Predation behaviour of land planarians

Predatory behaviour of land planarians is seldom observed or reported. Aspects reported are (1) finding prey; (2) attack behaviour; (3) capture using adhesive mucus, pharyngeal action, poisonous secretions, physical embrace; (4) feeding by extension of pharynx, releasing copious digestive fluid. The species Bipalium kewense, B. adventitium and B. pennsylvanicum attack earthworms, immobilizing them by physical holding, digesting by pharyngeal secretions and then ingesting the treated tissue. Group attacks on giant African land snails involving chemotactic tracking, occur in Platydemus manokwari and Endeavouria septemlineata. Specialized capture methods are used by some species; Rhynchodemus sylvaticus uses an expanded cephalic hood to capture small insects and in Africa, termites are captured by the elongated anterior of Microplana termitophaga as planarians wait within the colony air shaft openings to ensnare the workers in sticky mucus. The result of extensive predation by land planarians may seriously reduce the prey, e.g., providing effective population control of giant land snails by introduced Platydemus manokwari, or causing serious depletion of desirable earthworm populations by the exotic Artioposthia triangulata in North Ireland.     

Apparent competition: an impact of exotic shrub invasion on tree regeneration

Invasion of habitats by exotic shrubs is often associated with a decrease in the abundance of native species, particularly trees. This is typically interpreted as evidence for direct resource competition between the invader and native species. However, this may also reflect indirect impacts of the exotic shrubs through harboring high densities of seed predators––known as apparent competition. Here I present data from separate seed predation experiments conducted with two shrub species exotic to North America; Rosa multiflora, an invader of abandoned agricultural land, and Lonicera maackii, an invader of disturbed or secondary forest habitats. Both experiments showed significantly greater risks of seed predation for tree seeds located under shrub canopies when compared to open microhabitats within the same site. These results indicate the potential importance of indirect impacts of exotic species invasions on native biota in addition to the direct impacts that are typically the focus of research.     

Non-native plantation forests as alternative habitat for native forest beetles in a heavily modified landscape

The once extensive native forests of New Zealand’s central North Island are heavily fragmented, and the scattered remnants are now surrounded by a matrix of exotic pastoral grasslands and Pinus radiata plantation forests. The importance of these exotic habitats for native biodiversity is poorly understood. This study examines the utilisation of exotic plantation forests by native beetles in a heavily modified landscape. The diversity of selected beetle taxa was compared at multiple distances across edge gradients between each of the six possible combinations of adjacent pastoral, plantation, clearfell and native forest land-use types. Estimated species richness (Michaelis–Menten) was greater in production habitats than native forest; however this was largely due to the absence of exotic species in native forest. Beetle relative abundance was highest in clearfell-harvested areas, mainly due to colonisation by open-habitat, disturbance-adapted species. More importantly, though, of all the non-native habitats sampled, beetle species composition in mature P. radiata was most similar to native forest. Understanding the influence of key environmental factors and stand level management is important for enhancing biodiversity values within the landscape. Native habitat proximity was the most significant environmental correlate of beetle community composition, highlighting the importance of retaining native remnants within plantation landscapes. The proportion of exotic beetles was consistently low in mature plantation stands, however it increased in pasture sites at increasing distances from native forest. These results suggest that exotic plantation forests may provide important alternative habitat for native forest beetles in landscapes with a low proportion of native forest cover.     

Water flux and canopy conductance of natural versus planted forests in Patagonia, South America

In NW Patagonia, South America, natural shrublands and mixed forests of short Nothofagus antarctica (G. Forst.) Oerst. trees are currently being replaced by plantations with Pseudotsuga menziesii (Mirb) Franco. This land use change is controversial because the region is prone to drought, and replacement of native vegetation by planted forests may increase vegetation water use. The goal of this study was to examine the physiological differences, especially the response of water flux and canopy conductance to microclimate, that lead to greater water use by exotic trees compared to native trees. Meteorological variables and sapflow density of P. menziesii and four native woody species were measured in the growing season 2005–2006. Canopy conductance (gc) was estimated for both the exotic (monoculture) and native (multi-species) systems, including the individual contributions of each species of the native forest. Sapflow density, stand-level transpiration and gc were related to leaf-to-air vapor pressure difference (VPD). All native species had different magnitudes and diurnal patterns of sapflow density compared to P. menziesii, which could be explained by the different gc responses to VPD. Stomatal sensitivity to VPD suggested that all native species have a stronger stomatal control of leaf water potential and transpiration due to hydraulic limitations compared to P. menziesii. In conclusion, differences in water use between a P. menziesii plantation and a contiguous native mixed forest of similar basal area could be explained by different gc responses to VPD between species (higher sensitivity in the native species), in addition to particular characteristics of the native forest structure.     

Carnivore mollusks as natural enemies of invasive land flatworms

There are several records of the carnivorous behaviour of land flatworms, considered to be top‐predators in their micro‐habitats, by preying upon various species of invertebrates. However, there is little knowledge of predators on land‐flatworms. The possible impact of invasive land flatworms on prey populations has caused widespread concern, when considering their predatory behaviour, combined with recent human influence on the distribution of certain species. This work is the first record of predation on land flatworms by a carnivorous snail. Various‐sized land flatworms of 10 native species of the subfamily Geoplaninae, as well as the exotic species Bipalium kewense (subfamily Bipaliinae), were offered to Rectartemon depressus (Gastropoda, Streptaxidae), which accepted all. The predator also fed on the snail Bradybaena similaris. The snails were maintained in laboratory for an average period of 12 months based on a mixed diet of flatworms and B. similaris, suggesting that the snail is a polyphagous predator. Because certain land‐flatworm species have been described as invasive species which may have a potential impact on prey populations in native and man‐made ecosystems, it is proposed that carnivorous snails of other native species, as potential predators of flatworms, should be tested for possible use in biological control programmes of these invasive planarians.     

Forest fragment size and nutrient availability: complex responses of mycorrhizal fungi in native–exotic hosts

In the past few decades, it has been widely accepted that forest loss due to human actions alter the interactions between organisms. We studied the relationship between forest fragment size and arbuscular mycorrhizal fungi (AMF) and dark septate endophytes (DSE) colonization, and the AMF spore communities in the rhizosphere of two congeneric Euphorbia species (native and exotic/invasive). We hypothesized that these fungal variables will differ with fragment size and species status, and predicted that (a) AMF and DSE colonization together with AMF spore abundance and diversity would be positively related to forest fragment size; (b) these relationships will differ between the exotic and the native species; and (c) there will be a negative relationship between forest fragment size and the availability of soil nutrients (NH₄ ⁺, NO₃ ⁻, and phosphorus). This study was performed in the eight randomly selected forest fragments (0.86–1000 ha), immersed in an agricultural matrix from the Chaquean region in central Argentina. AMF root colonization in the native and exotic species was similar, and was positively related with forest fragment size. Likewise, AMF spore diversity and spore abundance were higher in the larger fragments. While DSE root colonization in the native host was positively related with forest fragment size, DSE colonization in the exotic host showed no relationship. Soil nutrients contents were negatively related with forest fragment size. In addition, NH₄ ⁺ and NO₃ ⁻ were negatively correlated with AMF spores abundance and root colonization and with DSE colonization in the native species. The results observed in this study show how habitat fragmentation might affect the interaction between key soil components, such as rhizospheric plant-fungal symbiosis and nutrient availability. These environmental changes may have important consequences on plant community composition and nutrient dynamics in this fragmented landscape.     

Predicting effects of large‐scale reforestation on native and exotic birds

Aim

Ecological restoration is critical for recovering biodiversity and ecosystem services, yet designing interventions to achieve particular outcomes remains fraught with challenges. In the extensive regions where non‐native species are firmly established, it is unlikely that historical conditions can be fully reinstated. To what degree, and how rapidly, can human‐dominated areas be shifted via restoration into regimes that benefit target species, communities or processes?

Location

We explore this question in a >20‐year‐old reforestation effort underway at Hakalau Forest National Wildlife Refuge in montane Hawaii. This large‐scale planting of Acacia koa trees is designed to secure populations of globally threatened bird species by transitioning the site rapidly from pasture to native forest.

Methods

We surveyed all forest birds in multiple corridors of young planted trees, remnant corridors of mature trees along gulches and at sites within mature forest. Using a Bayesian hierarchical approach, we identified which factors (distance from forest, habitat type and surrounding tree cover) had the most important influence on native and exotic bird abundance in the reforestation area.

Results

We found that 90% of native and exotic bird species responded quickly, occupying corridors of native trees approximately a decade after planting. However, native and exotic forest birds responded to markedly different characteristics of the reforested area. Native bird abundance was strongly predicted by proximity to mature forest and remnant corridors; conversely, exotic bird abundance was best predicted by overall tree cover throughout the area reforested.

Main conclusions

Our results demonstrate that large‐scale tree planting in corridors adjacent to mature forest can catalyse rapid recovery (both increased abundance and expanded distribution) of forest birds and that it is possible to design reforestation to benefit native species in novel ecosystems.

     

Dominance of an invasive earthworm in native and non-native grassland ecosystems

More attention is currently being focused on earthworm invasions; however, in many ecosystems the relative abundance of native and invasive earthworm species is unknown. We characterized earthworm populations of two grassland types within the Palouse region: native prairie remnants and Conservation Reserve Program (CRP) set asides planted with exotic grasses. The earthworm community in both grassland types was completely dominated by the exotic-invasive Aporrectodea trapezoides. Only one individual of a native species, Driloleirus americanus (the giant Palouse earthworm), was found in a prairie remnant. No differences were found between prairie remnants and CRP sites for mean earthworm density (24–106 individuals m⁻²) or fresh weight (12–45 g m⁻²). Our results suggest that the combined effects of land-use change, habitat fragmentation and competitive interactions have resulted in the decimation of native earthworm populations and dominance of invasive earthworms in native and non-native grasslands of the Palouse region.     

Native and exotic plants of fragments of sagebrush steppe produced by geomorphic processes versus land use

Habitat fragmentation and invasion by exotic species are regarded as major threats to the biodiversity of many ecosystems. We surveyed the plant communities of two types of remnant sagebrush-steppe fragments from nearby areas on the Snake River Plain of southeastern Idaho, USA. One type resulted from land use (conversion to dryland agriculture; hereafter AG Islands) and the other from geomorphic processes (Holocene volcanism; hereafter kipukas). We assessed two predictions for the variation in native plant species richness of these fragments, using structural equation models (SEM). First, we predicted that the species richness of native plants would follow the MacArthur–Wilson (M–W) hypothesis of island biogeography, as often is expected for the communities of habitat fragments. Second, we predicted a negative relationship between native and exotic plants, as would be expected if exotic plants are decreasing the diversity of native plants. Finally, we assessed whether exotic species were more strongly associated with the fragments embedded in the agricultural landscape, as would be expected if agriculture had facilitated the introduction and naturalization of non-native species, and whether the communities of the two types of fragments were distinct. Species richness of native plants was not strongly correlated with M–W characteristics for either the AG Islands or the **kipukas. The AG Islands had more species and higher cover of exotics than the kipukas, and exotic plants were good predictors of native plant species richness. Our results support the hypothesis that proximity to agriculture can increase the diversity and abundance of exotic plants in native habitat. In combination with other information, the results also suggest that agriculture and exotic species have caused loss of native diversity and reorganization of the sagebrush-steppe plant community.     

Linking dominant Hawaiian tree species to understory development in recovering pastures via impacts on soils and litter

Large areas of tropical forest have been cleared and planted with exotic grass species for use as cattle pasture. These often remain persistent grasslands after grazer removal, which is problematic for restoring native forest communities. It is often hoped that remnant and/or planted trees can jump‐start forest succession; however, there is little mechanistic information on how different canopy species affect community trajectories. To investigate this, I surveyed understory communities, exotic grass biomass, standing litter pools, and soil properties under two dominant canopy trees—Metrosideros polymorpha (‘ōhi‘a) and Acacia koa (koa)—in recovering Hawaiian forests. I then used structural equation models (SEMs) to elucidate direct and indirect effects of trees on native understory. Native understory communities developed under ‘ōhi‘a, which had larger standing litter pools, lower soil nitrogen, and lower exotic grass biomass than koa. This pattern was variable, potentially due to historical site differences and/or distance to intact forest. Koa, in contrast, showed little understory development. Instead, data suggest that increased soil nitrogen under koa leads to high grass biomass that stalls native recruitment. SEMs suggested that indirect effects of trees via litter and soils were as or more important than direct effects for determining native cover. It is suggested that diverse plantings which incorporate species that have high carbon to nitrogen ratios may help ameliorate the negative indirect effects of koa on natural understory regeneration.     

Fire enhances the ‘competition-free’ space of an invader shrub: <Emphasis Type="Italic">Rosa rubiginosa</Emphasis> in northwestern Patagonia

Fire can influence reproductive phenology of plants, enhancing the reproductive rate of many species. Disturbances such as fire can promote the proliferation of exotic species in native plant communities. In this study we analyze the effect of fire on reproductive phenology in three native species (a shrub: Berberis buxifolia and two small trees: Maytenus boaria and Schinus patagonicus) and in an exotic shrub (Rosa rubiginosa). Flowering and fruiting phenology was monitored in neighbouring burned and unburned forests. The shrubs flowered and fruited in both sites, but the small trees did so only in the unburned site. There is no overlapping in the flowering and fruiting phenophases between the natives and the exotic species. Therefore, they do not compete in resource offering to pollinators and seed dispersers. Consequently, R. rubiginosa has a ‘competition-free’ space enhanced by fire, from the reproductive phenology perspective.     

Plantations of exotic tree species in Britain: irrelevant for biodiversity or novel habitat for native species?

Novel or emergent ecosystems arising from human action present both threats and opportunities for biodiversity. It has been suggested that exotic species can “facilitate” or “inhibit” native biodiversity through habitat modification. In Britain, there is a discussion over the contribution to biodiversity of plantations of exotic conifer species as these are commonly thought to have little relevancy as a habitat for native biodiversity. To address this we compared the species richness of a range of different taxonomic groups (lichens, bryophytes, fungi, vascular plants, invertebrates and songbirds) in exotic and native forest stands of differing structural stages in northern and southern Britain. In terms of overall native species-richness there was no significant difference between the exotic and the native stands. In the north, six species groups showed higher values in the exotic Sitka spruce (Picea sitchensis) stands with the remaining six showing higher values in the native Scots pine (Pinus sylvestris) stands. Most notably, lichen species richness was much lower in the exotic stands compared to the native stands, whereas bryophyte and fungal species richness was proportionately higher in the exotic stands. In the south, five species groups (all invertebrate taxa) showed higher species richness in exotic Norway spruce (Picea abies) stands compared to native oak (Quercus robur) stands. Five species groups had higher species-richness in the oak stands, in particular lichens and fungi. It is concluded that emergent ecosystems of exotic conifer species are not irrelevant to biodiversity. Where already well-established they can provide habitat for native species particularly if native woodland is scarce and biodiversity restoration is an immediate priority.     

Relevance of exotic pine plantations as a surrogate habitat for ground beetles (Carabidae) where native forest is rare

Plantation forests are of increasing importance worldwide for wood and fibre production, and in some areas they are the only forest cover. Here we investigate the potential role of exotic plantations in supporting native forest-dwelling carabid beetles in regions that have experienced extensive deforestation. On the Canterbury Plains of New Zealand, more than 99% of the previous native forest cover has been lost, and today exotic pine (Pinus radiata) plantations are the only forest habitat of substantial area. Carabids were caught with pitfall traps in native kanuka (Kunzea ericoides) forest remnants and in a neighbouring pine plantation, grassland and gorse (Ulex europaeus) shrubland. A total of 2,700 individuals were caught, with significantly greater abundance in traps in young pine, grassland and gorse habitats than in kanuka and older pine. Rarefied species richness was greatest in kanuka, a habitat that supported two forest specialist species not present in other habitat types. A critically endangered species was found only in the exotic plantation forest, which also acts as a surrogate habitat for most carabids associated with kanuka forest. The few remaining native forest patches are of critical importance to conservation on the Canterbury Plains, but in the absence of larger native forest areas plantation forests are more valuable for carabid conservation than the exotic grassland that dominates the region.     

Exotic shrubs as ephemeral ecological traps for nesting birds

Certain exotic plants may increase risk of nest predation, and, in this way, may act as ecological traps. We hypothesized that the greater vulnerability to predation was a consequence of either (1) reduced nest height due to architectural differences among plant species or (2) seasonal changes in the distribution of nests among forest strata. To test this, we examined temporal variation in nest survival of 888 nests of Northern Cardinal (Cardinalis cardinalis) in native substrates and two exotic shrubs (Lonicera maackii and Rosa multiflora) in Ohio, USA, 2001–2006. We evaluated evidence for an ecological trap by monitoring the annual reproductive productivity of 245 breeding pairs of cardinals. Only nests in Rosa experienced relatively constant survival rates across the season, whereas probability of survival increased over the season for nests in other substrates. Interestingly, the relative vulnerability of nests in different substrates varied across the season. Most strikingly, nests in Lonicera in early spring showed the lowest survival rates but exceeded survival rates of nests in native substrates late in the season. Nest height failed to explain seasonal changes in nest survival, as only nests in native plants significantly increased in height as the season progressed. Rather, predation risk seemed to be a function of the proportion of nests within each substrate, as illustrated by the decreased predation in Lonicera as the relative proportion of nests in native substrates increased. The patterns of temporal variation in predation risk that we detected show that impacts of Lonicera are not a function of plant architecture alone and may be related to leaf phenology, changes in nest density, nest site location, and/or nest synchrony. Examination of the reproductive productivity of cardinals showed that pairs that made their first nest attempt in Lonicera fledged 20% fewer cardinal young than birds that began the season using other substrates. Thus, we suggest that exotic plants may represent an ephemeral ecological trap for certain nesting birds, where negative effects persist only during certain periods.     

Restoration of mangrove plantations and colonisation by native species in Leizhou bay,South China

To examine the natural colonisation of native mangrove species into remediated exotic mangrove stands in Leizhou Bay, South China, we compared soil physical–chemical properties, community structure and recruitments of barren mangrove areas, native mangrove species plantations, and exotic mangrove species—Sonneratia apetala Buch.Ham—between plantations and natural forest. We found that severely degraded mangrove stands could not regenerate naturally without human intervention due to severely altered local environments, whereas some native species had been recruited into the 4–10 year S. apetala plantations. In the first 10 years, the exotic species S. apetala grew better than native species such as Rhizophora stylosa Griff and Kandelia candel (Linn.) Druce. The mangrove plantation gradually affected soil physical and chemical properties during its recovery. The exotic S. apetala was more competitive than native species and its plantation was able to restore soil organic matter in about 14 years. Thus, S. apetala can be considered as a pioneer species to improve degraded habitats to facilitate recolonisation by native mangrove species. However, removal to control proliferation may be needed at late stages to facilitate growth of native species. To ensure sustainability of mangroves in South China, the existing mangrove wetlands must be managed as an ecosystem, with long-term scientific monitoring program in place.     

Is <Emphasis Type="Italic">Pinus radiata</Emphasis> invading the native vegetation in central Chile? Demographic responses in a fragmented forest

Forest fragmentation facilitates the invasion of exotic species. This threat may be especially severe if forest fragments are surrounded by plantations of exotic species like Pinus radiata, an aggressive colonizer and shade-intolerant tree that has invaded successfully several native ecosystems of the southern hemisphere. In this study, we experimentally tested if the conditions of a successful seedling establishment P. radiata are fulfilled at the Coastal Maulino forest, an endemic fragmented forest of central Chile. Results demonstrated that seeds are dispersed into the native forests, however seedling establishment occurs only at the edges. We conclude that this exotic species is not invading native forests up to date. However, we suggest to conduct evaluations of seed rain and seedling establishment in the long term, in order to monitor the fate of this exotic species in fragmented native forest of Central Chile.     

Impact of cocoa farming on vegetation in an agricultural landscape in Ghana

Cocoa production occurs almost wholly within areas identified as biodiversity hotspots in West Africa and it has been noted as a major contributor to deforestation at the forest‐agriculture interface. This study investigated the impact of cocoa farming on vegetation in relation to three land‐use types of increasing cocoa production intensity from remnant native forest through shaded to unshaded cocoa farmlands in Ghana. The study used transects and forty‐two 25 m × 25 m vegetation plots. The overall noncocoa plant species richness decreased significantly (95% CI) from the remnant native forest through shaded to the unshaded cocoa farmlands. Significant differences (P ≤ 0.05) were also found in the mean density and basal area of noncocoa plants per hectare with the remnant native forest recording the highest values and the unshaded cocoa farmlands the lowest. The relative density of about 44.7% out of the 41 most abundant plant species declined in cocoa farmlands. The results of this study showed that cocoa farming could result in a drastic forest plant species loss with subsequent recruitment of nonforest species, forest plant species population decline as well as changes in the structural characteristics of the vegetation. This impact increases with increasing cocoa production intensity.     

The effects of nitrogen addition on the growth of two exotic and two native forest understory plants

Many studies have linked the spread of exotic, invasive species to high nitrogen supply, but most of this work does not distinguish between various inorganic forms and different concentrations of available nitrogen. Previous research has suggested that exotic, invasive species common in eastern deciduous forests may preferentially utilize nitrate in contrast to native species that preferentially make use of ammonium. We tested this hypothesis by comparing the growth response of two common forest invaders, Berberis thunbergii and Microstegium vimineum, and two co-occurring native species (Vaccinium pallidum and Hamamelis virginiana) to different forms of nitrogen under varying concentrations in a greenhouse experiment. Two forms of nitrogen (nitrate or ammonium) were added at different concentrations (22, 106, and 212 mg N l⁻¹) to all species. Growth response variables included survival, stem length, stem diameter, above and belowground biomass, and estimated seed production (Microstegium only). Unexpectedly the exotic species did not respond preferentially to nitrate addition. Microstegium responded most strongly to both nitrogen forms depending on the response variable. Berberis and Hamamelis surprisingly reacted similarly to nitrogen additions. As expected, Vaccinium fared poorly under most treatment conditions, but did show some growth in NH₄ ⁺ treatments. Our findings suggest species response to nitrogen addition is complex, and that exotic species do not all respond similarly to nitrogen inputs. In this study, the response of exotic and native species to available nutrients does not provide a general mechanism of invasion success.