Cave invertebrate assemblages differ between native and exotic leaf litter

Abstract Allochtonous leaf litter is an important source of energy and nutrients for invertebrates in cave ecosystems. A change to the quality or quantity of litter entering caves has the potential to disrupt the structure and function of cave communities. In this study, we adopted an experimental approach to examine rates of leaf litter decomposition and the invertebrate assemblages colonizing native and exotic leaf litter in limestone caves in the Jenolan Caves Karst Conservation Reserve, New South Wales, Australia. We deployed traps containing leaf litter from exotic sycamore (Acer pseudoplatanus) and radiata pine (Pinus radiata) trees and native eucalypts (Eucalyptus spp.) in twilight zones (near the cave entrance) and areas deep within the caves for 3 months. Thirty‐two invertebrate morphospecies were recorded from the litter traps, with greater richness and abundance evident in the samples from the twilight zone compared with areas deep within the cave. Sycamore litter had significantly greater richness and abundance of invertebrates compared with eucalypt and pine litter in samples from the twilight zone, but there was no difference in richness or abundance among litter samples placed deep within the cave. Relative rates of decay of the three litters were sycamore > eucalypt > pine. We discuss the potential for the higher decomposition rates and specific leaf area in sycamores to explain their higher invertebrate diversity and abundance. Our findings have important implications for the management of exotic plants and the contribution of their leaf litter to subterranean ecosystems.     

Density‐dependent effects of non‐native brown trout Salmo trutta on the species–area relationship in stream fish assemblages

The spatial scale and density‐dependent effects of non‐native brown trout Salmo trutta on species richness of fish assemblages were examined at 48 study sites in Mamachi Stream, a tributary of Chitose River, Hokkaido, Japan. The density of age ≥1 year S. trutta was high in the upstream side of the main stem of Mamachi Stream. Fish species richness increased with increasing area of study sites (habitat size), but the increasing magnitude of the species richness with area decreased with increasing age of ≥1 year S. trutta density. The relationships between age ≥1 year S. trutta, however, and presence–absence of each species seemed to be different among species. Species richness was also determined by location and physical environmental variables, i.e. it was high on the downstream side and in structurally complex environments.     

Family 34 glycosyltransferase (GT34) genes and proteins in Pinus radiata (radiata pine) and Pinus taeda (loblolly pine)

Using a functional genomics approach, four candidate genes (PtGT34A, PtGT34B, PtGT34C and PtGT34D) were identified in Pinus taeda. These genes encode CAZy family GT34 glycosyltransferases that are involved in the synthesis of cell‐wall xyloglucans and heteromannans. The full‐length coding sequences of three orthologs (PrGT34A, B and C) were isolated from a xylem‐specific cDNA library from the closely related Pinus radiata. PrGT34B is the ortholog of XXT1 and XXT2, the two main xyloglucan (1→6)‐α‐xylosyltransferases in Arabidopsis thaliana. PrGT34C is the ortholog of XXT5 in A. thaliana, which is also involved in the xylosylation of xyloglucans. PrGT34A is an ortholog of a galactosyltransferase from fenugreek (Trigonella foenum‐graecum) that is involved in galactomannan synthesis. Truncated coding sequences of the genes were cloned into plasmid vectors and expressed in a Sf9 insect cell‐culture system. The heterologous proteins were purified, and in vitro assays showed that, when incubated with UDP‐xylose and cellotetraose, cellopentaose or cellohexaose, PrGT34B showed xylosyltransferase activity, and, when incubated with UDP‐galactose and the same cello‐oligosaccharides, PrGT34B showed some galactosyltransferase activity. The ratio of xylosyltransferase to galactosyltransferase activity was 434:1. Hydrolysis of the galactosyltransferase reaction products using galactosidases showed the linkages formed were α‐linkages. Analysis of the products of PrGT34B by MALDI‐TOF MS showed that up to three xylosyl residues were transferred from UDP‐xylose to cellohexaose. The heterologous proteins PrGT34A and PrGT34C showed no detectable enzymatic activity.     

Ecological parameters of the leaf‐litter frog community of an Atlantic Rainforest area at Ilha Grande,Rio de Janeiro state,Brazil

Abstract The Atlantic Rainforest originally covered much of the Brazilian coast and is now reduced to approximately only 7% of its original area. Data on abundance distribution and microhabitat characteristics of anuran amphibians living on the forest floor leaf litter in the Atlantic Rainforest are scarce. In this study, we analysed the effect of litter depth and structure on the abundance and species richness of leaf‐litter frogs in an area of Atlantic Rainforest at Ilha Grande, Rio de Janeiro State, south‐eastern Brazil. We performed monthly samples (nocturnal and diurnal) from August 1996 to October 1997 using small (2 m × 1 m) plots. We sampled 234 plots, totalling 468 m² of forest leaf litter. We estimated leaf‐litter depth and the proportion of leaves in the plot and tested their effect on the total abundance of frogs and species richness using multiple regression analysis. We found 185 frogs from eight species: Brachycephalus (=Psyllophryne) didactylus (Izecksohn, 1971) (Brachycephalidae), Dendrophryniscus brevipollicatus Jiménez de la Espada 1871 (Bufonidae), Adenomera marmorata Steindachner 1867, Eleutherodactylus parvus (Girard 1853), Eleutherodactylus guentheri (Steindachner 1864), Eleutherodactylus binotatus (Spix 1824) and Zachaenus parvulus (Girard 1853) (Leptodactylidae), and Chiasmocleis sp. (Microhylidae). Brachycephalus didactylus was the most abundant species, with 91 individuals, whereas Dendrophryniscus brevipollicatus was the rarest, with two individuals. Mean litter depth and the proportion of leaves in the leaf litter were significantly related to frog abundance (R² = 0.17; F_2,107 = 10.779; P = 0.0001) and species richness (R² = 0.11; F_2,107 = 6.375; P = 0.002) indicating that microhabitat characteristics may affect local distribution and abundance of frogs in the forest floor.     

Landscape characteristics of non‐native pine plantations and invasions in Southern Chile

The spread of non‐native conifers into areas naturally dominated by other vegetation types is a growing problem in South America. This process results in a landscape transformation as the conifers suppress native vegetation leading to reduced biodiversity, lower water availability and altered nutrient dynamics. Previous research highlights the broad spatial extents of land cover change in parts of Chile. However, in Southern Chile, the extent of plantations and the landscape characteristics associated with plantations and ongoing pine invasions are poorly understood. Here, we characterised non‐native pine land cover within one Landsat scene (World Reference System 2 Path 232/Row 92; ~34 000 km²) in Southern Chile. We created training data based on historical high‐resolution imagery, derived land cover predictors from time series of Landsat observations and used a Random Forest classifier to map the distribution of non‐native pines. The overall classification accuracy was 88%, and the accuracy of the non‐native pine class exceeded 90%. Although 71% of non‐native pine patches were within 500 m of other non‐native pine patches, isolated non‐native pine patches were found to occur up to 55 km from the nearest neighbour. These distant plantations could exacerbate invasion risk by creating propagule sources for novel invasion fronts. In relation to landscape characteristics, non‐native pines were found to be more likely to occur in low slope and mid‐elevation areas. Because most of the study area is native forest, most non‐native pine patches border native forest. However, non‐native pine patches were almost three times more likely than random patches to border grass/agriculture. This suggests that grasslands and disturbed sites, which have low resistance to non‐native pine invasion, are disproportionately exposed to pine propagules. Our results indicate that non‐native pine plantations are extensive across Southern Chile, and well poised to cause future invasion.     

Dicyclohexylamine uptake and effects on polyamine content in cultured cotyledons of radiata pine

Excised cotyledons of radiata pine ( Pinus radiata D. Don) were cultured in the presence or absence of benzyladenine and two concentrations of dicyclohexylamine, a potent inhibitor of spermidine synthesis in animals and bacteria. Cellular levels of the drug and of putrescine, spermidine and spermine were determined by dansylation followed by thin layer chromatography. The rate of uptake of the drug was rapid during the first 6 h of incubation and then diminished; nevertheless, its cellular content increased with time in culture and uptake was greater if more was present in the medium. Cotyledons cultured on benzyladenine-containing media accumulated more dicyclohexylamine than on benzyladenine-free media and this resulted in toxic side-effects. The drug had no significant effect on the elongation growth occurring in the absence of the hormone. In cotyledons treated with the drug, spermidine levels fell to zero after a 5- to 10-day exposure. Putrescine increased transiently at 24 h and then declined significantly. Spermine levels also declined to 11% of control values by day 5 and remained low throughout. Except for a marked decrease in all three polyamines during the first 24 h of culture, all control cotyledons maintained significantly higher polyamine levels than dicyclohexylamine-treated ones.     

Soil–atmosphere exchange of greenhouse gases in a Eucalyptus marginata woodland, a clover-grass pasture, and Pinus radiata and Eucalyptus globulus plantations

Soils provide the largest terrestrial carbon store, the largest atmospheric CO₂ source, the largest terrestrial N₂O source and the largest terrestrial CH₄ sink, as mediated through root and soil microbial processes. A change in land use or management can alter these soil processes such that net greenhouse gas exchange may increase or decrease. We measured soil–atmosphere exchange of CO₂, N₂O and CH₄ in four adjacent land‐use systems (native eucalypt woodland, clover‐grass pasture, Pinus radiata and Eucalyptus globulus plantation) for short, but continuous, periods between October 2005 and June 2006 using an automated trace gas measurement system near Albany in southwest Western Australia. Mean N₂O emission in the pasture was 26.6 μg N m⁻² h⁻¹, significantly greater than in the natural and managed forests (< 2.0 μg N m⁻² h⁻¹). N₂O emission from pasture soil increased after rainfall events (up to 100 μg N m⁻² h⁻¹) and as soil water content increased into winter, whereas no soil water response was detected in the forest systems. Gross nitrification through ¹⁵N isotope dilution in all land‐use systems was small at water holding capacity < 30%, and under optimum soil water conditions gross nitrification ranged between < 0.1 and 1.0 mg N kg⁻¹ h⁻¹, being least in the native woodland/eucalypt plantation < pine plantation < pasture. Forest soils were a constant CH₄ sink, up to −20 μg C m⁻² h⁻¹ in the native woodland. Pasture soil was an occasional CH₄ source, but weak CH₄ sink overall (−3 μg C m⁻² h⁻¹). There were no strong correlations (R < 0.4) between CH₄ flux and soil moisture or temperature. Soil CO₂ emissions (35–55 mg C m⁻² h⁻¹) correlated with soil water content (R < 0.5) in all but the E. globulus plantation. Soil N₂O emissions from improved pastures can be considerable and comparable with intensively managed, irrigated and fertilised dairy pastures. In all land uses, soil N₂O emissions exceeded soil CH₄ uptake on a carbon dioxide equivalent basis. Overall, afforestation of improved pastures (i) decreases soil N₂O emissions and (ii) increases soil CH₄ uptake.     

岷江上游本地种油松和外来种辐射松造林对土壤磷的影响

通过测定岷江上游16年生本地种油松和外来种辐射松人工林下不同土壤层次中各形态磷素含量以及磷酸酶活力,阐述两种林分对土壤磷素含量及其分布的影响.结果表明,在各个土层中,两种林分下的土壤含水量、pH值、有机碳的含量以及微生物生物量磷均无显著差异.在0～20cm土层中,油松林与辐射松林土壤全磷、Al-P、Fe-P、Ca-P含量以及磷酸酶活力均无显著差异,而油松林土壤有效磷和有机磷显著高于辐射松林;在20～40cm土壤中,油松林土壤全磷、有效磷、有机磷、Al-P、Fe-P含量与辐射松林差异不显著,油松林土壤Ca-P含量、酸性磷酸酶和中性磷酸酶活力显著高于辐射松林;在40～60cm土壤中,油松林土壤除中性磷酸酶活力与辐射松林的差异不显著外,其余各形态磷素含量和酸性磷酸酶活力变化与20～40cm土壤中的一致.此外,随土壤深度的增加,两种人工林土壤各形态磷素含量、磷酸酶活力都呈降低的趋势.单从土壤磷的状况看,油松林土壤中磷素含量高于辐射松林.     

Soil and solution chemistry under pasture and radiata pine in New Zealand

The conversion of hill country pasture to exotic forest plantations is occurring rapidly (70,000 ha yr^–1) in New Zealand. Impacts of this land-use change on soil properties, soil fertility, and water quality are only beginning to be investigated. This study examines the effects of radiata pine (Pinus radiata) on soil and soil solution chemistry, in a region of low atmospheric pollution, 20 years after plantation establishment, assuming that the pasture and pine research sites had comparable soil properties before planting pine. The primary effects of conversion on soil chemistry were a decrease of organic carbon in the mineral soil that was balanced by an accumulation of the surface litter layer, a decrease in soil N, soil acidification, and increased pools of exchangeable Mg, K, and Na. Soil solution studies revealed a large input of sea salts by enhanced canopy capture of sea salts that contributed to much larger solute concentrations and elemental fluxes in the pine soil. Sea salts appear to accumulate in the micropores of pine soil during the dry summer period and are slowly released to macropore flow during the rainy season. This results in a progressive decrease in solute concentrations over the period of active leaching. While chloride originating from sea salt deposition was the dominant anion in the pine soil, bicarbonate originating from root and microbial respiration was the dominant anion in the pasture soil. Carbon dioxide concentrations in the soil atmosphere were 12.5-fold greater in the pasture soil than in the pine soil due to greater rates of root and microbial respiration and to slower diffusion rates resulting from wetter soil conditions in the pasture. Although elemental fluxes from the upper 20 cm of the soil profile were substantially greater in the pine soil, these losses were compensated for by increased elemental inputs resulting from nutrient cycling and enhanced canopy capture of sea salts.     

Relative contribution of neutral and deterministic processes in shaping fruit‐feeding butterfly assemblages in Afrotropical forests

The unified neutral theory of biodiversity and biogeography has gained the status of a quantitative null model for explaining patterns in ecological (meta)communities. The theory assumes that individuals of trophically similar species are functionally equivalent. We empirically evaluate the relative contribution of neutral and deterministic processes in shaping fruit‐feeding butterfly assemblages in three tropical forests in Africa, using both direct (confronting the neutral model with species abundance data) and indirect approaches (testing the predictions of neutral theory using data other than species abundance distributions). Abundance data were obtained by sampling butterflies using banana baited traps set at the forest canopy and understorey strata. Our results indicate a clear consistency in the kind of species or species groups observed at either the canopy or understorey in the three studied communities. Furthermore, we found significant correlation between some flight‐related morphological traits and species abundance at the forest canopy, but not at the understorey. Neutral theory's contribution to explaining our data lies largely in identifying dispersal limitation as a key process regulating fruit‐feeding butterfly community structure. Our study illustrates that using species abundance data alone in evaluating neutral theory can be informative, but is insufficient. Species‐level information such as habitat preference, host plants, geographical distribution, and phylogeny is essential in elucidating the processes that regulate biodiversity community structures and patterns.     

Root development and configuration in intensively managed radiata pine plantations

Summary In south-east Australia, where radiata pine (Pinus radiata D. Don) is grown on sandy soils low in nutrients and short of water, early establishment, and rapid growth to canopy closure lead to increased productivity. At this stage demands for nutrients and water are high, and trees respond vigorously to silvicultural inputs.For several months after transplanting in winter roots are confined within a narrow planting wedge, low temperature restricts new root growth and slows recovery from water stress in plants. From spring, depending upon the configuration and vigour of the roots transplanted, lateral roots extend radially throughout the soil.Although there were small decreases in concentration of roots radially from the stems of very young trees, such spatial differences disappeared between ages 2 and 3, so that rooting density was independent of distance from the stem. The pattern of vertical distribution of lateral roots was not influenced by age and 80–90% of the lateral roots were within the top 30 cm soil. Roots developed rapidly as the trees grew towards canopy closure, but in general the rooting densities of these pines are among the lowest reported for plants. In rapidly growing trees approaching canopy closure, the secondary thickening of the lateral roots was sufficient to double the weight of roots without altering root length.Knowledge about root growth and root configuration during the early phase of plantation development will assist management decisions where intensive silviculture is practiced, and hence ensure the most efficient use of nutrients and water.     

Assessment and monitoring of damage from insects in Australian eucalypt forests and commercial plantations

Abstract   This paper presents a review of recent developments in the assessment and monitoring of health in Australian eucalypt forests and plantations of pine and eucalypt species, with an emphasis on damage caused by herbivorous insects. The diverse range of interests and priorities amongst Australian stakeholders of native forests and plantations influences the scale, resolution and accuracy of results sought, and this in turn influences how the assessment data are collected, analysed and reported. The authors discuss sampling systems that include extensive ground-based surveys, permanent plots and airborne technologies being developed in Australia. In all cases, there is an appreciation that the assessment protocols should be objective, repeatable and cost effective. Significant progress has been made in the application of digital, remotely sensed imagery to detect and classify damaged forest canopies. The success of this approach depends, in part, on a sound understanding of the progression of symptoms at the leaf, tree crown and stand scale, especially those symptoms that influence spectral reflectance behaviour.     

Land use diversification may mitigate on-site land use impacts on mammal populations and assemblages

Land use is a major cause of biodiversity decline worldwide. Agricultural and forestry diversification measures, such as the inclusion of natural elements or diversified crop types, may reduce impacts on biodiversity. However, the extent to which such measures may compensate for the negative impacts of land use remains unknown. To fill that gap, we synthesised data from 99 studies that recorded mammal populations or assemblages in natural reference sites and in cropland and forest plantations, with or without diversification measures. We quantified the responses to diversification measures based on individual species abundance, species richness and assemblage intactness as quantified by the mean species abundance indicator. In cropland with natural elements, mammal species abundance and richness were, on average, similar to natural conditions, while in cropland without natural elements they were reduced by 28% and 34%, respectively. We found that mammal species richness was comparable between diversified forest plantations and natural reference sites, and 32% lower in plantations without natural elements. In both cropland and plantations, assemblage intactness was reduced compared with natural reference conditions, but the reduction was smaller if diversification measures were in place. In addition, we found that responses to land use were modified by species traits and environmental context. While habitat specialist populations were reduced in cropland without diversification and in forest plantations, habitat generalists benefited. Furthermore, assemblages were impacted more by land use in tropical regions and landscapes containing a larger share of (semi)natural habitat compared with temperate regions and more converted landscapes. Given that mammal assemblage intactness is reduced also when diversification measures are in place, special attention should be directed to species that suffer from land use impacts. That said, our results suggest potential for reconciling land use and mammal conservation, provided that the diversification measures do not compromise yield.     

Non‐linearities in bird responses across urbanization gradients: A meta‐analysis

Urbanization is one of the most extreme forms of environmental alteration, posing a major threat to biodiversity. We studied the effects of urbanization on avian communities via a systematic review using hierarchical and categorical meta‐analyses. Altogether, we found 42 observations from 37 case studies for species richness and 23 observations from 20 case studies for abundance. Urbanization had an overall strong negative effect on bird species richness, whereas abundance increased marginally with urbanization. There was no evidence that city size played a role in influencing the relationship between urbanization and either species richness or abundance. Studies that examined long gradients (i.e. from urban to rural) were more likely to detect negative urbanization effects on species richness than studies that considered short gradients (i.e. urban vs. suburban or urban vs. rural areas). In contrast, we found little evidence that the effect of urbanization on abundance was influenced by gradient length. Effects of urbanization on species richness were more negative for studies including public green spaces (parks and other amenity areas) in the sampled landscapes. In contrast, studies performed solely in the urban matrix (i.e. no green spaces) revealed a strong positive effect on bird abundance. When performing subset analyses on urban–suburban, suburban–rural and suburban–natural comparisons, species richness decreased from natural to urban areas, but with a stronger decrease at the urban–suburban interface, whereas bird abundance showed a clear intermediate peak along the urban–rural gradient although abundance in natural areas was comparable to that in suburban areas. This suggests that species loss happens especially at the urban–suburban interface, and that the highest abundances occur in suburban areas compared to urban or rural areas. Thus, our study shows the importance of suburban areas, where the majority of birds occur with fairly high species richness.     

Equal diversity in disparate species assemblages: a comparison of native and exotic woodlands in California

The species richness of ecosystems can remain stable over time, despite changes in species composition and changes in the dominant plant species. While this pattern of stability is known to occur temporally, it has been examined poorly in a spatial context. To examine this spatially, the species richness, diversity and composition of native woodlands (of oak and bay trees) and exotic woodlands (of eucalypt trees) were compared in California. Species richness was nearly identical for understorey plants, leaf‐litter invertebrates, amphibians and birds; only rodents had significantly fewer species in eucalypt sites. Species diversity patterns (using the Shannon–Wiener Index) were qualitatively identical to those for species richness, except for leaf‐litter invertebrates, which were significantly more diverse in eucalypt sites during the spring. Species composition was different between sites, as evidenced by a principal components analysis, coefficients of similarity, and the relatively few species shared between native and eucalypt sites. Thus, the consistency in richness and diversity observed for most groups, in most seasons, occurred despite significant differences in species composition. These results are consistent with previous demonstrations of temporal stability, suggesting that species richness may often be stable, both temporally and spatially, despite changes in composition and regardless of the dominant vegetation.     

Degradation of native and exotic riparian plant leaf litter in a floodplain pond

1. A litter‐bag experiment was undertaken in a pond on the margins of a large temperate floodplain in south‐western France to assess the potential influence of the replacement of native by exotic riparian species on organic matter degradation. We determined initial litter chemical composition, breakdown rates and the invertebrate assemblages associated with the litter for five pairs of native dominant and exotic invasive species co‐occurring at different stages along a successional gradient. 2. Litter chemical composition, breakdown rates and abundance and diversity of detritivorous invertebrates were similar for the exotic and native species overall. No overall changes in organic matter degradation can thus be predicted from the replacement of dominant natives by exotic invasives. Breakdown rates were primarily driven by the C/N ratio. 3. One invasive species (Buddleja davidii) showed significantly higher breakdown rates than its native counterpart (Populus nigra), resulting in the disappearance of leaf litter 6 months prior to the next litterfall. In some cases, therefore, invasion by exotic species may result in discontinuity of resource supply for decomposers.     

The importance of sward architectural complexity in structuring predatory and phytophagous invertebrate assemblages

Abstract.  1. Although the importance of plant community assemblages in structuring invertebrate assemblages is well known, the role that architectural complexity plays is less well understood. In particular, direct empirical data for a range of invertebrate taxa showing how functional groups respond to plant architecture is largely absent from the literature.
2. The significance of sward architectural complexity in determining the species richness of predatory and phytophagous functional groups of spiders, beetles, and true bugs, sampled from 135 field margin plots over 2 years was tested. The present study compares the relative importance of sward architectural complexity to that of plant community assemblage.
3. Sward architectural complexity was found to be a determinant of species richness for all phytophagous and predatory functional groups. When individual species responses were investigated, 62.5% of the spider and beetle species, and 50.0% of the true bugs responded to sward architectural complexity.
4. Interactions between sward architectural complexity and plant community assemblage indicate that the number of invertebrate species supported by the plant community alone could be increased by modification of sward architecture. Management practices could therefore play a key role in diversifying the architectural structure of existing floral assemblages for the benefit of invertebrate assemblages.
5. The contrasting effects of sward architecture on invertebrate functional groups characterised by either direct (phytophagous species) or indirect (predatory species) dependence on plant communities is discussed. It is suggested that for phytophagous taxa, plant community assemblage alone is likely to be insufficient to ensure successful species colonisation or persistence without appropriate development of sward architecture.     

Ectomycorrhizal fungi associated with ponderosa pine and Douglas-fir: a comparison of species richness in native western North American forests and Patagonian plantations from Argentina

The putative ectomycorrhizal fungal species registered from sporocarps associated with ponderosa pine and Douglas-fir forests in their natural range distribution (i.e., western Canada, USA, and Mexico) and from plantations in south Argentina and other parts of the world are listed. One hundred and fifty seven taxa are reported for native ponderosa pine forests and 514 taxa for native Douglas-fir forests based on available literature and databases. A small group of genera comprises a high proportion of the species richness for native Douglas-fir (i.e., Cortinarius, Inocybe, and Russula), whereas in native ponderosa pine, the species richness is more evenly distributed among several genera. The comparison between ectomycorrhizal species richness associated with both trees in native forests and in Patagonia (Argentina) shows far fewer species in the latter, with 18 taxa for the ponderosa pine and 15 for the Douglas-fir. Epigeous species richness is clearly dominant in native Douglas-fir, whereas a more balanced relation epigeous/hypogeous richness is observed for native ponderosa pine; a similar trend was observed for Patagonian plantations. Most fungi in Patagonian Douglas-fir plantations have not been recorded in plantations elsewhere, except Suillus lakei and Thelephora terrestris, and only 56% of the fungal taxa recorded in Douglas-fir plantations around the world are known from native forests, the other taxa being new associations for this host, suggesting that new tree + ectomycorrhizal fungal taxa associations are favored in artificial situations as plantations.