species, those possessing appropriate life-history adaptations can survive and compete in any climate and particularly in areas where favorable conditions have been anthropogenically manipulated. A related trend is the signicance of latitude in our analysis of exotic species richness (Fig. 4), where exotic species richness decreases with increasing latitude. As growing seasons tend to decrease with increasing latitude, fewer species from warmer regions should be able to establish. Indeed, many exotics in European forests come from tropical or warm-temperate regions (Walther, 2003). Thus, in contrast to native riparian species, exotic species appear to respond vigorously to environmental gradients related to latitude. Grace (1987) related the similarity between the distribution of certain exotic species in the UK and winter low temperature isotherms to frost tolerance or to the lack of appropriate seasonal cycles. This nding has strong implications for increasing invasion potential with current global climate forecasts. As native ora respond to climate change by shifting their ranges northward (Svenning and Skov, 2006), more resources will become

Latitudinal gradients in species richness are well documented in the ecological literature (Grubb, 1987; Adams and Woodward, 1989). Nonetheless, results of our analysis did not reveal a direct relationship between latitude and species richness within the latitude range investigated here. This nding corroborates well with the lack of latitudinal gradients observed in studies of temperate and Mediterranean oodplain forests zel and Me dail, 2003; Schnitzler and Carbiener, 2006) (Que and of aquatic macrophytes (Willig et al., 2003). In oodplains, different habitat-types result from interactions among the hydrological regime, landscape heterogeneity, and availability of nutrient resources. Thus, diversity in river-oodplain systems responds more to local stresses, such as temporary anoxia or periodic ood-related disturbances, than to latituderelated factors (Dister, 1983; Deiller et al., 2001; Johnson, 2001). The lack of signicance for the environmental variables in our species richness model further supports this idea. The results do suggest that later successional communities generally possess higher levels of species diversity. These communities have typically developed greater structural diversity with time, which in turn supports a greater number of species (Pautou and Arens, 1994; Schnitzler, 1995b). This increase in the number of niches for species to occupy may also explain the positive relationship with percent exotics, as more niches would also be available for exotic species, as well as native species (Shea and Chesson, 2002). Our analyses also indicate lower levels of woody species in the communities that experience more extreme or more variable environmental conditions, such as those in swampy areas (Marsh Alder) or in colder regions (White Alder). In these harsher environments, it is no surprise that the number of woody species decreases. Perennial plants, such as woody plants, require special life-history characteristics to survive in more extreme environments (Grace, 1987; Kozlowski and Pallardy, 2002) In contrast annual grasses simply need nearby seed sources to persist. Areas experiencing frequent disturbances represent another form of extreme environmental conditions. Correspondingly, we found Willow Scrub communities, which typically experience the most frequent oods, also had lower levels of woody species. In contrast, those communities which represent less severe environmental conditions (i.e. warmer and wetter climates or in areas with intermediate levels of disturbance), the woody species richness increases sharply, particularly in the later-successional communities (e.g. AshOakElm, White Poplar, Ash-Alder). When woody species establish, they typically limit the amount of light reaching the ground layer and thus limit the number of species capable of persisting in the shade. One consequence would be a decrease in overall species richness, which could explain the negative woody

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available in the abandoned areas, which should thus enhance these habitats invasibility (Davis et al., 2000). The study by Svenning and Skov (2006) also found latitude to be the major predictor of decreases in habitat suitability for 36 European herbs. Our results indicate that certain topographies are more vulnerable to exotic species, such as communities in at plains. Rivers systems in at plains often tend to be the lower reaches of rivers that support a wide range of species and microhabitats (Ward et al., 2002). Since these areas typically represent the largest parts of the river system, the speciesarea relationship predicts higher species richness (and thus exotic species richness) simply due to their larger area. Larger oodplains will also have more habitats buffered from extreme high-ow and low-ow events of the river and thus provide habitat for exotic species which may not be adapted to the hydrologic regime of the river oodplain. A closer examination of the communities that had significant coefcient in the exotic species model may also provide insights into community and species characteristics that are suitable for establishment of exotic populations. Willow-Poplar communities appear most vulnerable to the integration of exotic species, according to our results. The canopy of these communities is typically more open and allows moderate levels of light to reach the forest oor, which can facilitate establishment (Schnitzler, 1994, 1995b). Further, exotics are also favored by the high frequency of ood disturbances in these communities, which often creates bare substrate and facilitates the spread of seeds and other propagules. The relatively short life span of these communities, generally on the order of a few decades (Schnitzler et al., 2005b), and open architecture favor the development of tall, nutrient-demanding hemicryptophytes with widespreading stolons or rhizomes. This effect appears in other habitats-types, such as grasslands along transportation corridors that also lack a closed canopy and possess altered disturbance regimes (Hansen and Clevenger, 2005). In contrast, Marsh Alder communities appear likely more resilient to invasion. Nonetheless, a high potential number of exotic amphiphytes have colonized from other continents, including Ludwigia peplo des, L. grandiora, Myriophyllum brassilensis, Lemna minuta, L. turionifera. The fact that these species have been able to colonize Marsh Alder communities despite seasonal inundations likely relates to shade and frost tolerance (many records come from the continental region of the temperate zone), varying rates of spread across species (Perrins et al., 1993; Pysek and Prach, 1993), or simply the age of the data (some references were published two, even three decades ago, before decline in water levels following drainages). As mentioned in the previous section, factors that favor overall species richness appear to favor exotics as well; this may explain the positive relationship between the two seen in our exotic species model.

4.3.

Exotic vs. invasive species

As discussed in Section 1, the presence of an exotic species in a community in itself does not necessarily represent a problem, except when it presents a potential for invasion. Invasive species typically have spread over large areas and at consider-

able distances from the parent plants (Pysek et al., 2004). Invasive species take advantage of the disturbance regimes of the riparian forests more readily than many of the native species (Planty-Tabacchi et al., 1995; Andersson et al., 2000). Some of them also integrate into the seed bank, allowing rapid invasion of gaps, such as A. donax, A. fruticosa in the submeridional zone and Impatiens noli-tangere and Solidago spp. in the temperate zone (Schnitzler et al., in preparation). Twelve of the exotics in Table 3 (Ailanthus altissima, A. negundo, R. pseudoacacia, Phytolacca americana, Solidago gigantea, I. glandulifera, I. parviora, Ambrosia artemisifolia, A. fruticosa, H. tuberosus, F. japonica, Erigeron canadensis) can be considered invasive species in Europe. They can form dense stands in various habitats, ranging from ruderal sites to deforested river banks. In intact oodplains, these tendencies appear to be less extensive, since some of species have been classied as local and scarce: Fallopia spp., A. negundo, S. gigantea remain limited to a few pioneer communities within which their abundance is often low, except locally (Schnitzler, 1995b). None of them attain cover of more than + or 1 (i.e., <5% of cover per record), probably due to shade and competition with native plants. Should these forests be harvested, these species could become highly invasive and impede natural forest regeneration. This is the case, for example, for F. japonica (Schnitzler et al., 2005a). Other plants may only be invasive in some regions. Two examples are the ornamental lianas, Ipomoae spp. and Sycios angulatos; which are locally invasive in the Mediterranean area in riparian forests near residential areas. The residential area serve as source populations for the camps and forests after the forests have been destroyed (De camps, 2002). De The classication of plants as invasive may depend on subjective motives (Simberloff, 2003). This may be the case for the archeophyte F. carica. It is widely represented and found in many records; thus, it is classied as widespread and abundant. However, this species is not considered invasive, probably because it has been present in riparian habitats for millennia. Understanding the transition from exotic to invasive (exotic species) is important, particularly given that many exotics species have only recently been established in the new habitat. Exotic species may simply represent additions to the local ora, while invasive exotic species can actually exclude local native species, such as Phalaris arundinacea in the Northwestern US (Perkins and Wilson, 2005). The eventual classication as an invasive species derives both from the suitability of their life-history traits for the local environment (i.e. their invasiveness) and the intensity of human use in the local environment, which increases invasibility of riparian habitats. At least ve factors have been invoked to explain invasibility: evolutionary history, community structure, propagule pressure, disturbance, and stress (Alpert et al., 2000). For example, two American vines, Parthenocissus quinquefolia and V. riparia, now show signs of future invasion in riparian habitats, as they appear to integrate easily in riparian forests, in contrast to upland environments (Schnitzler, unpublished data). P. quinquefolia is an exclusive liana, linked to gaps and forest edges, and has been widely planted in gardens as an ornamental plant. It has already become locally invasive on oodplain islands, such as the Loire. Vitis x riparia, a stock

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Table 5 Species richness in Lower Mississippi comparison systems (Klimas, 1988) System
Lower Mississippi Willow Cottonwood Sycamore-Maple-Elm Sugarberry-Elm-Ash Sweetgum-Oaks Cypress-Tupelo Overcup oak-Hickory

of the riparian forests (Denk et al., 2001; Schnitzler et al., 2005b). % Exotic
6.2 6.3 4.6 3.9 5.1 2.2 3.4 0.0

Totala
227 64 109 155 158 89 58 26

% Woody
34.4 26.6 35.8 38.7 34.8 57.3 51.7 61.5

4.5.

Implications for exotic species management

a Due to differing sampling efforts, richness values are not comparable.

of the European cultivated vine, has expanded after abandonment of vineyards during the 20th century; in particular its abundance is increasing in particularly western Mediterranean oodplains. Other exotic species that show potential invasiveness include Trachycarpus fortunei, which has been invading the early successional communities of southern France, and Gledistchia triacanthos, which has become very abundant on the sand bars of a tributary of the Rho ne, probably because of local plantations (Tabbachi, personal communication). One question offered by the results of this analysis is how applicable are the ndings to other systems outside of Europe. The lack of use of the Braun-Blanquet system in other regions of the world make the methodology of this study more difcult to apply to other regions; nonetheless a comparison with data from a study of the Lower Mississippi River by Klimas (1988) suggests some similarities (Table 5). With respect to community composition, woody species appear to exist at greater percentages in the later-successional communities (Overcup oak-Hickory, Sweetgum-Oaks, and Cypress-Tupelo). Further, higher levels of exotics are also present in the counterparts to the Willow-Poplar community, the Willow and the Cottonwood communities of the Lower Mississippi Lower levels are seen in the frequently inundated Cypress-Tupelo community, a counterpart to the Marsh Alder community.

Our results suggest that certain types of communities in temperate riparian forests are at potentially greater risk for exotic species, indicating that management efforts should focus particularly on such sites. Communities that tend to be more open (e.g. Willow-Poplar) are one example. This has large implications for anthropogenic disturbances that create similar disturbances, such as timber harvests, and underscores the need for invasive species management in such systems. Lastly, the importance of latitude in the exotic species model and its positive relationship with exotic species richness suggest that the spread of exotics should be enhanced under currently predicted future climate scenarios. Global climate change will also increase human pressure on river systems by altering hydrologic regimes and thus further encourage the spread of many invasive species (Pearce, 2006). Although exotic species do not necessarily pose an immediate threat to the native system, they certainly possess the potential to alter system sustainability in the long-term, particularly if riparian habitats become further altered by other human impacts. Exotic plants, when invasive, are one of the leading direct causes of biodiversity loss; exotic dominance could also be the indirect consequence of habitat modication driving native species loss (Didham et al., 2005). Therefore, it behooves society to promote greater scientic rigor in assessing the mechanistic causes of invasibility in riparian habitats and to initiate and strengthen conservation and management efforts to prevent the spread of additional exotic species.

Acknowledgements
We would like to thank Thierry Cornier for his bibliographic assistance and four anonymous reviewers for their helpful comments which have improved the quality of this manuscript. All remaining errors are our own.

4.4.

Caveats

A study such as this has its limitations. As a meta-analysis, it relies on pre-existing data. These data, although collected within a standardized system, span across several decades (19652006), which limits the strength of the comparisons. Further, as some studies indicate that increasing river alterations in the past 50 years have enhanced the possibilities for ood intolerant exotics to penetrate in oodplains (Dynesius the mont, 2000), it and Nilsson, 1994; De Waal et al., 1995; Be would be valuable to this incorporate variables on river modications present at the study sites. Unfortunately adequate data were not available in the studies accessed for our analysis to investigate this important question, however we hope to do so in the future. Lastly, many areas that could inform us about exotic species dynamics in rivers systems no longer exist in Europe. Human activity in areas along the Mediterranean or around the Black or Caspian Seas has destroyed most

Appendix.

Studies used in meta-analysis

zel, P., 1994. Place, role and historic value Barbero, M., Que of laurifolious elements in western Mediterranean preforest and forest vegetation. Annali di Botanica 12, 80133. Bensettiti, F., 1992. Approche phytosociologique des aulna gion dEl Kala (Alge rie). Documents Phytosocioies de la re logiques 14, 231240. Bensettiti, F., Lacoste A., 1999. Les ripisylves du nord de rie: essai de synthe ` se synsyste matique a ` le chelle lAlge diterrane e occidentale. Oecologia Mediterranea de la Me 25 (1), 1339. Bergmeier, E., 1990. Wa sche des Niederen lder und Gebu Olymp (Kato Olimbos, NO Thessalien). Ein Beitrag zur systematischen und orographischen Vegetationsgliederung Griechenlands. Phytocoenologia 18 (2/3), 161342.

154

B I O L O G I C A L C O N S E RVAT I O N

1 3 8 ( 2 0 0 7 ) 1 4 6 1 5 6

the mont, J., 2000. Les grands euves. Armand Colin, Be Paris. Birse, E.L., 1982. The main types of woodland in north Scotland. Phytocoenologia 10, 955. Brullo, S., Spampinato, G., 1990. La vegetazione dei corsi dacqua della Sicilia. Bolletino della Accademia Gioenia di Scienze Naturali 23, (336), 119252. Bulokhov A.D., Solomeshch A.I., 2003. Syntaxonomy of forests of Russian South Nechernosemic. Brjanks BGU. ` le tude de lassociation StelCoste, I., 1984. Contribution a lario nemori-Alnetum glutinosae (Ka stner 38) Lohm 57 en Roumanie. In: Cramer, J. (Ed.), Colloques phytosociologi ge tation des fore ques La ve ts alluviales, Strasbourg, Vaduz, 1980, pp. 6674. Denk, T., Frotzler, N, Navitashvili, N., 2001. Vegetational patterns and distribution of relict taxa in humid temperate forests and wetlands of Georgia (Transcaucasia). Biological Journal of the Linnean Society 72, 287332. Dierschke, H., 1975. Die Schwarzerlen (Alnus glutinosa) Uferwa lder Korsikas. Phytocoenologia 2, 229243. Dubyna, D.V., Neuhauslova, Z., Dzinba, T.P., Sheyag Sosonko, V.R., 2004. Classication and prodrome of vegetation of reservoirs, oodlands and arenas of the northern Black Sea region. Phytosociocente 23, 153199. Dumont, J.M., 1984. Les fore ts de lAlnion glutinosae et de gion du plateau des Tailles (Haute lAlno-Padion de la re Ardenne belge). In: Cramer, J. (Ed.), Colloques phytosoco ge tation des fore logiques, la ve ts alluviales, Strasbourg, pp. 259297. ` res observations sur Estrade J., Rameau J.C., 1984. Premie les fore ts riveraines des Vosges et du Morvan. In: Cramer, ge tation des J. (Ed.), Colloques phytosocologiques, la ve fore ts alluviales, Strasbourg, pp. 411429. Francalancia, C., Orsomando, E., 1984. Les fore ts riveraines de la Valnerina (Italie centrale). In: Cramer, J. (Ed.), Collo ge tation des fore ques phytosocologiques, la ve ts alluviales, Strasbourg, pp. 155160. Gafta, D., Pedrotti, F., 1995. Tipicazione di due nuove associazioni forestali ripariali per la penisola italiana. Documents Phytosociologiques 15, 413415. ge tation de la Corse. Editions des Gamisans, J., 1991. La ve `ve, conservatoires et jardin botaniques. Ville de Gene Edisud. Gehu, J.M., Franck J., 1984. Observations sur les saulaies e de la Loire, des sources a ` lembouriveraines de la valle chure. In: Cramer, J. (Ed.), Colloques phytosocologiques, ge tation des fore la ve ts alluviales, Strasbourg, pp. 305 323. Gellini, R., Pedrotti, F. Venanzoni, P., 1986. Le associazioni forestali ripariali e palustri della selva di San Rossore (Pisa). Documents Phytosociologiques 12, 2741. Girault, D., Timbal, J., 1984. Les fore ts alluviales de la Woevre. In: Cramer, J. (Ed.), Colloques phytosocologiques, la ge tation des fore ve ts alluviales, Strasbourg, pp. 393405. Golub, V., 1997. Communities of the Medicagini caeruleae Salicenion albae suball nova on the territory of the Lower Volga valley. Ukrainian Phytosociology 1, 6166. Golub, V.B., Kuzmina, E.G., 1997. The communities of Querco-Fagetea Br.Bl of the Lower Volga Valley. Peddes Repertorium 108, 205218.

Golub, V.B., Kuzmina, E.G., Golub, V.B., Kuzmina, E.G., Yuritsyna, N.A., 1998. Communities with dominance of Tamarix ramossisima: the lower Volga valley. Ukrainian Phytosociological Collection 1, 5260. Horvat, I., Glavac, V., Ellenberg H. 1974. Vegetation Su dosteuropas. Geobotanica selecta Band IV, Fischer Verlag Stuttgart. Jelem, H., 1974. Die Auenwa lder der Donau in Osterreich. Mitteilungen der Forstlichen Bundesversuchsanstalt 109, 1287. Johansson, M.E., Nilsson, C., Nilsson, E. 1996. Do rivers function as corridors for plant dispersal? Journal of Vegetation Science 7, 593598. Kalliola, R., Puhakka, M. 1988. River dynamics and vegetation mosaicism: a case study of the river Kamajohka, northernmost Finland. Journal of Biogeography 15, 703 719. Killiand-Lund, J., 1981. Die Waldgesellschaften SO Norwegen. Phytocoenologia 9, 53250. Krausch, H.D. 1965. Vegetationskundliche Beobachtungen im Donau Delta; Limnologica, 3, 271313. Lapraz, G., 1966 Recherches phytosociologiques en Cata ge tation primitive des rivie ` res logne Les vestiges de la ve tage bioclimatique me diterranen et des ruisseaux dans le sub-humide. Collectanea Botanica VI, IV, 548557. ge tation de le tage me diterrane en Loisel, R., 1976. La ve dans le sud est continental franc ais. Thesis, University of Aix-Marseille III. Manzi, A., 1988 Relitto di bosco ripariale lungo il corse planiziare del ume Sangro (Italia centarle). Documents phytosociologiques 11, 562571. Margl, H., 1973. Panzengesellschaften und ihre standortgebundene Verbreitung in teilweise abgeda mmten Donauauen. (Untere Lobau). Verhandlungen der ZoologischBotanisch Gesellschaft in Wien 113, 551. Merloni, N., Piccoli, F., 2001. La vegetazione del complesso Punte Alberete et valle Mandriole (parco regionale del delta del Po, Italia). Braun-Blanquetia, 29, Camerino. s phytosoMolinier, R., Tallon, G., 1970. Prodrome des unite es en Camargue. Bulletin du Museum ciologiques observe National dHistoire Naturelle de Marseille 30, 5110. Natkhutsrishwili, G., 1999. The vegetation of Georgia (Caucasus). Braun-Blanquetia 15, 717. ` gre, R., 1984. Dynamique ve ge tal de lOuve ` ze. In: CraNe ge tation mer, J. (Ed.), Colloques phytosocologiques, la ve des fore ts alluviales, Strasbourg, pp. 171190. Oberdorfer, E., 1992. Suddeutsche Panzengesellschaften. Part IV: Wa rsche, vol. 2, G. Fisher-Verlag, lder und Gebu Jena, Stuttgart, New York. Passarge, H., 1985. Phanerophyten Vegetation der ma rkischen Oderaue. Phytocoenologia 13, 505603. Piccoli, F., Gerdol, R., 1984. Typology and dynamics of wood in the Po plain (N-Italy): the Bosco della Mesola. In: Cra ge tation mer, J. (Ed.), Colloques phytosocologiques, la ve des fore ts alluviales, Strasbourg, pp. 161170. Prieditis, N., 1997. Alnus glutinosa dominated wetland forests of the Baltic Region: community structure, syntaxonomy and conservation. Plant Ecology 129, 4994. Rameau, J., Schmitt, C., 1984. Les fore ts alluviales de la plaine de la Sao ne. In: Cramer, J. (Ed.), Colloques phytoso-

B I O L O G I C A L C O N S E RVAT I O N

1 3 8 ( 2 0 0 7 ) 1 4 6 1 5 6

155

ge tation des fore cologiques, la ve ts alluviales, Strasbourg, pp. 93114. Rivas-Martinez, S., 1968. Contribucion al estudio geobotanico de los bosque araneses (Pirineo ilerdense). P. Inst. Biol. Aplic. 45, 81105. Rivas-Martinez, S., Lousa, M., Dias, T., Fernandez, F., Costa, J., 1990. La vegetacion del sur de Portugal (Sado, Alentejo y Algarve). Itinera Geobotanica 3, 5126. Rivas-Martinez, S., Diaz, T., Fernandez-Gonzalez, F., Izco, J., Loidi, J., Lousa, M., Penas, A., 2001. Syntaxonomical checklist of vascular plant communities of Spain and Portugal to association level. Itinera Geobotanica 14, 5341. Rodwell, J.S., Pigott, C.D., Ratcliffe, D.A., 1991. British Plant Communities. Cambridge University Press, Cambridge, New York. e du Sartori, F., 1984. Les fore ts alluviales de la basse valle Tessin (Italie du nord). In: Cramer, J. (Ed.), Colloques phyt ge tation des fore osocologiques, la ve ts alluviales, Strasbourg, pp. 201216. Sartori, F., Bracco, F., 1996. Present vegetation of the Po plain in Lombardy. Alliona 34, 112135. Schnelle, E., 1981. Die Panzengeselleschaften des Naturschutzgebietes Steckby-Lo dderitzer Forst. Hercynia 18, 387398. cologie Schnitzler, A., 1988. Typologie phytosociologique, e omoret dynamique des fore ts alluviales du complexe ge nan (plaine rhe nane centrale dAlsace). phologique ello-rhe Thesis, University Louis Pasteur, Strasbourg. compare e Schnitzler, A., Carbiener, R., 2006. Biodiversite ` ze, Ouve ` ze, des ripisylves du Rho ne et de ses afuents (Ce gion me diterrane enne. Fore diterDurance) dans la re ts Me ennes 27, 4358. rane Schwabe, A., 1985. Monographie Alnus incana reicher Waldgesellschaften in Europa. Variabilita t und Ahnlichkeiten inner azonal verbreiteten Gesellschaftsgruppe. Phytocoenologia 13 (2), 197302. Silc, U., 2003. Vegetation of the class Salicetea purpureae in Dolenjska (SE Slovenia). Phytocoenologia 40 (2), 227. Skogen, A., 1972. The Hippophae rhamnoides alluvial forest at Leinora, Central Norway. A phytosociological and ecological study. Kongelige Norske Videnskabers Selskabs Skrifter 4, 1115. Taran, G.S., 1994. Willow forests of Ob oodplain between mouths of Tym and Vakh rivers (Salicetea purpureae Moor 1958). Botanical investigations of Siberia and Kazakhstan. Transactions of V.V. Sapozhnikov Herbarium Issue 5 Barnaul, 4756. ` se, P., 1993. Les groupements ligneux riverains de la Vare Basse Durance (Provence). In: Cramer, J. (Ed.), Colloques ge tation des fore phytosocologiques, la ve ts alluviales, Strasbourg, pp. 565593.
R E F E R E N C E S

Abbot, R.J., 1992. Plant invasions, interspecic hybridization, and the evolution of new plant taxa. Tree 7, 401403. Adams, J.M., Woodward, F.I., 1989. Patterns in tree species richness as a test of the glacial extinction hypothesis. Nature 339, 699701.

Allen, B.P., Sharitz, R.R., Goebel, P.C., 2005. Twelve years post-hurricane liana dynamics in an old-growth southeastern oodplain forest. Forest Ecology and Management 218, 259269. Alpert, P., Bone, E., Holzapfel, C., 2000. Invasiveness, invasibility and the role of environmental stress in the spread of nonnative plants. Perspectives in Plant Ecology, Evolution and Systematics 3, 5266. Andersson, E., Nilsson, C., Johansson, M.E., 2000. Effects of river fragmentation on plant dispersal and riparian ora. Regulated Rivers-Research & Management 16, 8389. Bailey, J., 1999. The Japanese knotweed invasion of Europe: the potential for further evolution in non-native regions. In: Vano, E.E.A. (Ed.), Biologica Invasions of Ecosystems: Pests and Benecial Organisms. NIAES, Tsukuba. e du Japon, une Bailey, J., Schniztler, A., 2003. La renoue dangereuse conque te de lOuest. La Recherche 5, 5055. the mont, J., 2000. Les grands euves. Armand Colin, Paris. Be Bohn, U., Gollub, G., Hettwer, C., Neuha uslova, Z., Schlu ter, H., Weber, H., 2003. Map of the Natural Vegetation of Europe. Federal Agency for Nature Conservation, Bonn. Braun-Blanquet, J., 1965. Plant Sociology. Hafner Publishing Company, New York. Buckton, S.T., Ormerod, J., 2002. Global patterns of diversity among the specialist birds of riverine landscapes. Freshwater Biology 47, 695709. Callaway, R.M., Newingham, B., Zabinski, C.A., Mahall, B.E., 2001. Compensatory growth and competitive ability of an invasive weed are enhanced by soil fungi and native neighbours. Ecology Letters 4, 429433. Carbiener, R., 1970. Un exemple de type forestier exceptionnel pour lEurope occidentale: la fore t du lit majeur du Rhin au rhe nan (Fraxino-Ulmetum, Oberd. 53). Inte re niveau du fosse t cologique et bioge ographique. Comparaison a ` dautres fore e ts thermophiles. Vegetatio 20, 97148. Davis, M.A., Grime, J.P., Thompson, K., 2000. Fluctuating resources in plant communities: a general theory of invasibility. Journal of Ecology 88, 528534. De Waal, L.C., Large, A.R.G., Wade, P.M., 1995. River and oodplain rehabilitation in western Europe: opportunities and constraints. Archiv fu r Hydrobiologie 101 (suppl.), 679693. camps, H., De camps, O., 2002. Ripisylves me diterrane ennes. De Tour du Valat, Arles, France. Deiller, A.F., Walter, J.M.N., Tremolieres, M., 2001. Effects of ood interruption on species richness, diversity and oristic composition of woody regeneration in the upper Rhine alluvial hardwood forest. Regulated Rivers-Research & Management 17, 393405. Denk, T., Frotzler, N., Navitashvili, N., 2001. Vegetational patterns and distribution of relict taxa in humid temperate forests and wetlands of Georgia (Transcaucasia). Biological Journal of the Linnean Society 72, 287332. Didham, R., Tylianakis, J., Hutchison, M., Ewers, R., 2005. Are invasive species the drivers of ecological change? Trends in Ecology & Evolution 20, 470474. Dister, E., 1983. Zur Hochwassertoleranz von Auenwaldba umen an lehmigen Standorten. Verhandlungen der Gesellschaft fur kologie X, 325336. O Dudley, T., 2005. Arundo donax. In: California Invasive Plant Council. Dynesius, M., Nilsson, C., 1994. Fragmentation and ow regulation in the Northern third of the world. Science 266. Dynesius, M., Jansson, R., Johansson, M.E., Nilsson, C., 2004. Intercontinental similarities in riparian-plant diversity and sensitivity to river regulation. Ecological Applications 14, 173191. Grace, J., 1987. Climate tolerance and the distribution of plants. New Phytologist 106, 113130.

156

B I O L O G I C A L C O N S E RVAT I O N

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Grubb, P., 1987. The Organization of Communities Past and Present. Blackwell, Oxford. Hansen, M., Clevenger, A., 2005. The inuence of disturbance and habitat on the presence of non-native plant species along transport corridors. Biological Conservation 125, 249259. Johnson, W.C., 2001. Tree recruitment and survival in rivers: inuence of hydrological processes. Hydrological Processes 14, 30513074. Klimas, C.V., 1988. Forest vegetation of the leveed oodplains of the Lower Mississippi River. Lower Mississippi River Environmental Program Report 11, Mississippi River Commission, Vicksburg, MS. Knutson, M., Klaas, E., 1998. Floodplain forest losses and changes in forest community composition and structure in the Upper Mississippi River: a wildlife habitat at risk. Natural Areas Journal 18, 138150. Kozlowski, T.T., 2002. Physiologicalecological impacts of ooding on riparian forest ecosystems. Wetlands 22, 550561. Kozlowski, T.T., Pallardy, S.G., 2002. Acclimation and adaptive responses of woody plants to environmental stresses. Botanical Review 68, 270334. Lieth, H., 1975. Modelling the primary productivity of the world. In: Lieth, H., Whittaker, R.H. (Eds.), Primary Productivity of the Biosphere. Springer-Verlag, New York, pp. 237264. camps, H., 1997. The ecology of interfaces: Naiman, R.J., De riparian zones. Annual Review of Ecology and Systematics 28, 621658. Nolet, B.A., Rosell, F., 1998. Comeback of the beaver Castor ber: an overview of old and new conservation problems. Biological Conservation 83, 165173. Pautou, G., Arens, M.F., 1994. Theoretical habitat templets, species traits, and species richness oodplain vegetation in the upper Rhone river. Freshwater Biology 31, 507522. Pearce, P., 2006. Quand meurent les grands euves. Enque te sur la vy, Paris. crise mondiale de leau. Calman-Le Perkins, T., Wilson, M., 2005. The impacts of Phalaris arundinacea (reed canarygrass) invasion on wetland plant richness in the Oregon Coast Range, USA depend on beavers. Biological Conservation 124, 291295. Perrins, J., Fitter, A., Williamson, M., 1993. Population biology and rates of invasion of three introduced Impatiens species in the British Isles. Journal of Biogeography 20, 3344. Planty-Tabacchi, A.M., Tabacchi, E., Naiman, R.J., DeFerrari, C., camps, H., 1995. Invasibilty of species-rich communities in De riparian zones. Conservation Biology 10, 598607. Planty-Tabacchi, A.M., Tabacchi, E., Naiman, R.J., Deferrari, C., Decamps, H., 1996. Invasibility of species rich communities in riparian zones. Conservation Biology 10, 598607. Pysek, P., Prach, K., 1993. Plant invasions and the role of riparian habitats: a comparison of four species alien to Central Europe. Journal of Biogeography 20, 413420. nek, M., Webster, G.L., Pysek, P., Richardson, D.M., Rejma Williamson, M., Kirschner, J., 2004. Alien plants in checklists and oras: towards better communication between taxonomists and ecologists. Taxon 53, 131143. zel, P., Me dail, F., 2003. Ecologie et bioge ographie des fore Que ts du diterrane en. Elsevier, Paris. bassin me Richardson, D.M., Pysek, P., Rejmanek, M., Barbour, M.G., Panetta, F.D., West, C.J., 2000. Naturalization and invasion of alien plants: concepts and denitions. Diversity and Distributions 6, 93107. molie ` res, M., Schnitzler, A., Badre, B., Sanchez-Perez, J.M., Tre Carbiener, R., 1993. Nutrient content in alluvial soils submitted to ooding in the Rhine alluvial deciduous forest. Acta Oecologica 14, 371387. Sanchez-Perez, J.M., Tremolieres, M., Takatert, N., Ackerer, P., Eichhorn, A., Maire, G., 1999. Quantication of nitrate removal by a ooded alluvial zone in the Ill oodplain (Eastern France). Hydrobiologia 410, 185193.

Schnitzler, A., 1994. European alluvial hardwood forests of large oodplains. Journal of Biogeography 21, 604623. Schnitzler, A., 1995a. Community ecology of arboreal lianas in gallery forests of the Rhine valley, France. Acta Oecologia 16, 219236. Schnitzler, A., 1995b. Successional status of trees in gallery forest along the river Rhine. Journal of Vegetation Science 6, 479486. compare e des Schnitzler, A., Carbiener, R., 2006. Biodiversite ` ze, Ouve ` ze, ripisylves du Rho ne et de ses afuents (Ce gion me diterrane enne. Foret Durance) dans la re dite rane enne XXVII, 4358. me Schnitzler, A., Gafta, D., Cornier, T., 2003. Concepts architecturaux s e cosyste miques des ripisylves. In: Pie gay, H., et particularite Pautou, G., Rufnoni, C. (Eds.), Les fore ts riveraines des cours cologie, fonctions et gestions. Institut pour le deau, e veloppement forestier, pp. 3045. de Schnitzler, A., Bailey, J., Hansen, C., 2005a. Genotypic and phenophytic variation in a Fallopia bohemica population in north-Eastern France. In: Acts of Ecology and Management of Alien Plant Invasions, 8th International Conference EMAPI, Katowice, Poland. Schnitzler, A., Hale, B.W., Alsum, E., 2005b. Biodiversity of oodplain forests in Europe and eastern North America: a comparative study of the Rhine and Mississippi Valleys. Biodiversity and Conservation 14, 97117. rault, B., Aran, D., in preparation. Schnitzler, A., Lourenc o, S., He Soil seed bank and vegetation dynamics in oodplain forests of the upper Rhine valley. Shea, K., Chesson, P., 2002. Community ecology theory as a framework for biological invasions. Trends in Ecology & Evolution 17, 170176. Simberloff, D., 2003. Confronting introduced species: a form of xenophobia? Biological Invasions 5, 179192. Sutley, F., 1982. Britannica Atlas. Encyclopaedia Britannica, Chicago, IL. Svenning, J., Skov, F., 2006. Potential impact of climate change on the northern nemoral forest herb ora of Europe. Biodiversity and Conservation 15, 33413356. Taran, G.S., 1994. Willow forests of Ob oodplain between mouths of Tym and Vakh rivers (Salicetea purpureae Moor 1958). In: Botanical Investigations of Siberia and Kazakhstan. Transactions of V.V. Sapozhnikov Herbarium, pp. 4756. Tickner, D.P., Angold, P.G., Gurnell, A.M., Mountford, J.O., 2001. Riparian plant invasions: hydrogeomorphological control. Progress in Physical Geography 25, 2252. Walter, J.M., 1979. Architectural proles of ood forests in Alsace. In: Cramer, J. (Ed.), Struktur and Dynamik von Wa lder, Vaduz, pp. 187234. Walther, G., 2003. Plants in a warmer world. Perspectives in Plant Ecology, Evolution and Systematics 6, 169185. Ward, J.V., Tockner, K., Schiemer, F., 1999. Biodiversity of oodplain river ecosystems: ecotones and connectivity. Regulated Rivers: Research & Management 15, 125139. Ward, J.V., Tockner, K., Arscott, D.B., Claret, C., 2002. Riverine landscape diversity. Freshwater Biology 47, 517539. Willig, M.R., Kaufman, D.M., Stevens, R.D., 2003. Latitudinal gradients of biodiversity: pattern, process, scale, and synthesis. Annual Review of Ecology, Evolution, and Systematics 34, 273309. Zavagno, F., DAuria, G., 2001. Synecology and dynamics of Amorpha fruticosa communities in the Po plain (Italy). In: Brundu, G., Brock, J., Camarda, I., Child, L., Wade, M. (Eds.), Plant Invasions: Species Ecology and Ecosystem Management. Backhuys, Leiden, Netherlands, pp. 175182. Zedler, J.B., Kercher, S., 2004. Causes and consequences of invasive plants in wetlands: opportunities, opportunists, and outcomes. Critical Reviews in Plant Sciences 23, 431452.