Landscape structure affects food quality of sika deer (<Emphasis Type="Italic">Cervus nippon</Emphasis>) evidenced by fecal nitrogen levels

Evaluating the quality of wildlife habitat is essential for understanding and predicting population dynamics in heterogeneous environments. We used fecal nitrogen levels as an indicator of habitat quality of sika deer (Cervus nippon) and explored important landscape elements influencing nitrogen levels, taking deer density into account. We established 92 plots differing in deer density and landscape structure on the Boso Peninsula, central Japan, and collected fecal samples along a 1-km transect at each plot. The regression models involving two independent variables, i.e., deer density and the length of forest edge within an area of 100 or 200 m from the transect, were selected based on the Akaike’s Information Criterion (AIC). Levels of fecal nitrogen were positively correlated with the length of the forest edge and negatively correlated with population density of deer. The area of 100 or 200 m from the transect most likely reflected the behavioral scale of the deer. Coverage of palatable understory vegetation increased with proximity to forest edge and decreased with deer density. Variability in the level of fecal nitrogen could thus be explained by food availability in the landscape. These results suggest that landscape alterations increase the carrying capacity of sika deer and thereby increase impacts upon the ecosystem.     

Suppression of Regeneration in New Zealand Mountain Beech Forests is Dependent on Species of Introduced Deer

We compared the impacts on forest regeneration of introduced sika (Cervus nippon) and red (Cervus elaphus) deer in New Zealand. Plot data were used to compare mountain beech (Nothofagus solandri var. cliffortioides) regeneration between a region with sika deer, and four regions without sika deer. All regions surveyed had red deer present. In the region where sika deer had been present for more than a decade, there was evidence of poor mountain beech seedling regeneration. In the four regions without sika deer, there was evidence of a strong regenerative response at stands with low occupancy by trees. When compared to larger deer species, sika deer have a digestive morphology allowing greater dietary versatility, which may result in them impeding forest regeneration where red deer do not. In contrast to mountain beech, some small-leaved shrub species may have been competitively advantaged by intensive browsing from sika deer. This is contra to a current view that small-leaved shrub species with interlacing branches were able to tolerate browsing from extinct ratite birds, but not introduced deer. Sika deer have been introduced into countries where other deer species are indigenous, such as Canada, Denmark, Great Britain, Czech Republic, Ireland and the United States of America. Because of their dietary advantage, sika deer may have a greater potential to impede forest regeneration and competitively exclude larger deer species, particularly at low basal area sites where impacts on tree regeneration are likely to be greatest.     

Food habits of sika deer in the Shiranuka Hills,eastern Hokkaido: a northern example from the north–south variations in food habits in sika deer

Seasonal changes in the composition of the diet of the sika deer population in the Shiranuka Hills, eastern Hokkaido, in 1998 were determined by fecal analyses. The deer were dependent on Sasa nipponica, a dwarf bamboo, throughout the year, particularly in winter when it accounted for as much as 77.7% of the diet. It accounted for 33.1% and 45.6% in spring and summer, respectively, and this decreased to 12.2% in autumn. Besides S. nipponica, all the graminoid categories accounted for large amounts (66–96.7%), while dicotyledonous plants accounted for little (3–8%) except in autumn when they accounted for 31%. The strong dependence of the Shiranuka population on graminoids was different from other Hokkaido deer populations, for example the population from Ashoro/Onbetsu and the extremely high density population on Nakanoshima Island. In spite of these differences, food for all Hokkaido sika deer was poor in winter. Along the north–south geographical cline in the food composition of sika deer along the Japanese archipelago, the Shiranuka population was positioned as a grazer type, in contrast to the southern populations. However, it is important to note that variations are great among local populations in Hokkaido.     

Simulation of the effects of deer browsing on forest dynamics

I introduced forest dynamics to a deterministic herbivore-vegetation model to examine the effects of vertically stratified forest structure on the dynamics of the herbivore–vegetation system, the resilience and stable states of vegetation, and the interactions between deer populations and vegetation. I constructed a model based on data from field studies performed in Hokkaido, northern Japan. Three phases of state were identified for a given deer density: (1) understory vegetation is maintained with a equilibrium proportion of canopy gaps in the absence of deer; (2) if the equilibrium proportion of canopy gaps is greater than that in the equilibrium state in the absence of deer, the understory vegetation can be maintained; and (3) the understory vegetation cannot be maintained. At the boundary between phases 2 and 3, the herbivore population level had discontinuous effects on vegetation. When the deer density was held below the threshold, forest vegetation had resilience to recover to the equilibrium stable state at the given deer density, although the equilibrium canopy gap ratio and vegetation biomass differed with deer density. However, the forest vegetation–herbivore system could not be maintained in a stable state without artificial deer population management if food limitation was the only mechanism to keep the deer population at low levels. The deer population must be kept below the boundary between phases 1 and 2 to maintain the forest regeneration processes. The level cannot be determined by observing the deer population; careful observation of forest regeneration processes is required.     

Relationship between a high density of sika deer and productivity of the short-grass (<Emphasis Type="Italic">Zoysia japonica</Emphasis>) community: a case study on Kinkazan Island,northern Japan

An extremely high-density (ca. 800 deer km^–2) wild sika deer population uses a short-grass community dominated by Zoysia japonica on Kinkazan Island in northeastern Japan. To explain why the density of wild deer is quite high on the Zoysia community, (1) we quantified the seasonal productivity of the Zoysia community, (2) we compared food availabilities among the plant communities, and (3) we described the habitat selection by the deer in different seasons. Food availability was greater on the Zoysia community than in the forest understory from spring to fall. The productivity of the Zoysia community was high enough to support the actual high density of the deer (814 deer km^–2) in summer. However, the productivity markedly decreased in winter, when the deer density decreased to less than half (358 deer km^–2) of the summer value. In contrast, the deer density of the adjacent forests was highest in winter (154 deer km^–2) and lowest in spring (19 deer km^–2). These results suggest that the deer using the Zoysia community in summer left and were absorbed into the adjacent forest in winter. If such an adjacent community were absent, many deer would not survive, and consequently the deer density on the Zoysia community in summer would not be so high. This intercommunity movement is particularly important for the deer using a plant community like the Zoysia community, which is highly productive but has a small standing biomass.     

江西桃红岭国家级自然保护区梅花鹿生境适宜性评价

华南梅花鹿(Cervus nippon)被IUCN列入濒危物种,也是我国国家Ⅰ级重点保护动物。目前种群仅分布于江西、浙江、安徽等狭窄的区域内,形成多个孤立种群,生境破碎和丧失被认为是限制梅花鹿种群增长的主要原因。于2011年3月至2013年3月采用样线法和样方法对桃红岭国家级自然保护区梅花鹿栖息地进行了野外调查,利用空间模拟方法,结合地理信息系统(GIS)技术的空间分析功能,以植被类型、坡度、坡向、海拔和人类干扰活动作为评价因子进行了生境适宜性评价。结果表明,桃红岭地区以森林为主,各类林地面积约9 488.15 hm~2,占75.90%,植被类型分为落叶阔叶林、针叶林、常绿阔叶林、针阔混交林、竹林、灌丛、草丛和芭茅丛,面积分别为1664.57、1638.63、3438.21、1247.15、87.85、1143.88、60.92 hm~2和206.94 hm~2。在不考虑人类活动影响时,梅花鹿的适宜生境和次适宜生境面积分别是2233.99 hm~2和2980.24 hm~2,分别占保护区总面积的18.61%和24.83%;而考虑人类活动影响时,梅花鹿的适宜生境和次适宜生境面积分别是1224.04 hm~2和2164.70 hm~2,分别占保护区总面积的10.20%和18.04%。由于梅花鹿的生境受到居民点、主要道路、农田耕作、森林采伐等人类活动的强烈影响,导致大量适宜和次适宜生境丧失、隔离,景观破碎度指数由0.4345增加到0.5898。以潜在可利用生境面积计算,保护区梅花鹿环境容纳量为(568±160)只,而以实际可利用生境面积计算,则只能容纳(368±105)只。适宜生境的丧失和破碎可能是限制桃红岭梅花鹿国家级自然保护区梅花鹿种群恢复的重要因素,在此基础上,通过实际调查提出了管理措施。     

Edge effects in a tropical montane forest mosaic: experimental tests of post-dispersal acorn removal

Forest edges created by scattered-patch clear-cutting have become common in tropical montane cloud forests in the highlands of Chiapas, Mexico. It was hypothesised that forest edges may influence regeneration of oak species, which are canopy dominants in these forests, by affecting the activities of small mammal species. Acorns of different oak species varying in germination timing were offered to predators and/or dispersers at different positions along replicated forest edges during 2 consecutive years. We investigated the effects of (1) edge type (hard and soft), (2) distance from the edge (0, 15, 30, 45 and 60 m inside forest fragments) and (3) oak species, on the rate of acorn removal mainly by small mammals. During a non-masting year, acorn removal was affected by the interaction of edge type and distance from the edge (P<0.05), with acorn removal being highest near hard edges compared to adjacent forest interiors. As predicted, acorn removal was greater along soft (100%) than along hard edges (82%), but this pattern was recorded only during the non-masting year. This study partly supports previous studies of rodents preferentially consuming acorns with early germination rather than acorns exhibiting dormancy, however these patterns may change with variation in acorn abundance. These results suggest that patch clear-cutting affects regeneration processes within forest fragments by influencing the activities of small mammals, but the nature of this effect also depends on acorn abundance and the characteristics of the forest edge created.     

Road edge effect on the abundance of the lizard <Emphasis Type="Italic">Gallotia galloti</Emphasis> (Sauria: Lacertidae) in two Canary Islands forests

Transportation infrastructure is a main cause of environmental change in forest landscapes worldwide. In the Canary Islands, a dense road system fragment the native Canarian pine and laurel forests causing potential changes in population densities of endemic lacertid lizards (genus Gallotia). Our aim was to assess road edge effects on relative abundance patterns of the endemic Gallotia galloti in both forests. We also explored the species–habitat relationships in this road-fragmentation context. We found that lizard relative density in relation to road edges differed between forests. Lizards were more abundant along edges and leeward interior, but virtually absent from the interior of the windward laurel forest. In the pine forest, lizards were present at three distances from edge, with a net decrease in abundance from edge to interior. These patterns may be explained partly by differences in vegetation structure regarding road proximity in each forest that potentially affect the helio- and thigmothermic character of G. galloti, and thus its habitat use. A general suggestion of this study is that road margins create corridors that may be used by native lizards for dispersal through inhospitable forest matrix. The high road density in Tenerife may have negative implications for the conservation of the genetic variability of G. galloti. At the island scale, increased communication between lizard populations through road corridors might increase homogenization of the gene pool. Ecological processes in which this lizard plays important roles may also experience changes along road edges.     

Contrasting responses of web-building spiders to deer browsing among habitats and feeding guilds

We examined web-building spider species richness and abundance in forests across a deer density gradient to determine the effects of sika deer browsing on spiders among habitats and feeding guilds. Deer decreased the abundance of web-building spiders in understory vegetation but increased their abundance in the litter layer. Deer seemed to affect web-building spiders in the understory vegetation by reducing the number of sites for webs because vegetation complexity was positively correlated with spider density and negatively correlated with deer density. In contrast, the presence of vegetation just above the litter layer decreased the spider density, and deer exerted a negative effect on this vegetation, possibly resulting in an indirect positive effect on spider density. The vegetation just above the litter layer may be unsuitable as a scaffold for building webs if it is too flexible to serve as a reliable web support, and may even hinder spiders from building webs on litter. Alternatively, the negative effect of this vegetation on spiders in the litter may be as a result of reduced local prey availability under the leaves because of the reduced accessibility of aerial insects. The response to deer browsing on web-building spiders that inhabit the understory vegetation varied with feeding guild. Deer tended to affect web-invading spiders, which inhabit the webs of other spiders and steal prey, more heavily than other web-building spiders, probably because of the accumulated effects of habitat fragmentation through the trophic levels. Thus, the treatment of a particular higher-order taxon as a homogeneous group could result in misleading conclusions about the effects of mammalian herbivores.     

Effect of increased food abundance near forest edges on flocking patterns of Coal Tit Parus ater winter groups in mountain coniferous forests

CapsuleIndividuals concentrated near forest edges in bigger social groups than in forest interiors and foraged more on pine cones which were more abundant there.

Aims To evaluate differences in food distribution between forest edges and forest interiors and their effects on the non-breeding flocking patterns of Coal Tit populations inhabiting mountain coniferous forests.

Methods We collected cone production data at forests edges and interiors in mountain pine forests located in the Pyrenees (northeast Iberian peninsula). At the same sites, we also quantified Coal Tit abundance, flocking patterns and foraging behaviour by means of paired bird surveys during autumn and early winter.

Results We recorded a larger abundance of pine cones available on trees along forest edges compared with forest interiors. Coal Tit groups were of bigger size along forest edges, although the number of social groups detected did not differ from forest interiors. Our observations on foraging behaviour supported the hypothesis that differences in flock sizes and overall abundances associated with distance to the edge are due to differences in the availability of pine cones and to the heavier use of these foraging substrates by birds along forest edges.

Conclusions Our results suggest that by changing food distribution, edge effects on pine cone production may be significantly involved in local changes in the social structure of the Coal Tit. An increase in resource heterogeneity and local population density may have important implications at a population level, such as favouring mobility of individuals searching for food resources and thus a transient life, and increasing the costs of territory defence to resident individuals.     

Down by the riverside: Riparian edge effects on three monkey species in a fragmented Costa Rican forest

Rivers represent natural edges in forests, serving as transition zones between landscapes. Natural edge effects are important to study to understand how intrinsic habitat variations affect wildlife as well as the impact of human-induced forest fragmentation. We examined the influence of riparian and anthropogenic edge on mantled howler, white-faced capuchin, Central American spider monkeys, and vegetation structure at La Suerte Biological Research Station (abbreviated as LSBRS), Costa Rica. We predicted lower monkey encounter rate, tree species richness, and median dbh at both edge types compared to interior and that monkeys would show species-specific responses to edge based on size and diet. We expected large, folivorous–frugivorous howler monkeys and small, generalist capuchins would be found at increased density in forest edge, while large, frugivorous spider monkeys would be found at decreased density in forest edge. We conducted population and vegetation surveys along interior, riparian, and anthropogenic edge transects at LSBRS and used GLMM to compare vegetation and monkey encounter rate. Tree species richness and median dbh were higher in forest interior than anthropogenic edge zones. Although spider monkey encounter rate did not vary between forest edges and interior, howler monkeys were encountered at highest density in riparian edge, while capuchins were encountered at highest density in anthropogenic edge. Our results indicate that diverse forest edges have varying effects on biota. Vegetation was negatively affected by forest edges, while monkey species showed species-specific edge preferences. Our findings suggest that riparian zones should be prioritized for conservation in Neotropical forests.     

Food habits of sika deer on the Boso Peninsula,central japan

The rumen contents of sika deer (Cervus nippon Temminck) on the Boso Peninsula, central Japan, were analyzed to identify local, sexual and age-specific differences in food habits. Graminoids and woody plants were the primary foods throughout the year. In winter, the use of evergreen broad leaves increased. The food habits of sika deer on Boso Peninsula were intermediate between those of populations inhabiting northern and southern Japan. Acorns, mainlyLithocarpus edulis Nakai, were consumed in fall and winter with a peak in October. Since the availability of acorns is not influenced by foraging in previous years, it can be regarded as a stable food supply and hence may be important for deer on the Boso Peninsula. The local difference between the Amatsukominato (AT) area, having a large plantation ofLithocarpus producing acrons, and the Kamogawa-Katsuura (KK) area, having a small plantation ofLithocarpus, was recognized; seeds and fruit were consumed more in AT than in KK in fall and winter. Males consumed more seeds and fruit than females at both sites in fall. This can be attributed to sexual differences in nutritional requirement.     

Food habits of sika deer and nutritional value of sika deer diets in eastern Hokkaido, Japan

Rumen content analysis and field observations were used to investigate the food habits and diet quality of sika deer (Cervus nippon yesoensis Heude) from 1991 to 1993 in eastern Hokkaido, Japan. Diets varied seasonally, with deer consuming graminoids and browse in winter, forbs and agricultural crops in spring and summer and all of these plant foods in autumn. Eighty-four plant species with sika deer bite marks were identified and their use also varied seasonally. The diversity of food resources available provided both critical protein and digestible energy, allowing for physiological maintenance and seasonal growth. With these high-quality diets, deer maintained good body condition in eastern Hokkaido, where the population density was relatively low.     

Deer density affects ground-layer vegetation differently in conifer plantations and hardwood forests on the Boso Peninsula, Japan

Deer overabundance reduces forest ground-layer vegetation and can cause cascading impacts on a forest ecosystem. To predict these effects, we must elucidate the relationship between deer density and the status of ground-layer vegetation. This relationship was studied in the Boso Peninsula, where the deer population density exhibits a clear geographical gradient. We examined species richness and cover of ground-layer vegetation at several cedar plantations and hardwood forests in the Boso Peninsula. We also examined whether deer impacts were altered by light condition, soil water content and forest type (cedar and hardwood). Species richness of ground-layer vegetation was maximized at an intermediate level of deer density, suggesting an intermediate disturbance effect. This phenomenon was compatible with the observation that evergreen species, which were competitive dominants, decreased with increasing deer density, whereas less competitive deciduous species increased until herbivory was intermediate. As deer density increased, cover of ground-layer vegetation gradually decreased, but species unpalatable to deer increased in abundance, suggesting indirect positive effects of deer for unpalatable species. Cedar plantations tended to have greater species richness and ground-layer cover than hardwood forests with similar deer levels. Canopy openness, an indicator of light conditions, increased species richness of hardwood forests and ground-layer cover of cedar plantations, even under deer herbivory. Topographic wetness index, an indicator of soil water content, significantly increased the ground-layer cover of cedar plantations under deer herbivory. These results emphasize the importance of environmental productivity and forest type in the management of ground-layer vegetation experiencing deer overabundance.     

Inflow of seeds through the forest edge: evidence from seed bank and vegetation patterns

To determine the influence of the proximity of a forest edge on seed bank composition and diversity, we performed a seed bank sampling at ancient deciduous forests bordering intensive arable fields. Also vegetation patterns were taken into account. We hypothised that forest edges may facilitate the entrance of diaspores of invasive species into the forest and the successive incorporation of these species in the forest seed bank. We noticed a substantial influence of the proximity of an edge on seed bank composition at as well the forested side of the edge as the field side. The forest edge zone was limited to 3 m into the forest and the field edge zone extended 3m into the field. The seed bank samples of field and forest edge are characterised by a higher species diversity and seed density and a higher similarity between seed bank and vegetation, compared to field or forest samples. The forest edges contains fewer pioneer species in comparison with the forest interior and more competitive species and species of edges and clearings compared with field and forest samples. The seed longevity index increases towards the forest interior. We can conclude from our data that the forest and edge seed bank are composed by both seeds from recent dispersal processes and local seed set and by seeds originating from past vegetation on the site. Near the edge, actual seed input seems of primal importance. Further towards the forest interior seed input decreases and long-living seeds of past vegetation become more important. Ancient forest edges thus act as a barrier for seeds of species of the surrounding arable field.     

Rain Forest Structure at Forest-Pasture Edges in Northeastern Costa Rica

Land-use change in the Sarapiquí region of Costa Rica has resulted in a fragmented forest landscape with abrupt edges between forest and pasture. Forest responses to edge effects vary widely and can significantly affect ecosystem integrity. Our objective was to examine forest structure at 20+ yr old forest-pasture edges in Sarapiquí. Three transects with 0.095-ha plots at seven distances from forest edges were established in each of six forest patches. Stem density, basal area, and aboveground biomass in trees and palms ≥ 10-cm diameter at breast height were measured in all plots. In addition, hemispherical photographs were taken to determine leaf area index, understory light availability, and percent canopy openness. Linear mixed-effects models showed significantly higher tree stem density at forest edges, relative to interiors, a pattern reflected by increased stem density, basal area, and aboveground biomass in small diameter trees (≤ 20 cm) growing near edges. No differences in total tree basal area, aboveground biomass, or hemispherical photograph-derived parameters were detected across the forest edge to interior gradient. The recruitment of small diameter trees following edge creation has contributed to the development of dense vegetation at the forest edge and has aided in the maintenance of similar tree basal area and aboveground biomass between edge and interior environments. These data reflect on the robustness of forest edges in Sarapiquí, a characteristic that will likely minimize future detrimental edge effects and promote a number of high-value environmental services in these forests.     

Diversity of dung-beetle community in declining Japanese subalpine forest caused by an increasing sika deer population

The Ohdaigahara subalpine plateau in Japan has recently suffered a reduction in primary forest land caused by an increasing population of sika deer (Cervus nippon). Deer have debarked many trees, causing dieback, gradually changing the primary forest first to light forest with a floor that is densely covered with sasa grass (Sasa nipponica) and then to S. nipponica grassland. To examine the effects of vegetative transformation on the dung-beetle community, we compared the diversity and abundance of dung-beetle assemblages in the primary forest, transition forest, and S. nipponica grassland using dung-baited pitfall traps. The species richness and species diversity (Shannon-Wiener index) were significantly highest in the primary forest and lowest in the S. nipponica grassland. The evenness (Smith-Wilson index) was highest in the primary forest and nearly equal in the transition forest and S. nipponica grassland. The abundance was apparently greater in the transition forest than in the primary forest and S. nipponica grassland. These results suggest that loss of primary forest resulting from an increasing deer population decreases the diversity of the dung-beetle community while increasing the abundance of dung beetles in the transition forest. Sika deer use transition forests and grasslands more frequently than primary forests as habitat, but an increase in dung supply there does not necessarily increase the diversity or abundance of dung-beetle assemblages.     

Densities of Two Frugivorous Primates with Respect to Forest and Fragment Tree Species Composition and Fruit Availability

Conservation of wildlife populations requires extensive knowledge of their habitat requirements, efficient methods to evaluate habitat quality, and an understanding of the value of fragments and edges. Kibale National Park, Uganda has areas that differ in the densities of 2 species of frugivorous monkeys—Cercopithecus mitis and Lophocebus albigena—including one on an edge and forest fragments outside the park that lack both species. We compared the basal area densities of important food trees with primate densities. The density of Cercopithecus mitis correlates most strongly with the basal area density of all types of food trees combined. The density of Lophocebus albigena does not correlate with the basal area densities of any category of food trees or with fruit availability. An index of their density—number of groups seen per km walked—correlates to fruit availability but with marginal significance. Lack of a relationship between the basal area densities of food trees and density of Lophocebus albigena may be the result of a mismatch in scale between the forest area measured and their large home ranges. We compared the unused area of forest to the other areas of the forest and the fragments and found it had higher basal area densities in all food tree categories for both species than the fragments and lower basal area densities of most categories than the other parts of the forest, indicating that the fragments are poor quality and would probably be unused even if dispersal were likely.     

Comparative rumen morphology of sympatric sika deer (Cervus nippon) and red deer (C. elaphus scoticus) in the Ahimanawa and Kaweka Ranges,central North Island,New Zealand

Eighteen sika deer (Cervus nippon) and 14 red deer (C. elaphus scoticus) were sampled from two areas where these closely related species are sympatric. Total body weight, carcass weight, age class, sex, and internal parameters (e.g. liver weight, kidney weight, rumen volume) were recorded. Samples of rumen wall mucosa taken from the dorsal rumen wall, atrium ruminis, caudoventral blindsac, and ventral rumen wall were used to compare rumen morphology between the two species (and also any area, sex, and age effects). Sika deer had significantly lower papilla densities in three of the four rumen wall sites and significantly smaller papillae than red deer in two of the four sites. Surface enlargement factors (SEFs) were calculated to provide comparisons of the effective absorptive surface within the rumens of the two species. The mean SEF for sika deer (4.76) was significantly less that for red deer (6.77), which suggests a greater degree of adaptation to digesting fibrous forage. In the central North Island, New Zealand, where the habitat has been considerably modified by introduced herbivores over the last century and food resources are depleted, such an adaptation would confer a competitive advantage on sika deer over red deer. It is postulated that this diet-related difference largely accounts for the ongoing replacement of red deer by sika deer where these two species are sympatric.     

Risk factors affecting the probability of damage by sika deer in plantation forests in Yamanashi Prefecture, Japan

Using a generalized linear mixed model approach, we determined the most important risk factors affecting the probability of damage by sika deer in a forest plantation in Japan. Candidate risk factors included tree species, stand age, peripheral dwarf bamboo community, topographical factors, snow depth, and human disturbance factors. Based on this model, we developed a risk map of forest damage. The model indicated that the most important risk factor was stand age, followed by tree species and maximum snow depth. Our predictive model has practical use due to its high classification accuracy (83.9%). To decrease damage from sika deer, an afforestation plan that incorporates these factors should be implemented. Because it is based on common, forest GIS data that have recently been compiled by several local governments in Japan, our modeling method of deer damage can easily be adapted to other areas.