Patterns of use and damage by exotic deer on native plant communities in northwestern Patagonia

Ungulate overabundance can strongly alter composition and structure of forest communities through selective damage independently of the herbivory history of the recipient system. In the early 1900s, red and fallow deer (Cervus elaphus and Dama dama, respectively) were introduced to northwestern Patagonia. We studied patterns of relative habitat use, damage, and species selection by introduced deer populations in three major plant communities that have the longest history of invasion in the region. We also evaluated community structure and composition characteristics as possible explanatory factors for the observed patterns of browsing, fraying, and bark-stripping. At the species level, exotic deer browsed more than expected on two evergreen species (Austrocedrus chilensis and Schinus patagonicus) and two spiny species (Colletia hystrix and Dasyphyllum diacanthoides), while one dominant shrubland tree (Lomatia hirsuta) was selected for fraying and bark-stripping. Browsing was the more widespread damage in all plant communities, while fraying and bark-stripping occurred at low incidence. Furthermore, species identity was found as the main driver of deer damage over plant community type. Finally, variation in damage and in habitat use was explained by community structure and composition characteristics. Bark-stripping and fraying are best predicted by community composition, whereas browsing and habitat use are best predicted by structural properties including understory cover, cover of fallen logs, and tree density. The patterns of damage and the association with community structure characteristics reported here provide insight for developing management strategies.     

Ungulates increase forest plant species richness to the benefit of non‐forest specialists

Large wild ungulates are a major biotic factor shaping plant communities. They influence species abundance and occurrence directly by herbivory and plant dispersal, or indirectly by modifying plant‐plant interactions and through soil disturbance. In forest ecosystems, researchers’ attention has been mainly focused on deer overabundance. Far less is known about the effects on understory plant dynamics and diversity of wild ungulates where their abundance is maintained at lower levels to mitigate impacts on tree regeneration. We used vegetation data collected over 10 years on 82 pairs of exclosure (excluding ungulates) and control plots located in a nation‐wide forest monitoring network (Renecofor). We report the effects of ungulate exclusion on (i) plant species richness and ecological characteristics, (ii) and cover percentage of herbaceous and shrub layers. We also analyzed the response of these variables along gradients of ungulate abundance, based on hunting statistics, for wild boar (Sus scrofa), red deer (Cervus elaphus) and roe deer (Capreolus capreolus). Outside the exclosures, forest ungulates maintained higher species richness in the herbaceous layer (+15%), while the shrub layer was 17% less rich, and the plant communities became more light‐demanding. Inside the exclosures, shrub cover increased, often to the benefit of bramble (Rubus fruticosus agg.). Ungulates tend to favour ruderal, hemerobic, epizoochorous and non‐forest species. Among plots, the magnitude of vegetation changes was proportional to deer abundance. We conclude that ungulates, through the control of the shrub layer, indirectly increase herbaceous plant species richness by increasing light reaching the ground. However, this increase is detrimental to the peculiarity of forest plant communities and contributes to a landscape‐level biotic homogenization. Even at population density levels considered to be harmless for overall plant species richness, ungulates remain a conservation issue for plant community composition.     

Long-Term Regional Shifts in Plant Community Composition Are Largely Explained by Local Deer Impact Experiments

The fact that herbivores and predators exert top-down effects to alter community composition and dynamics at lower trophic levels is no longer controversial, yet we still lack evidence of the full nature, extent, and longer-term effects of these impacts. Here, we use results from a set of replicated experiments on the local impacts of white-tailed deer to evaluate the extent to which such impacts could account for half-century shifts in forest plant communities across the upper Midwest, USA. We measured species'' responses to deer at four sites using 10–20 year-old deer exclosures. Among common species, eight were more abundant outside the exclosures, seven were commoner inside, and 16 had similar abundances in- and outside. Deer herbivory greatly increased the abundance of ferns and graminoids and doubled the abundance of exotic plants. In contrast, deer greatly reduced tree regeneration, shrub cover (100–200 fold in two species), plant height, plant reproduction, and the abundance of forbs. None of 36 focal species increased in reproduction or grew taller in the presence of deer, contrary to expectations. We compared these results to data on 50-year regional shifts in species abundances across 62 sites. The effects of herbivory by white-tailed deer accurately account for many of the long-term regional shifts observed in species'' abundances (R² = 0.41). These results support the conjecture that deer impacts have driven many of the regional shifts in forest understory cover and composition observed in recent decades. Our ability to link results from shorter-term, local experiments to regional long-term studies of ecological change strengthens the inferences we can draw from both approaches.     

Effects of bird predation on arthropod abundance and tree growth across an elevational gradient

Considerable uncertainty surrounds the conditions under which birds can cause trophic cascades. In a three‐year experiment, we studied the direct and indirect effects of insectivorous birds on arthropod abundance, herbivory, and growth of striped maple Acer pensylvanicum saplings in a northern hardwood forest of central New Hampshire, USA. We manipulated bird predation by erecting exclosures around saplings and directly manipulated herbivory by removing herbivores. We also examined how climate modifies these interactions by replicating the experiment at three locations along an elevational gradient. Effects of bird predation were variable. Overall, mean arthropod biomass was 20% greater on saplings within bird exclosures than on controls (p<0.05). The mean biomass of leaf‐chewing herbivores, primarily Lepidoptera larvae, was 25% greater within exclosures but not statistically different from controls. To a lesser degree, mean herbivore damage to foliage within exclosures exceeded that of controls but differences were not significant. We also did not detect significant treatment effects on sapling shoot growth. The high understory vegetation density relative to bird abundance, and low rate of herbivory during the study (mean 5% leaf area removed, controls), may have limited the ability of birds to affect sapling growth. Climate effects operated at multiple scales, resulting in a complex interplay of interactions within the food web. Regional synchrony of climatic conditions resulted in annual fluctuations in herbivore abundance and tree growth that were shared across elevations. At the same time, local environmental variation resulted in site differences in the plant, herbivore, and bird communities. These patterns resulted in a mosaic of top–down strengths across time and space, suggesting an overall pattern of limited effects of birds on plant growth, possibly interspersed with hotspots of trophic cascades.     

Facilitative interactions between an exotic mammal and native and exotic plants: hog deer (<Emphasis Type="Italic">Axis porcinus</Emphasis>) as seed dispersers in south-eastern Australia

Endozoochory by exotic mammalian herbivores could modify vegetation composition by facilitating the dispersal and establishment of exotic and native plant species. We examined the potential for endozoochoric dispersal of native and exotic plants by exotic hog deer (Axis porcinus) in south-eastern Australia. We quantified the germinable seed content of hog deer faecal pellets collected in five vegetation types within a 10,500-ha study area that was representative of their Australian range. Twenty exotic and 22 native species germinated from hog deer faecal pellets and significantly more native species germinated compared to exotic species. Seedlings of the encroaching native shrub Acacia longifolia var. sophorae emerged, but no native trees emerged and the percentage of grasses that germinated was low (11%). The species composition of germinants was similar among the five vegetation types. We estimated that the hog deer population in our study area could potentially disperse >130,000 viable seeds daily. Our study shows how an exotic mammal can disperse seeds from both native and invasive plants and highlights the need for endozoochory to be considered more widely in studies assessing the impacts of exotic mammals on plant communities.     

Do riparian plant community characteristics differ between <Emphasis Type="Italic">Tamarix</Emphasis> (L.) invaded and non-invaded sites on the upper Verde River,Arizona?

Invasion by Tamarix (L.) can severely alter riparian areas of the western U.S., which are globally rare ecosystems. The upper Verde River, Arizona, is a relatively free-flowing river and has abundant native riparian vegetation. Tamarix is present on the upper Verde but is a minor component of the vegetation (8% of stems). This study sought to determine whether riparian vegetation characteristics differed between sites where Tamarix was present and sites where Tamarix was absent during the invasion of the upper Verde. We hypothesized that herbaceous understory and woody plant communities would differ between Tamarix present and absent sites. Our hypothesis was generally confirmed, the two types of sites were different. Tamarix present sites had greater abundance of all vegetation, native understory species, graminoids, and native trees, and a positive association with perennial native wetland plant species. Tamarix absent sites had greater abundance of exotic plants and upland adapted plants and an association with greater abiotic cover and litter. These results are contrary to other reports of Tamarix association with depauperate riparian plant communities, and suggest that Tamarix invasion of a watershed with a relatively natural flow regime and a robust native plant community follows similar establishment patterns as the native riparian plant community.     

Resistance to wildfire and early regeneration in natural broadleaved forest and pine plantation

The response of an ecosystem to disturbance reflects its stability, which is determined by two components: resistance and resilience. We addressed both components in a study of early post-fire response of natural broadleaved forest (Quercus robur, Ilex aquifolium) and pine plantation (Pinus pinaster, Pinus sylvestris) to a wildfire that burned over 6000 ha in NW Portugal. Fire resistance was assessed from fire severity, tree mortality and sapling persistence. Understory fire resistance was similar between forests: fire severity at the surface level was moderate to low, and sapling persistence was low. At the canopy level, fire severity was generally low in broadleaved forest but heterogeneous in pine forest, and mean tree mortality was significantly higher in pine forest. Forest resilience was assessed by the comparison of the understory composition, species diversity and seedling abundance in unburned and burned plots in each forest type. Unburned broadleaved communities were dominated by perennial herbs (e.g., Arrhenatherum elatius) and woody species (e.g., Hedera hibernica, Erica arborea), all able to regenerate vegetatively. Unburned pine communities presented a higher abundance of shrubs, and most dominant species relied on post-fire seeding, with some species also being able to regenerate vegetatively (e.g., Ulex minor, Daboecia cantabrica). There were no differences in diversity measures in broadleaved forest, but burned communities in pine forest shared less species and were less rich and diverse than unburned communities. Seedling abundance was similar in burned and unburned plots in both forests. The slower reestablishment of understory pine communities is probably explained by the slower recovery rate of dominant species. These findings are ecologically relevant: the higher resistance and resilience of native broadleaved forest implies a higher stability in the maintenance of forest processes and the delivery of ecosystem services.     

Enemy release or invasional meltdown? Deer preference for exotic and native trees on Isla Victoria,Argentina

Abstract How interactions between exotic species affect invasion impact is a fundamental issue on both theoretical and applied grounds. Exotics can facilitate establishment and invasion of other exotics (invasional meltdown) or they can restrict them by re‐establishing natural population control (as predicted by the enemy‐release hypothesis). We studied forest invasion on an Argentinean island where 43 species of Pinaceae, including 60% of the world's recorded invasive Pinaceae, were introduced c. 1920 but where few species are colonizing pristine areas. In this area two species of Palearctic deer, natural enemies of most Pinaceae, were introduced 80 years ago. Expecting deer to help to control the exotics, we conducted a cafeteria experiment to assess deer preferences among the two dominant native species (a conifer, Austrocedrus chilensis, and a broadleaf, Nothofagus dombeyi) and two widely introduced exotic tree species (Pseudotsuga menziesii and Pinus ponderosa). Deer browsed much more intensively on native species than on exotic conifers, in terms of number of individuals attacked and degree of browsing. Deer preference for natives could potentially facilitate invasion by exotic pines. However, we hypothesize that the low rates of invasion currently observed can result at least partly from high densities of exotic deer, which, despite their preference for natives, can prevent establishment of both native and exotic trees. Other factors, not mutually exclusive, could produce the observed pattern. Our results underscore the difficulty of predicting how one introduced species will effect impact of another one.     

Comparisons of arthropod assemblages on an invasive and native trees: abundance,diversity and damage

The success of exotic plants may be due to lower herbivore loads than those on native plants (Enemies Release Hypothesis). Predictions of this hypothesis include lower herbivore abundances, diversity, and damage on introduced plant species compared to native ones. Greater density or diversity of predators and parasitoids on exotic versus native plants may also reduce regulation of exotic plants by herbivores. To test these predictions, we measured arthropod abundance, arthropod diversity, and foliar damage on invasive Chinese tallow tree (Triadica sebifera) and three native tree species: silver maple (Acer saccharinum), sycamore (Platanus occidentalis), and sweetgum (Liquidambar styraciflua). Arthropod samples were collected with canopy sweep nets from six 20 year old monoculture plots of each species at a southeast Texas site. A total of 2,700 individuals and 285 species of arthropods were caught. Overall, the species richness and abundance of arthropods on tallow tree were similar to the natives. But, ordination (NMS) showed community composition differed on tallow tree compared to all three native trees. It supported an arthropod community that had relatively lower herbivore abundance but relatively more predator species compared to the native species examined. Leaves were collected to determine damage. Tallow tree experienced less mining damage than native trees. The results of this study supported the Enemies Release Hypothesis predictions that tallow tree would have low herbivore loads which may contribute to its invasive success. Moreover, a shift in the arthropod community to fewer herbivores without a reduction in predators may further limit regulation of this exotic species by herbivores in its introduced range.     

Effects of introduced ungulates on forest understory communities in northern Patagonia are modified by timing and severity of stand mortality

Natural disturbances such as fires, windstorms, floods, and herbivory often act on plant communities, affecting their structure and the abundance and composition of their species. Most research has focused on the effects of single disturbances on plant communities whereas the synergistic effects of several disturbances have received less attention. In this study, we evaluated how timing and severity of tree mortality modified plant use by introduced deer and early post-mortality successional trajectories in northern Patagonian conifer forests. We sampled understory composition and deer use in Austrocedrus chilensis (ciprés de la cordillera) forest stands undergoing varying timing and severity of forest mortality as reconstructed using dendroecological techniques. In addition, we evaluated the effect of fallen logs on plant composition and deer use of plants by monitoring areas of massive dieback where fallen logs had been removed for fire hazard reduction, and nearby control areas not subjected to such removal. Stepwise regression analyses showed that history and severity of tree mortality strongly influence plant composition and deer use of plants. For deer use (with pellet counts and browsing index as response variables), results showed a positive relationship with degree of stand mortality and a negative relationship with cover of fallen logs. Similarly, cover of unpalatable shrub species was explained by canopy mortality history, whereas cover of palatable shrub species was positively associated with severity of canopy mortality. In areas where fallen logs had been removed, pellet counts were six times higher than those in control areas. Though total shrub species cover was similar between log removal and control areas, proportion of unpalatable shrubs increased in areas where fallen logs had been removed. In conclusion, deer use of plants was strongly limited by tall fallen logs, allowing palatable species to establish and grow. Fallen log removal accelerated deer entrance and changed understory composition toward more browse-resistant and unpalatable species. These results underscore the importance of considering the dynamics (timing, severity, and extent) of fallen woody debris influencing understory herbivory and post-disturbance succession. In addition, experimental results underpin the importance of maintaining snags and large woody debris in disturbed landscapes where salvage logging is a routine procedure.     

Experimental test of the Invasional Meltdown Hypothesis: an exotic herbivore facilitates an exotic plant,but the plant does not reciprocally facilitate the herbivore

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Understory Invasion by <Emphasis Type="Italic">Acacia longifolia</Emphasis> Alters the Water Balance and Carbon Gain of a Mediterranean Pine Forest

In water-limited ecosystems, where potential evapotranspiration exceeds precipitation, it is often assumed that plant invasions will not increase total ecosystem water use, because all available water is evaporated or transpired regardless of vegetation type. However, invasion by exotic species, with high water use rates, may potentially alter ecosystem water balance by reducing water available to native species, which may in turn impact carbon assimilation and productivity of co-occurring species. Here, we document the impact of invasion by an understory exotic woody species (Acacia longifolia) in a semi-arid Mediterranean dune pine forest. To quantify the effects of this understory leguminous tree on the water use and carbon fixation rates of Pinus pinaster we compare an invaded and a non-invaded stand. A. longifolia significantly altered forest structure by increasing plant density and leaf area index in the mid-stratum of the invaded forest. A. longifolia contributed significantly to transpiration in the invaded forest (up to 42%) resulting in a slight increase in stand transpiration in the invaded relative to non-invaded forest. More importantly, both water use and carbon assimilation rates of P. pinaster were significantly reduced in the invaded relative to non-invaded stand. Therefore, this study shows that exotic plant invasions can have significant impacts on hydrological and carbon cycling even in water-limited semi-arid ecosystems through a repartitioning of water resources between the native and the invasive species.     

Forest community structure and composition following containment treatments for the fungal pathogen oak wilt

In temperate oak forests in Ohio, USA, we examined variability in forest communities within containment treatment sites for oak wilt (Bretziella fagacearum), a fungal pathogen lethal to susceptible oak species. Containment treatments included quarantine lines in soil for limiting belowground fungal spread and sanitation cutting of 1–3 mature black oak (Quercus velutina) trees within oak wilt infection patches. At 28 sites, we compared tree structure and understory plant communities across a gradient of 1- to 6-year-old treatments and reference forest (untreated and without evidence of oak wilt). While oak seedlings were abundant, oak saplings (1–10 cm in diameter) were absent. In contrast, many native understory plant community measures were highest in oak wilt treatments. Plant species richness 100 m^?2 doubled in treatments, regardless of age, compared with reference forest. Plant cover increased with treatment age, with 6-year-old treatments exhibiting 5?×?more cover than reference forest. Non-native plants averaged only a small proportion (<?0.12) of cover across treatments and reference forest. Variability in understory communities was mostly predictable using treatment age, tree canopy cover, and geographic location, as 20 of 25 understory measures had at least 72% of their variance modeled. While oak wilt treatments did not facilitate oak regeneration nor many conservation-priority species of open savanna-woodland habitats, the treatments did diversify and increase cover of native understory communities with minimal invasion of non-native plants.

     

Biogeographical contrasts to assess local and regional patterns of invasion: a case study with two reciprocally introduced exotic maple trees

Quantitative comparisons of distribution and abundance of exotic species in their native and non‐native ranges represent a first step when studying invaders. However, this approach is rarely applied 2 particularly to tree species. Using biogeographical contrasts coupled with regional dispersal surveys, we assessed whether two exotic maple tree species, Acer negundo and Acer platanoides, can be classified as invasive in the non‐native regions surveyed. We also examined the importance of biogeography in determining the degree of invasion by exotic species using this reciprocal approach. Local‐scale surveys were conducted in a total of 34 forests to compare density, relative abundance, age structure of native and introduced populations, and whether the two introduced maple species negatively affected native tree species density. Regional‐scale surveys of a total of 136 forests were then conducted to assess distribution in the introduced regions. Introduced populations of A. negundo were denser than populations measured in their native range and negatively related to native tree species density. Age structure did not differ between regions for this species. At the regional scale, this species has invaded most of the riparian corridors sampled in France. Conversely, the density of A. platanoides introduced populations was similar to that of native populations and was not related to native tree species density. Although seedling recruitment was higher away than at home, this species has invaded only 9% of the forests sampled in southern Ontario, Canada. Although reported invasive, these two exotic maple species differed in their relative demographic parameters and regional spread. Acer negundo is currently invasive in southern France while A. platanoides is not aggressively invasive in southern Ontario. Importantly, this study effectively demonstrates that biogeography through structured contrasts provide a direct means to infer invasion of exotic species.     

Exotic deer diminish post-fire resilience of native shrub communities on Santa Catalina Island, southern California

Browsing by exotic mule deer on Santa Catalina Island (SCI) off the coast of southern California may diminish the post-fire resilience of native shrublands. To assess this, deer exclosures were established following a wildfire to monitor post-fire recovery of three dominant, native shrub species (Heteromeles arbutifolia, Rhus integrifolia, and Rhamnus pirifolia). Post-fire resprout growth, mortality, and tissue water status as well as pre- and post-fire shrub density and cover were measured inside and outside of deer exclosures. We found that deer browsing significantly limited post-fire resprout growth and led to increased mortality of resprouting H. arbutifolia shrubs (88 % mortality outside compared to 11 % inside exclosures). Post-fire resprouts maintained favorable water status during the study despite drought conditions, indicating that water stress was not a proximate cause of resprout mortality. Deer browsing resulted in a >93 % reduction in canopy coverage of dominant shrub species. The dramatic reduction of native shrubs at this site may create opportunities for displacement by exotic species, resulting in eventual vegetation-type conversion. The observed link between intense browsing and post-fire shrub mortality provides much needed information concerning the environmental impact of exotic deer on SCI and illustrates the interaction between exotic herbivores and fire on an island system.     

Ecological filtering of exotic plants in an Australian sub‐alpine environment

We investigated some of the factors influencing exotic invasion of native sub‐alpine plant communities at a site in southeast Australia. Structure, floristic composition and invasibility of the plant communities and attributes of the invasive species were studied. To determine the plant characteristics correlated with invasiveness, we distinguished between roadside invaders, native community invaders and non‐invasive exotic species, and compared these groups across a range of traits including functional group, taxonomic affinity, life history, mating system and morphology. Poa grasslands and Eucalyptus‐Poa woodlands contained the largest number of exotic species, although all communities studied appeared resilient to invasion by most species. Most community invaders were broad‐leaved herbs while roadside invaders contained both herbs and a range of grass species. Over the entire study area the richness and cover of native and exotic herbaceous species were positively related, but exotic herbs were more negatively related to cover of specific functional groups (e.g. trees) than native herbs. Compared with the overall pool of exotic species, those capable of invading native plant communities were disproportionately polycarpic, Asteracean and cross‐pollinating. Our data support the hypothesis that strong ecological filtering of exotic species generates an exotic assemblage containing few dominant species and which functionally converges on the native assemblage. These findings contrast with those observed in the majority of invaded natural systems. We conclude that the invasion of closed sub‐alpine communities must be viewed in terms of the unique attributes of the invading species, the structure and composition of the invaded communities and the strong extrinsic physical and climatic factors typical of the sub‐alpine environment.     

Giant invasive Heracleum persicum: Friend or foe of plant diversity?

The impact of invasion on diversity varies widely and remains elusive. Despite the considerable attempts to understand mechanisms of biological invasion, it is largely unknown whether some communities’ characteristics promote biological invasion, or whether some inherent characteristics of invaders enable them to invade other communities. Our aims were to assess the impact of one of the massive plant invaders of Scandinavia on vascular plant species diversity, disentangle attributes of invasible and noninvasible communities, and evaluate the relationship between invasibility and genetic diversity of a dominant invader. We studied 56 pairs of Heracleum persicum Desf. ex Fisch.‐invaded and noninvaded plots from 12 locations in northern Norway. There was lower native cover, evenness, taxonomic diversity, native biomass, and species richness in the invaded plots than in the noninvaded plots. The invaded plots had nearly two native species fewer than the noninvaded plots on average. Within the invaded plots, cover of H. persicum had a strong negative effect on the native cover, evenness, and native biomass, and a positive association with the height of the native plants. Plant communities containing only native species appeared more invasible than those that included exotic species, particularly H. persicum. Genetic diversity of H. persicum was positively correlated with invasibility but not with community diversity. The invasion of a plant community by H. persicum exerts consistent negative pressure on vascular plant diversity. The lack of positive correlation between impacts and genetic diversity of H. persicum indicates that even a small founder population may cause high impact. We highlight community stability or saturation as an important determinant of invasibility. While the invasion by H. persicum may decrease susceptibility of a plant community to further invasion, it severely reduces the abundance of native species and makes them more vulnerable to competitive exclusion.     

The invasive plant species Centaurea maculosa alters arbuscular mycorrhizal fungal communities in the field

While several recent studies have described changes in microbial communities associated with exotic plant invasion, how arbuscular mycorrhizal fungi (AMF) communities respond to exotic plant invasion is not well known, despite the salient role of this group in plant interactions. Here, we use molecular methods (terminal restriction fragment length polymorphism analyses based on the large subunit of the rRNA gene) to examine AMF community structure in sites dominated by the invasive mycorrhizal forb, Centaurea maculosa Lam. (spotted knapweed), and in adjacent native grassland sites. Our results indicate that significant AMF community alteration occurs following C. maculosa invasion. Moreover, a significant reduction in the number of restriction fragment sizes was found for samples collected in C. maculosa-dominated areas, suggesting reduced AMF diversity. Extraradical hyphal lengths exhibited a significant, on average 24%, reduction in C. maculosa-versus native grass-dominated sites. As both AMF community composition and abundance were altered by C.maculosa invasion, these data are strongly suggestive of potential impacts on AMF-mediated ecosystem processes. Given that the composition of AMF communities has the potential to differentially influence different plant species, our results may have important implications for site restoration after weed invasion.     

Community-level consequences of invasion: impacts of exotic clonal plants on riparian vegetation

Biological invasions by exotic species are occurring at exceptional rates and spatial scales worldwide and are increasingly recognized as key forms of global environmental change. Despite this growing prominence, surprisingly few ecological studies have quantified the impacts of exotic taxa on the plant communities they invade, and this is especially evident in riparian ecosystems. Along the Russian River in northern California, we used both comparative and experimental studies to investigate the influence of two exotic clonal plant species—giant reed (Arundo donax) and blue periwinkle (Vinca major)—on the composition of riparian plant communities. Our results indicate that Arundo invasion was associated with significantly lower richness of native perennial plant species on stream banks and floodplains, whereas there was no relationship on gravel bars. Additional research showed that plots invaded by Arundo and Vinca, both individually and collectively, exhibited significantly lower native and exotic species richness and abundance of both established plants and seedlings than uninvaded plots. Finally, after 2 years, experimental reductions of Arundo biomass via cutting and herbicide resulted in significantly increased native plant species richness and abundances of both established plants and seedlings, while having no effects on other exotics. In summary, our results indicate that Arundo and Vinca have strongly negative effects on diverse components of a riparian plant community, which must be addressed via effective control and restoration efforts.     

Invasion of Ligustrum lucidum (Oleaceae) in the southern Yungas: Changes in habitat properties and decline in bird diversity

Ligustrum lucidum is the major exotic tree in NW Argentina montane forests (Yungas). To assess the effects of its expanding invasion on avian communities we (1) measured different habitat properties (vertical forest structure and composition, vegetation cover, light availability, air temperature, air relative humidity and soil litter depth), (2) compared bird species composition and diversity in Ligustrum-dominated and native-dominated secondary forests and (3) analyzed seasonal patterns and changes in these variables between forest types. The study was conducted during 2010–2011 wet and dry seasons, at two altitudinal zones: 500–800 and 1100–1450 masl. Compared with native forests, Ligustrum dominated forests had a more homogeneous vertical forest structure and denser canopy cover (resulting in lower understory solar radiation), significantly lower understory cover and lower litter depth. Air temperature and relative humidity did not differ between forests in either season. Solar radiation was higher in the dry season in both forest types, but litter depth showed opposite patterns between seasons depending on forest type. We recorded 59 bird species in 21 families. Bird species abundance, richness and diversity indexes were significantly lower in Ligustrum-dominated relative to native forests of similar successional age, which had almost twice as many species as the former. Avian communities differed between altitudinal zones, but the difference was stronger between Ligustrum and native-dominated forests. Avian community composition was less variable in time and space in native forests than in Ligustrum-dominated ones. Our results suggest that L. lucidum invasion generates structurally homogeneous and simpler forests that represent a less suitable habitat for a diverse avifauna. This illustrates the wide ecological changes (from habitat properties and ecosystem functioning to vertebrate community composition) that the subtropical mountain forests of Argentina are experiencing with this invasion.