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1007/s10531-010-9866-1 ORIGINAL PAPER
Encroachment of upland Mediterranean plant species in riparian ecosystems of southern Portugal
Maria J. Santos
Received: 8 September 2009 / Accepted: 14 May 2010 / Published online: 6 June 2010 Ó The Author(s) 2010. This article is published with open access at Springerlink.com
Abstract Riparian ecosystems have unique biodiversity, are highly sensitive to disturbance and anthropogenic inﬂuence. As world water resources become scarcer, scientists predict greater competition among species for water resources. Indeed, increased encroachment of upland plants into the riparian zone is already occurring, irreversibly changing riparian plant communities. Since semi-arid regions such as Mediterranean-type ecosystems are likely to follow this same trajectory, assessing the contributions of riparian versus upland (sclerophyllous) plants to community composition is important. A survey of seventy 2 km-long riparian transects on the Sado and Guadiana watersheds in southern Portugal assessed (1) the woody riparian plant community composition, (2) how much richness is due to strictly riparian plants versus sclerophyllous upland plants, and (3) which combinations of biotic and abiotic factors allow higher species richness in the strictly riparian, sclerophyllous, and overall plant communities. The survey detected 53 different woody plant species (28 endemic) across all communities. Riparian community richness was on average 16 species, seven of which were strictly riparian and the remainder being sclerophyllous, exotic species or fruit trees. Sclerophyllous plant species occurred consistently across sampling units (90% of transects). On average, 46% of the total woody plant community richness was due to strictly riparian plants and 28% was due to sclerophyllous plants. Community richness was positively affected by the area of shrubs in the riparian zone and by the absence of human activities and goats. Surrounding landscape pattern only affected the strictly riparian plant richness. These results suggest that natural and human-mediated disturbances in riparian ecosystems create gaps and clearings for which riparian and sclerophyllous plants compete. Establishment success seems to be related to the propagule pressure of the neighbouring landscape, its diversity and density, as well as the presence of herbivores. Preserving strictly riparian plants, removing exotic
M. J. Santos (&) ˆ Departamento de Biologia Animal, Faculdade de Ciencias, Centro de Biologia Ambiental, Universidade de Lisboa, Campo Grande, Bloco C2, 1749-016 Lisbon, Portugal e-mail: firstname.lastname@example.org M. J. Santos Department of Land, Air and Water Resources, University of California Davis, One Shields Avenue, Davis, CA 95616, USA
2005). 2000). 2009). spatially segregated from the strictly riparian plant patches. and promoting riparian values (recreation. especially in areas like the Iberian Peninsula which have become more arid. 2005). 1996. Schnitzler et al. dynamic and complex biophysical terrestrial ecosystems (Miller 2002. 2000. Despite their high biological value. These changes led to the loss of unique riparian species (Sabo et al. 2000) and likely resulted in woody plant encroachment in the riparian ecosystem (Huxman et al. Scarascia-Mugnozza et al. which intensiﬁes water loss through increased transpiration. aesthetics and the provision of ecosystem services) will aid the future conservation of the unique biodiversity of riparian ecosystems. riparian ecosystems have seldom been included in systematic conservation planning (Nel et al. preventing grazing. Salinas et al.2668 Biodivers Conserv (2010) 19:2667–2684 species. 2003). Iverson et al. 2009). These systems are transitional zones between aquatic and upland terrestrial environments with a linear spatial conﬁguration. Salinas et al. The impacts of woody plant encroachment on water availability are exacerbated by climate change impacts on riparian areas. Malanson 1993). Matos et al. which decreases scouring ﬂows. Tabacchi et al. or herbivory) that may eventually completely replace strictly riparian plants (species with a life cycle that requires an inundated period for seed establishment and germination). adapted to disturbance (e. Malanson 1993). Riparian ecosystems contain a high and unique number of plant species (Sabo et al. More speciﬁcally. drought) (Lyon and Gross 2005. 2001. especially in the semi-arid Mediterranean region (Nel et al. 2005. ﬂoods. 2007) such as agriculture (Aguiar and Ferreira 2005. and are becoming increasingly threatened by human activities (Salinas et al. in a restricted area of land (Lyon and Gross 2005. the study asked (1) how much of the riparian richness is due to strictly riparian species and how much is due to sclerophyllous 123 . Woody plant encroachment causes major shifts in hydrological dynamics by decreasing surface and subsurface ﬂow. 2002. ´ Virgos 2001. Machtans et al. Spackman and Hughes 1994. 2000) and upland plant encroachment (Huxman et al. and other riparian functions (for a review see Huxman et al. and transportation infrastructures (Jongman and Pungetti 2003. leading to an increase in woody plant survival. This results in higher forest cover along the channel. and decreases water availability to other plant and wildlife species. Keywords Riparian forest Á Plant richness Á Landscape metrics Á Landscape ecology Á Restoration Introduction Riparian ecosystems are highly diversiﬁed. To test if this is this pattern holds along two main watersheds in southern Portugal. 2008. 2006. water-land interface and terrestrial habitats for animal species. It is likely that such areas will be more susceptible to incursion by sclerophyllous plants (species which have developed leathery leaves to minimize water loss. as a response to nutrient poor soil. Naiman et al. Riparian plant communities in Mediterranean climates have been impoverished and threatened by human activities (Aguiar et al.. I investigated the ratio of strictly riparian and sclerophyllous species. as well as drinking water for upland animals (Brookshire et al. 2005). Riparian ecosystems also provide aquatic. Williams et al. 2005). land development for industry or tourism. Such a transition is likely to occur when space becomes available to sclerophyllous species. 2005). 2005). 2002). Hilty and Merenlender 2004. Rivers have already been inﬂuenced by changing ¨ precipitation regimes resulting from climate change (Schroter et al.g.
Topography in this area ranges from coastal areas to low altitude mountains (400 m). streams n = 24 and rivers n = 22) 123 .. Dominant plant communities are characterized by cork oak (Quercus suber) and holm oak (Q. Materials and methods Study area The study area was a 6. 1). more strictly riparian plants) woody riparian community. Carmel and Flather 2004). more intact (i.e. Mean precipitation is 500 mm/year. (2) are the strictly riparian and sclerophyllous woody plant species spatially segregated. southern Portugal. Highlighted areas indicate the riparian ecosystems sampled (creeks n = 24. rotundifolia) woodlands with ´ terrestrial shrubs such as rock-rose (Cistus spp. and (3) which combination of biotic and abiotic factors allows for a richer. 1 Location of the study area in Alentejo province. this paper assesses watershed level impacts in riparian ecosystems and proposes measures that enhance conservation of these unique riparian ecosystems in face of current threats and climate change.2°C. Portuguese Meteorology Institute.400 km2 region located within the Alentejo province of southern Portugal (Fig.Biodivers Conserv (2010) 19:2667–2684 2669 woody plant species. Based on the results of this survey. Climate varies from cold to mild during the winter to hot and dry during the summer (with a temperature range of -8 to 45.) dominating the sub-canopy (Chıcharo Fig. with a dry period from June to September (Rivas-Martinez 1987).
holm oak woodland (6%).2670 Biodivers Conserv (2010) 19:2667–2684 et al. and other (including Eucalyptus spp. 12%). resulting in a ﬁnal sample of 70 transects. Field data collection I visited all sites once for plant identiﬁcation between December 2003 to February 2004. All plant species were identiﬁed in the ﬁeld.. Each 2 km transect was subdivided into 200 m segments. intensive agriculture (e. and samples of unknown species were collected and identiﬁed in the laboratory. This distance was selected as subsamples because it matches the minimum resolvable unit in the land cover map (approximately 200 m2). holm oak woodland. drying during the summer. Since there was a clear signiﬁcant difference in river bed width between creeks. moor and heath species. African tamarisk (Tamarix africana). plantations. cereal production and cork extraction) and livestock production (cattle.5 km wide buffer of the waterway. and among surrounding dominant land-cover (cork oak woodland. a key factor affecting plant establishment and growth. vineyards.. goats and pigs). 1). The identiﬁcation was resolved to the ﬁnest taxonomic status possible. which I walked while recording the presence or absence of every woody plant species within 5 m of the bank. the distinction was considered reliable. Each waterway was surveyed using a transect parallel to the right waterway margin. and Pinus spp. and is comparable to similar surveys along riparian systems in the Iberian Peninsula (Aguiar and Ferreira 2005). and intermediate order waterways) and rivers ([10 m wide and high order waterways). in which plant species presence was recorded. grey willow (Salix atrocinerea). The stretches were equally distributed among creeks (1–5 m wide and lower order waterways). especially in the case of the willow. The majority of watercourses are intermittent. streams (5– 10 m wide. Human activities include traditional agriculture (olive groves. which lacked diagnostic features during the sampling months. streams and rivers were deﬁned by their progressively higher order. the dominant riparian species are the white poplar (Populus alba). Herbaceous species were excluded from 123 . tomato. which require water extraction from neighbouring watercourses. I derived ﬁve land-cover classes from the 1990 CORINE land-cover data (derived from classiﬁcation of Landsat TM 30m resolution multispectral imagery) within a 1. sheep. and this classiﬁcation was conﬁrmed by testing if the classiﬁed stretches had signiﬁcantly different river bed width. 2001). oleander (Nerium oleander) and alder (Alnus glutinosa) (Aguiar et al. and revisited each transect between June to September 2004 to assess any change in environmental context variables (see below). streams and rivers. cork oak woodland (23%). ´ 2006. 1%). Seventy-two sampling sites were randomly selected from this layer and screened for site accessibility. urban areas. etc. I used a digital data layer of watercourses in the study area overlain on the land-cover data to identify all possible 2 km stretches dominated by a single land-cover type within the waterway buffers. corn. 2001). Along watercourses. Chıcharo et al. raywood ash (Fraxinus angustifolia). Two river transects surrounded by holm oak woodlands were inaccessible. and result in reduction of riparian vegetation by clear-cutting or grazing. The classes are: extensive agriculture (cereal plantations) (58%). and agriculture) giving eight replicates of each waterway-type/landscape-context combination (Fig.g. with all specimens categorized at genus or species levels. Sampling design I used a stratiﬁed random sampling method to select 72 water stretches. The two seasons represent the variability of surface water in the watercourses. Creeks.
The Shannon diversity index emphasizes rare landscapes whereas the Simpson diversity index more heavily weights common landscapes.). width and area. Further. livestock. conﬁguration (number of patches).5 km from each side of the riparian zone because this was the average distance from the waterway to the top of the nearest hill. Therefore. development (houses. calculated by multiplying length by width. I made qualitative assessments of the degree of human presence through registering presence and absence of human activities (houses. and used the average value to characterise the water availability of each transect. fences and roads). Water characteristics were measured twice at the same locations. 3. I also calculated the evenness of the landscapes using the Shannon’s equitability index (J’). etc.Biodivers Conserv (2010) 19:2667–2684 2671 the analysis because of the lack of consistently identiﬁable features (due to either phenology or herbivory). holm oak. Finally. I assessed the potential impact of type of surrounding landscape (area of cork oak. I measured water depth and width every 200 m along the transect (a total of 11 measurements per transect). its degree of contact with the riparian area (edge density) and its shape complexity (area weighted mean shape index and area weighted mean fractal dimension). Richness values for strictly riparian species (species with a life cycle 123 .2 (ESRI 1996). where k is the number of classes. and for each of the land cover types its extent (patch size). which account for both the abundance and evenness of landscape (Krebs 1998). Data analysis Woody plant richness was deﬁned as the total number of identiﬁed species per sampling unit (per transect). As a proxy for the effect of propagule connectivity (Li and Wu 2004). Equitability assumes values between 0 and 1. water availability. hunting. irrigated agriculture and others). with 1 being complete evenness in landscape composition and corresponds to samples receiving the maximum value of the Shannon-Wiener index. H’ varies between 0 to log(k). the landscape diversity index and landscape equitability were calculated using Shannon-Wiener (H’) and Simpson (D) diversity indexes. Table 1) and the continuity of the riparian vegetation (vegetation was deﬁned as sparse when [100 m were traversed without ﬁnding woody species).1’’ (Rempel and Carr 2003) extension for ArcView 3. All the metrics were calculated for a buffer of 1. I measured the length and average width of the tree and shrub layers within each transect. farming. Landscape metrics were calculated using the ‘‘Patch Analyst v. each of the sampling sites was characterized according to characteristics of the riparian zone itself. characteristics of the surrounding landscape. to assess the effect of the presence of multiple surrounding landscapes on the seed sources to surveyed patches in the riparian areas. along with their length. and D varies from 0 to ??. Characteristics of the riparian zone included the structure of the riparian area (presence of tree and shrub layers. dry agriculture. Environmental variables To assess the role of biotic and abiotic factors in determining the richness of the woody riparian plant communities. once in the wet season (winter) and once in the dry season (summer). It is likely that surrounding land cover contributes a signiﬁcant source of seeds and propagules to riparian areas. and the presence of human disturbance (Table 1). I characterized the surrounding landscape using a suite of landscape metrics calculated from the available digital CORINE landcover data following their formulation in McGarigal and Marks (1995) (Table 1). and livestock along each transect (Table 1).
localized. Description Number of species in a transect Number of strictly riparian plant species Measurement Watercourse Watercourse Watercourse Type Field data Field data Field data Field data Field data Field data Field data Field data Field data Field data GIS Classes – – – – – – – Absenta. plant species Length tree Width tree Length shrub Width shrub Length of trees in a transect Watercourse Width of trees in a transect Watercourse Length of shrubs in a transect Width of shrubs in a transect Watercourse Watercourse Watercourse Continuity veg Dominant woody vegetation continuity Water width Water depth Area cork oak Average width of waterway Watercourse parallel to the transect Average depth of waterway Watercourse parallel to the transect Area within the buffer covered by cork oak woodlands Buffer Area holm oak Area within the buffer covered by holm oak woodlands Area agriculture Area of irrigated agriculture Area within the buffer covered by dry agriculture Area within the buffer covered by irrigated agriculture Buffer GIS – m2 Buffer GIS – m2 Buffer GIS – m2 Area of others Area within the buffer covered by other land covers NUMP MPS ED AWMSI AWMFD SDI SEI Human activities Number of patches in the buffer Mean patch size Edge density Area weighted mean shape index Buffer GIS – m2 Buffer––patch Buffer––patch Buffer––edge GIS GIS GIS – – – – – – – None.2672 Biodivers Conserv (2010) 19:2667–2684 Table 1 Environmental parameters and their description. where they were measured. Densec – – – cm cm m2 Units – – – m m m m Sclerophyllous Number of sclerophyllous spp. Scarceb. type of variable. richness Riparian spp. classes and units at the 70 sampled riparian stretches in southern Portugal Variable Spp. frequent ha m/ha ha-1 ha-1 Buffer––shape GIS Area weighted mean fractal Buffer––shape GIS dimension Shannon’s diversity Index Shannon’s equitability Index Presence of human activities Buffer–– diversity Buffer–– diversity Watercourse GIS GIS Field data 123 .
intense Absent. with values close to 0 representing no spatial autocorrelation. intense 2673 Units Cattle Sheep Goats Pigs a b c Degree of use of the riparian area by cattle Degree of use of the riparian area by sheep Degree of use of the riparian area by goats Degree of use of the riparian area by pigs Watercourse Watercourse Watercourse Watercourse Field data Field data Field data Field data Vegetation present but with no deﬁned structure in the segment Vegetation scattered and scarce throughout the segment Vegetation continuous and dense throughout the segment that requires an inundated period for seed establishment and germination) and sclerophyllous species (species which have developed leathery leaves to minimize water loss. the more samples collected the fewer new species are expected to be found. Since no signiﬁcant spatial autocorrelation was found at distances above 1. intense Absent. To test how much of the total richness is a function of the riparian and the sclerophyllous plants. i. scarce. This indicates that the sample size was sufﬁcient to characterize the variability in the study area. scarce. walls. and as a response to poor nutrient soils and herbivory) were also calculated. 1989). The effects of spatial autocorrelation on transect location were tested using Moran’s I index (Moran 1950). A sample was considered sufﬁcient when the curve of the cumulative number of identiﬁed species plotted against the number of samples reaches an asymptote. Spatial autocorrelation was calculated using ROOKCASE Microsoft Excel Add-in (Sawada 1999). Species-transect curves were calculated in PC-ORD (McCune and Grace 2002). artiﬁcial water channels Absent.. One-way ANOVA was used to determine if the riparian plant community richness was a function of the watercourse type.5 km.Biodivers Conserv (2010) 19:2667–2684 Table 1 continued Variable Human structures Description Human infrastructures found in the transect Measurement Watercourse Type Field data Classes Roads. after testing for normality in the distribution of the variables and transforming accordingly (log transforming area of landcover) (Zar 1999). it was concluded that spatial autocorrelation was not affecting the data and therefore it could be used for further analysis. scarce.e. the neighborhood distance was progressively increased from a radius of 1000–5000 m in 1000 m increments and I measured Moran’s I index for each radius distances. a regression was ﬁtted between the total species richness and the richness of riparian 123 . fences. In order to assess if the samples were sufﬁcient to describe study-area-wide riparian vegetation richness I used a species transect curve. The number of samples at which the asymptote is reached corresponds to the sufﬁcient sample size required (Krebs 1998). streams (n = 24) and rivers (n = 22). houses. intense Absent. even when separating between creeks (n = 24). and varies from -1 (perfect negative spatial autocorrelation) to 1 (perfect positive spatial autocorrelation). To estimate the distance threshold at which spatial autocorrelation could be considered negligible. scarce. This index measures the similarity in the spatial patterns of the variable (Fortin et al. and an asymptote was reached with 22 sampling transects. in our case woody species richness.
05 was used. model ﬁt (explanatory power) was assessed using R-square values. Since there was not signiﬁcant collinearity between any of the predictor variables. A linear regression was also ﬁtted between the richness of riparian and sclerophyllous plants to identify a relationship between the two. willows (40. olive tree (31%). African tamarisk.9 species.2674 Biodivers Conserv (2010) 19:2667–2684 and sclerophyllous plants. streams and rivers (P [ 0.46. Strictly riparian had a mean richness of 6. d.f. Strictly riparian species included white willow and other willows. whereas sclerophyllous species had a mean richness of 5. I tested whether the two vegetation types were present in the same spatial location (i. Sclerophyllous plant species were consistently found across all sampling units.6 ± 7. Species richness had a mean of 15. Model signiﬁcance was assessed using F-test values. the percentage of the total richness due to riparian and sclerophyllous plant species was estimated and a linear regression ﬁtted between the total richness and the percent of riparian and sclerophyllous in the community.5 species per transect. For these tests a signiﬁcance level of 0. black poplar. which included strictly riparian and sclerophyllous plant species (Appendix Table 3).1%). as they were not signiﬁcantly different among creeks.2%) were the most common tree species. quinces. and others (see Appendix Table 3).e. When considering all species.5%) and rockrose (36. except for 10% of transects (7 out of 70) where no sclerophyllous species were detected.01).1%) were the most common shrubs. complete replacement (slope = 0) or partial replacement (0 \ slope \ 1). Exotic species such as acacia and eucalyptus were also commonly found. cork oak (40.3 species. Linear regression was used to test if within each segment higher richness of strictly riparian plants was correlated with higher richness of sclerophyllous vegetation. they were maintained for further analysis. black poplar (33. the same 200 m sample) or spatially segregated in the same riparian zone. Strictly riparian species richness was signiﬁcantly higher than sclerophyllous plants (F = 6. and for statistically signiﬁcant models (a = 0. lentisc and rock-roses. richness of strictly riparian plants and richness of sclerophyllous plant species. To assess the proportion of the richness that each of these groups contributes. The slope of the regression line indicates additive richness (slope = 1). species richness did not vary as a function of watercourse type. = 138. and holm oak (30. P = 0. Raywood ash (60. A generalized linear model (GLM) was used to test the effect of each of the environmental context variables in the total riparian plant richness. and raywood ash.05 123 . All statistical analyses were performed using JMP 5..6 ± 2. and Bonferroni corrections were applied to correct signiﬁcance values for multiple comparisons (Zar 1999).2 ± 3.0 (SAS Institute) for Windows. including pears. The correlation between each of the environmental context variables (Table 1) was tested using Pearson correlation coefﬁcients (Zar 1999). Sclerophyllous species included cork and holm oak. The patch structure of the riparian zones was analysed by comparing the segments of each transect in terms of their riparian and sclerophyllous composition. with a maximum of 33 different species in one transect and a minimum of two species.7%).1). and so were fruit trees. If the slope of the regression was negative it would indicate spatial segregation. and blackberry (79. Results Riparian plant richness Riparian plant communities were composed of 53 different woody plant species.1%).6%).
Strictly riparian plant richness was also signiﬁcantly increased by a surrounding landscape with high number of large holm oak patches. and 28% were sclerophyllous plants. indicating no spatial segregation. Sclerophyllous plants richness alone was also not signiﬁcantly different along watercourses (F = 0. The ﬁnal statistical model explained about 70% of the variability in total woody species richness. 2006). indicating that most of the total riparian richness is due to sclerophyllous plant species. 46% of the woody plant species were strictly riparian plants. the percentage of strictly riparian plants signiﬁcantly decreased (P \ 0. Since the areas that were surveyed in this study are similar to those studied previously and.Biodivers Conserv (2010) 19:2667–2684 2675 for all tests). and diversity of interfaces between ´ aquatic and terrestrial habitats (Naiman and Decamps 1997). these communities have on average 16 woody riparian plant species in 100 m (Aguiar and Ferreira 2005. was signiﬁcantly higher along rivers (F = 5. Aguiar et al.0001) and the percentage of sclerophyllous plants signiﬁcantly increased (P \ 0. ﬂow-facilitated dispersal of seeds and propagules (Deferrari and Naiman 1994).0001.19) but signiﬁcant positive correlation between strictly riparian and sclerophyllous plant species richness within riparian areas (Fig.02. it is not likely that the discrepancy in the 123 .51) (Fig. d. P = 0. Fig. 2a). All GLMs were signiﬁcant (Table 2). Riparian species richness. there is a stronger relationship between total riparian richness and the sclerophyllous plant richness (R2 = 0. disturbance (Pollock et al. On average. soil variability (Pollock et al.51. 1998). However. as well as strictly riparian and sclerophyllous richness were mainly a function of the areas of shrubs in the riparian ecosystem (except for sclerophyllous plants richness). and similar values were attained for both strictly riparian (69%) and sclerophyllous species (71%). and with low diversity of land covers (Table 2). Regression between strictly riparian and sclerophyllous plants in each 200 m segment was not signiﬁcant (P [ 0. with less than half of the sites (31 out of 70) having more than 15 species. Environmental variables associated with riparian plant richness Higher total woody plant richness. Discussion Riparian plant richness Previous studies of richness of comparable riparian systems in the Iberian Peninsula have shown that in the last 5 years. = 69. There was a weak (R2 = 0. 3). geographical and topographical variability (Naiman and ´ Decamps 1997). Sclerophyllous plants richness increased in reduced areas of agriculture and reduced human activities and goats (Table 2).009) than either creeks or streams. d. such as productivity (Pollock et al. P = 0. in some cases. Lowest total richness was associated with a community dominated by strictly riparian and high total richness was due to the combined presence of strictly riparian and encroachment by sclerophyllous species. 2b). as well as the absence of human activities and goats (Table 2). at the same locations of studies from other authors.f.05). = 69. when considered alone.f. The remainder of the variability was accounted for by exotic and fruit trees. 1998). As the total richness of the community increased.6). 1998). Several factors contribute to richness in riparian plant communities.84) than that between total riparian richness and strictly riparian plant species (R2 = 0. The results presented in this study show lower values (average richness of eight species per 100 m. Table 2).
right axis.6 0.2 0. However.6 0. dotted line) and sclerophyllous (open circles. The stronger explanatory power of the sclerophyllous regression indicates an additive effect of sclerophyllous species to total richness.0129x + 0.9 0. full line) plant species with total plant species richness.9 0.1 0 0 5 10 15 20 25 30 35 (b) % Strictly Riparian Plant Species 1 0. Furthermore.5068 14 12 10 8 Sclerophyllous Plant Species Richness 6 4 2 0 Sclerophyllous y = 0.1 0 % Sclerophyllous Plant Species Total Plant Species Richness % Riparian Linear (% Riparian) % Sclerophyllous Linear (% Sclerophyllous) Fig. A recent study focusing on ranking the health of riparian systems in southern Portugal found that most riparian stretches are in poor condition (Matos et al.014x + 0.8426 6 4 2 0 0 5 10 15 20 25 30 35 Total Plant Species Richness Strictly riparian Linear (Strictly riparian) Sclerophyllous Linear (Sclerophyllous) 1 y = -0. right axis. dotted line) and sclerophyllous (open circles.7863 R² = 0. left axis. current riparian clear cut practices. The proportion of each group changes as total richness increases results is due to differences in productivity. 123 .2 0. (b) Regression of % strictly riparian (closed circles. and geographical and topographical variability. left axis.8 0.4 0. soil variability. full line) plant species with total plant species richness. the degree of disturbance of the sites in the current study may be higher than that of previous studies.507 R² = 0.2424x + 2. ﬂow-facilitated dispersal of seeds and propagules.3 0. 2008).2676 Biodivers Conserv (2010) 19:2667–2684 (a) 14 Strictly Riparian Plant Species Richness 12 10 8 16 Strictly riparian y = 0.5 0.4908x .3538 y = 0.2.3 0.3669 0.5 0.7 0. 2 (a) Regression of strictly riparian (closed circles.8 0.4 0.6878 R² = 0. such as those present in the study area.7 0.0859 R² = 0.
009 (0. and 123 .Biodivers Conserv (2010) 19:2667–2684 16 2677 Sclerophylous Plant Species Richness 14 12 10 8 6 4 2 0 0 2 4 6 8 y = 0. Bold values indicate signiﬁcance at P-value less than 0.26 (0.55) 0. winter and summer water depth and width. plant equitability. This is not unexpected as riparian plant richness is unique (Sabo et al.09) 0.08) 25. 2005).005 (0.002 (0.71 2. There is a positive relationship between the two plant groups.06 (0.1) 20. 3 Relationship between strictly riparian and sclerophyllous species richness.23 (0.16 (0.12 (0.03) 210.44) 0. sheep and pigs did not retrieve signiﬁcant results thus were excluded from the table have been identiﬁed as a cause of decreasing riparian species richness in the Iberian Peninsula (Salinas et al.09) 0.76 (0.6651 R² = 0.03) 0. Higher total plant richness was recorded when more of the sclerophyllous species were present. but highly variable Table 2 Generalized linear models for the total riparian plant richness.067 (0.12 (0.01) 22. patch complexity (Area-weighted mean shape index and area-weighted mean fractal dimension).6836x + 0. edge density.08) 20.69 1. area of cork oak.95 (0.69 (0.42 (0.07) 2.02) 65 Sclerophyllous Estimate (P-value) 93.09) 22.65 (0.6.70 2.05 Measurements of area of tree.04) 61 23.03) 66 21.01) 24.07) Strictly riparian Estimate (P-value) 44.005 (0.09) 0.62 (0.29) Intercept Area shrubs N patches Mean patch area Shannon diversity index Area holm oak Area agriculture Human activities Human structures Goats R-square F-test (P-value) d.66 (0.94 (0. strictly riparian and sclerophyllous plant richness found along watercourses in southern Portugal Variable Total richness Estimate (P-value) 99.81 (0.1895 10 12 14 Strictly Riparian Plant Richness Fig.06) 0. 2000). presence of cattle.05) 0.f.9 (0.
Salinas et al. Human activities in the riparian ecosystem included development of roads. houses. In fact. These results may be explained by the heterogeneity of the riparian ecosystem itself at the ﬁner scale (segments) and that of the landscape at the larger scale (transects). 1998. These factors have also been identiﬁed by other authors as major causes of the decrease in strictly riparian richness in other riparian areas (Aguiar and Ferreira 2005. changes in water rights policies have altered the management prescriptions for riparian zones. Environmental variables associated with riparian plant richness Pollock et al. Pollock et al. which was almost twice the average contribution of the sclerophyllous plants (28%).2678 Biodivers Conserv (2010) 19:2667–2684 thus contribution from upland plants can only result in an increase in local richness. The ﬁndings in this study corroborate this prediction. 2001). Hilty and Merenlender 2004. 2002). higher absolute numbers of sclerophyllous than riparian plant species were recorded. Salinas et al. As both natural and human-mediated disturbances create gaps in the riparian ecosystem ´ (Naiman and Decamps 1997. Malanson 1993. Since strictly riparian plants occur in limited numbers (Pollock et al. as total richness increases. which in turn create higher fragmentation and gaps within the riparian vegetation. and the limitation on maximum richness of strictly riparian species explains the negative slope in the regression between total species richness and strictly riparian plants. 2006). (1998) demonstrated that species richness in riparian ecosystems is a dynamic equilibrium between disturbance frequency and community productivity. On average 48% of the total plant richness was due to the presence of strictly riparian plants. My results however. Tabacchi et al. 2000. I predicted that a more heterogeneous riparian ecosystem would have higher total woody species richness. and increases are only possible with the addition of sclerophyllous species. allowing neighbouring land-owners to clear-cut riparian trees for easier access to water. sclerophyllous species richness increases at a similar rate. This shift towards a water resistant community was also observed in the Tagus river system in Portugal (Aguiar et al. Alternatively. fences. These authors suggested that richness is increased by the presence of patches of different seral stages. This indicates that the riparian community is mainly dominated by strictly riparian plants. which may be explained by encroachment of upland species as a response to climate induced reduction in water levels in the upland habitats (Gasith and Resh 1999). and some of those seeds could have become established either naturally or following disturbance (Aguiar et al. and artiﬁcial water channels. 2007). 1998. 2005) the maximum richness is truncated. walls. these results also indicate a constant presence of sclerophyllous species in riparian ecosystems. which contain different suites of species. Spatial segregation between strictly riparian and sclerophyllous plant patches may be an important factor in determining propagule pressure from sclerophyllous plants species into riparian areas. 2000). 123 . Miller 2002. However. upland and exotic species are known to use riparian ecosystems for dispersal (Schnitzler et al. Sabo et al. Furthermore. this creates opportunities to the establishment of both riparian and sclerophyllous plants in similar locations (Tabacchi et al. In the context of this study. only show spatial segregation between transects (between 2 km transects) and not within transects (between 200 m segments of each transect). 2). from the negative relationship between richness and presence of human activities it can be inferred that increased sclerophyllus richness does not seem to be a function of the structure of the riparian ecosystem. while riparian species richness has a lower effect (Fig. which would be mostly due to the presence of sclerophyllous plants in addition to (rather than replacing) strictly riparian plants. However.
blackberry bushes may facilitate the germination seeds from these species. and holm oak woodlands do not seem to be exporting seeds elsewhere. mixed mosaics of riparian and sclerophyllous plant species appear as older seral stages. Finally.) have a very negative effect on the strictly riparian plant richness. At the same time. and also prevent seed mortality from desiccation by providing shade (Karrenberg et al. This study results also revealed that riparian species richness (total and strictly riparian) was positively affected by the presence of a developed shrub layer and it was negatively affected by the presence of goats. however. the second is a sclerophyllous shrub. which had a marked negative effect on the overall plant richness. 2002). 2004. Blackberry shrubs tend to create a very dense canopy. which may prevent light from reaching the riparian species seeds. However. as the total riparian community is inundated by propagules from different types of plant species. suggesting that these fewer larger patches may be contributing greater numbers of sclerophyllous plant propagules to the riparian ecosystem. olive yards. The most commonly found shrub species in the study area were blackberry shrubs (79. Thus. In the Mediterranean. forbs and woody species. and rock-rose (36. Blackberry shrubs are probably the most related to the observed positive inﬂuence on riparian richness. The results showed that strictly riparian plant richness was positively affected by fragmentation (higher number of patches) of the surrounding landscape. willows and poplar seeds are known to germinate in the dark (Karrenberg et al. sheep are intermediate foragers that consume grasses. 2002). if the surrounding land cover is mainly holm oak woodlands. since they are the ones most detected. currently below replenishment rates. Goat foraging period (May–June. which may have different establishment success rates in the different open patches within the riparian area. (1998) pattern of different seral stages. and goats are browsers that consume forbs and woody species and avoid grasses (Vallentine 2001). Furthermore. Portuguese Associations for Bovine and Ovine and Caprice livestock production. sheep and goats share the same forage areas and are separated temporally and behaviorally (Vallentine 2001) by different foraging preferences. Cows are grazers that consume grasses and avoid woody species. matching Pollock et al. this effect may be only temporary. reduced sclerophyllous seeds exist in the seed pool to colonize the riparian zone. patches of a variety of different land covers (holm oak. and it was negatively affected by the presence of patches of different landscapes (as measured by the landscape diversity indexes). unpublished data) coincides with the time when young woody riparian plants have reached the sapling stage and become more conspicuous. diversity of land covers and their density. cork oak woodlands. making them more vulnerable to herbivores. As more agricultural land exists around the riparian area. the thorns and the dense canopy of these shrubs may prevent the saplings from being eaten by goats. resulting ultimately in an increase in total species richness. This can also explain the negative effect of the area of agriculture on the richness of sclerophyllous plants in the riparian ecosystem.5%). the results indicate that fewer riparian plants are found when larger sclerophyllous patches surround the riparian ecosystem. Younger seral stages will be dominated by riparian plants. While the ﬁrst is mostly found in riparian areas. etc. Ramirez and Diaz 2008). and as sclerophyllous species may colonize gaps.Biodivers Conserv (2010) 19:2667–2684 2679 2002). First.1%). which occurs in a short period (a few days). Three factors may contribute to this pattern: the total area covered by the different land covers. the frequency of seeds and propagules may actually be reduced since this landscape is characterized by a sparse canopy that is experiencing a decreasing trend in recruitment (Plieninger et al. cows. 123 .
Margaret Andrew. and collaborative campaigns to increase the awareness of these valuable resources that involve land-owners. as a large sample size (n = 70) for a study of this type was achieved. Alex Fremier. the range of variability among riparian systems found in Mediterranean climates was covered. Open Access This article is distributed under the terms of the Creative Commons Attribution Noncommercial License which permits any noncommercial use. which could be used for restoration and reforestation of riparian zones. For both ecophysiological groups of plants (strictly riparian and sclerophyllous) all the species that had diagnostic features were identiﬁed. microclimate moderation. The maintenance of riparian ecosystems and their role as ecosystem service providers may require creating a riparian plant seed bank and nursery. MJS: POCTI/MGS/47435/2002). water temperature control. and service as corridors for wildlife (Miller 2002). Judson Wynne. and J. 70% of explained variability). 123 . and reproduction in any medium.2680 Biodivers Conserv (2010) 19:2667–2684 Data quality assessment The quality of the interpretation of the results also depends upon the quality of the data input to the models. its effect is likely negligible. richness estimates would be increased at most by eight species. remove existing exotic plants. creation of distinct habitats and communities. water quality regulation. Implications for conservation of riparian ecosystems in semi-arid Mediterranean climates The change in the composition and structure of riparian ecosystems observed in this study may result in the loss of the suite of ecosystem services they provide. Furthermore. Francis Bozzolo and Erin Hestir provided helpful improvements to this manuscript. and willow. If all the taxa were identiﬁed to species level. Further conservation measures should promote riparian ecosystem aesthetic. and the statistical models were signiﬁcant and returned a very good ﬁt to the data (ca. It is acknowledged that some underestimation may have occurred of species richness as some species lacked key characters that allowed their differentiation. All these measures can be included in a more encompassing and stronger protection policy especially designed for the conservation of the unique biodiversity and services of riparian ecosystems. recreational and service provider values through economic incentives and subsidies. ˆ Acknowledgments The present paper was funded by the Fundacao para Ciencia e Tecnologia project ¸˜ ‘‘Riparian galleries as Corridors and Linkage Habitats in the Fragmented Landscape of Southern Portugal: Applications to Conservation Planning’’ (POCTI/MGS/47435/2002. The author is extremely grateful to Hugo M. shelter and nutrients for aquatic and terrestrial organisms. water storage. and preventing or limiting grazing. Even though this underestimation may make comparison of these results to those of other authors more difﬁcult. heath and moor species were the ones harder to diagnose (see Appendix Table 3). provided the original author(s) and source are credited. managers and users. Future preservation of riparian ecosystems should include monitoring of edge encroachment by upland plants and management to prevent their establishment within a buffer zone around the riparian system. Further thanks are due to Ana Rita Alves and Luıs Miguel Rosalino for their help during ﬁeldwork. which resulted in comparable underestimations. including soil and bank stabilization. distribution. Matos for the ﬁeld work and to Margarida Santos-Reis for the ´ logistic and ﬁnancial support. At the larger scale. conservation of riparian ecosystems through limitations in grazing and tramping by livestock and water use regimes is recommended.
S.3 2.R C.9 7.S. Ficus carica Fraxinus angustifolia (*) Olea europaea Phillyrea angustifolia (*) Pinaceae Poaceae Punicaceae Rhamnaceae Pinus pinaster Pinus silvestris Arundo donax Phyllostachys spp.S.6 15.R C.S. Fagaceae Quercus coccifera Quercus faginea Quercus rotundifolia Quercus suber Lamiaceae Lauraceae Myrtaceae Moraceae Oleaceae Lavandula stoechas Laurus nobilis Eucalyptus globulus Myrtus spp.R C.S.R C. Ceratonia siliqua Genista spp.9 18.R C.R C.R C.3 2.R C.R C. PlPlantation). Ex-Exotic.4 4.R C S C.R C.6 10 10 1.S.S.Biodivers Conserv (2010) 19:2667–2684 2681 Appendix Table 3 List of species detected and frequency of detection in the 70 sampled transects (percent of transects).R C.R C S.R C.6 Sc SR SR Sc Sc Sc Sc Sc Sc Sc Sc Sc Ex Sc Sc Ex F Sc SR Sc Sc Sc Sc Sc Sc Sc Ex Sc F SR Sc Sc Pl Ex SR Ex F Sc C.R C.R C.4 1. and R-river) Family Scientiﬁc name Common name Frequency Eco-phys.S.6 14.S.S.S.4 2.9 1.6 4.S.R Anacardiaceae Pistacia lentiscus (*) Apocynaceae Betulaceae Caprifoliaceae Cistaceae Nerium oleander Alnus glutinosa Lonicera implexa (*) Viburnum tinus Cistus albidus (*) Cistus crispus (*) Cistus ladaniferus Cistus monspeliensis Cistus populifolius (*) Cistus salvifolius Halimium halimifolium (*) Cupressaceae Ericaceae Fabaceae Arbutus unedo Erica arborea Acacia spp.3 77.S.R R C.1 68.S.9 28.R C.S. Punica granatum Rhamnus alaternus (*) Mastic Oleander Black alder Honeysuckle Laurestine White-leaved rockrose Rockrose Gum rockrose Montpellier rockrose Rockrose Sage-leaf rockrose Halimium Strawberry tree Briar root Wattle Carob tree Spanish broom Retama Gorse Kermes oak fff Holm oak Cork oak French lavender Sweet bay Tasmanian bluegum Myrtle Fig tree White ash Olive tree False olive Maritime pine Scotch Pine Giant reed Bamboo Pomegranate Italian buckthorn Chamaecyparis lawsoniana Oregon cedar 123 .S C.R C C.S.R C.R S. eco-physiological group (SR-Strictly Riparian.9 14.R C.7 60 1. S-stream.S.S.R C.S.6 1.R C.S.S. Ulex spp.R C.R C.S.S. F-fruit tree.S.S.4 18.7 30 14. Sc-Sclerophyllous.7 14.3 21.3 25. Retama spp.9 2.9 58.4 38.R S S C. and type of river system (C-creek.3 15. Waterway (%) G 32.S.9 12.1 22.4 60 62.3 40 24.R S C.
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