Predicting Summer Site Occupancy for an Invasive Species,the Common Brushtail Possum (Trichosurus vulpecula), in an Urban Environment

Invasive species are often favoured in fragmented, highly-modified, human-dominated landscapes such as urban areas. Because successful invasive urban adapters can occupy habitat that is quite different from that in their original range, effective management programmes for invasive species in urban areas require an understanding of distribution, habitat and resource requirements at a local scale that is tailored to the fine-scale heterogeneity typical of urban landscapes. The common brushtail possum (Trichosurus vulpecula) is one of New Zealand’s most destructive invasive pest species. As brushtail possums traditionally occupy forest habitat, control in New Zealand has focussed on rural and forest habitats, and forest fragments in cities. However, as successful urban adapters, possums may be occupying a wider range of habitats. Here we use site occupancy methods to determine the distribution of brushtail possums across five distinguishable urban habitat types during summer, which is when possums have the greatest impacts on breeding birds. We collected data on possum presence/absence and habitat characteristics, including possible sources of supplementary food (fruit trees, vegetable gardens, compost heaps), and the availability of forest fragments from 150 survey locations. Predictive distribution models constructed using the programme PRESENCE revealed that while occupancy rates were highest in forest fragments, possums were still present across a large proportion of residential habitat with occupancy decreasing as housing density increased and green cover decreased. The presence of supplementary food sources was important in predicting possum occupancy, which may reflect the high nutritional value of these food types. Additionally, occupancy decreased as the proportion of forest fragment decreased, indicating the importance of forest fragments in determining possum distribution. Control operations to protect native birds from possum predation in cities should include well-vegetated residential areas; these modified habitats not only support possums but provide a source for reinvasion of fragments.     

An inventory of mammalian pests in a New Zealand city

Abstract

Urban ecosystems include many habitat types supporting native flora and fauna. These habitats may also sustain populations of introduced mammalian pests, although relatively little is known about the composition or distributions of these species in urban environments. We made a preliminary survey of the distribution and relative abundance of pest mammals across three urban habitat types (gully, amenity park, residential) in Hamilton, New Zealand. Tracking tunnel and WaxTag® surveys showed that: (1) rats (Rattus rattus or R. norvegicus), mice (Mus musculus) and brushtail possums (Tricho‐surus vulpecula) were detected most often and in highest abundances in gullies, were relatively rare in amenity parks, and were not detected in residential areas; (2) hedgehogs (Erinaceus europaeus) were regularly detected in all habitats; and (3) mustelids were not detected in any habitat. Live trapping in areas where rats were detected captured 21 rats, of which 19 were R. rattus. Scat and animal sign surveys found evidence of lagomorphs in amenity parks and gullies but not in residential areas. Cats (Felis catus) were detected in all habitats. These data suggest that important mammalian pests other than cats are either absent (mustelids) or comparatively rare (rats) in urban Hamilton and largely confined to gully habitats. Further research is needed to determine whether detection probabilities of target species vary significantly between the species and habitats we sampled, and to quantify the impact that mammalian pests might have in limiting populations of other animals, especially avifauna.     

Bird communities of natural and modified habitats in Panama

Only a small proportion of land can realistically be protected as nature reserves and thus conservation efforts also must focus on the ecological value of agroecosystems and developed areas surrounding nature reserves. In this study, avian communities were surveyed in 11 habitat types in central Panama, across a gradient from extensive forest to intensive agricultural land uses, to examine patterns of species richness and abundance and community composition. Wooded habitats, including extensive and fragmented forests, shade coffee plantations, and residential areas supported the most species and individuals. Nearctic-Neotropical migratory species were most numerous in lowland forest fragments, shade coffee, and residential areas. Introduced Pinus caribbea and sugar cane plantations supported the fewest species compared to all other habitats. Cattle pastures left fallow for less than two years supported more than twice as many total species as actively grazed pastures, such that species richness in fallow pastures was similar to that found in wooded habitats. Community similarities were relatively low among all habitat types (none exceeding the observed 65% similarity between extensive and fragmented lowland forests), but communities in shade coffee and residential areas were 43% and 54′% similar to lowland forest fragments, respectively. Fallow pastures and residential areas shared 60% of their species. Bird communities in shade coffee and residential areas were characterized by higher proportions of frugivorous and nectarivorous species than in native forests. These same guilds also were better represented in fallow than in grazed pastures. Raptors and piscivorous species were most prevalent in cattle pastures and rice fields. These results, though based upon only species richness and abundance, demonstrate that many human-altered habitats have potential ecological value for birds, and conservation efforts in tropical areas should focus greater attention on enhancement of agricultural and developed lands as wildlife habitat. To understand the true conservation value of these modified lands will require examination not only of numbers but also of the types of species supported by these habitats, their reproductive output and survival rates.     

Landscape context of plantation forests in the conservation of tropical mammals

Plantation forests have been expanding in many tropical and subtropical environments. Howerver, even when they replace less wildlife friendly land uses such as pastures and annual crops, the biodiversity levels of pristine natural habitats often have not been recovered. Here we addressed how the landscape context of plantation forests located in South-eastern Brazil affects species richness and community resilience of medium and large size mammals. The area covered by native habitat fragments surrounding plantation forests is positively related to functional richness, including the presence of species more vulnerable to extinction in fragmented landscapes. In addition, the degree of aggregation of plantation forest stands is negatively related to more vulnerable species. No primates were recorded in our seven plantation forest sites (ranging from 272 to 24,921 ha), even when they were seen in native habitat fragments adjacent to commercial tree stands. Two invasive species (Sus scrofa and Lepus capensis) were recorded in four plantation forest sites. The impoverishment of fauna in plantation forests is due to two factors. First, plantation forests generally are structurally simplified habitats when compared to highly diverse tropical forests. Secondly, the isolation from habitat fragments which act as source of individuals in the landscape precludes the establishment of individual in plantation forest. We also highlighted the management practices to improve the complexity of vegetation in commercial tree stands should be taken cautiously, insofar as reduced productivity per area entails a greater demand for land. Thus, an alternative would be intensify the management of the commercial tree stands for wood production together with the restoration of adjacent areas set aside to conservation and native habitat fragments protection.     

Urban and coastal breeding lesser black-backed gulls (Larus fuscus) segregate by foraging habitat

Despite urbanization's general erosion of biodiversity, towns and cities provide novel opportunities for some species. During the 20th century, gulls (Laridae) colonized urban areas around the world where they flourished. At the same time, some coastal populations declined. The reasons for this difference are not fully understood, partly because little is known about any ecological differences between urban and non-urban gulls, such as their foraging ecology. Here we compare the movement ecology and habitat selection of Lesser Black-backed Gulls Larus fuscus graellsii breeding at two neighbouring colonies – one urban and one coastal – in north-west England. We used bird-borne GPS loggers to first compare colony-level movement behaviour and habitat selection and then investigated individual-level habitat use. We observed clear colony-level habitat segregation: urban breeders preferentially foraged in urban areas whereas coastal breeders foraged primarily in coastal habitats and avoided urban areas. Coastal breeders also had larger core and home-ranges than urban breeders, possibly due to differences in colony size. However, we also found inter-individual differences in habitat use, which may have important management implications. These findings suggest a link between nesting and foraging ecology, and thus management or environmental change altering food availability will impact gulls at the coastal and urban sites differently.     

Tree colonization by the Asian longhorn beetle,Anoplophora glabripennis (Coleoptera: Cerambycidae): effect of habitat and tree suitability

Tree colonization and feeding activity of the invasive wood‐borer Asian longhorn beetle (Anoplophora glabripennis), an Asian pest introduced into North America and Europe, was studied in a newly invaded area in Italy. The hypothesis being tested was that the reproductive success of the insect depend on habitat type and tree suitability. Adult beetles were caged on branches of host and nonhost species, in both urban and forest habitats. Two months later, number and size of feeding patches on plant tissues, eggs laid, and surviving larvae were assessed. Bark concentration of C and N was also measured from the same trees. Results indicated that the mean area of plant tissues consumed by adult feeding was significantly larger on trees growing in forest than in urban habitat, although within the same habitat there were no differences between susceptible and nonsusceptible trees. ALB tree colonization, in terms of number of eggs laid and young larvae survival, was not affected by habitat while it was higher on susceptible trees. Although trees growing in forests had a lower nitrogen concentration, they allowed colonization rates similar to those of trees growing in the urban habitat. Hence, the amount of carbon and nitrogen did not fully explain tree suitability or habitat selection. We suggest compensatory feeding as a potential mechanism that might explain this peculiar situation, as supported by a more intensive feeding activity recorded on trees in the forest. Suitability of different trees may be due to other factors, such as secondary chemical compounds.     

北京不同类型人工林中蛾类群落特征

近年来,绿化造林工程在北京市大规模展开,涉及林地生境类型多样.林地昆虫群落组成和多样性特征受林分树种组成、立地条件、管护方式的影响.通过对北京市5种人工林样地中蛾类群落结构、多样性和相似性及其时空动态特征的调查,探讨了蛾类多样性变化对生境类型的生态响应.结果 表明:共获得标本7046头,隶属于25科269种,以夜蛾科(...     

The effect of forest clear-cutting on habitat use in Sichuan snub-nosed monkey (<Emphasis Type="Italic">Rhinopithecus roxellana</Emphasis>) in Shennongjia Nature Reserve,China

The habitat use of two groups of Sichuan snub-nosed monkey (Rhinopithecus roxellana) was observed, using the focal group method, for 7 months in four seasons from June 2000 to October 2001. The habitats were classed into primary forest and three successional habitats: after clear-cutting: grassland, shrub forest and young forest. The results showed that the large group of monkeys had larger range areas than the small group in the same season. Both groups had larger range sizes in summer or autumn than in winter or spring. They spent most of their time using primary forest and young forest, rarely used shrub forest and did not use grassland. In each season, they used the habitats non-randomly and preferred primary forest. The preference order of habitats for both groups every season was the same: primary forest > young forest > shrub forest grassland. The results suggested that primary forest was high-quality habitat and should be conserved as a top priority. Clear-cutting would cause habitat loss and habitat fragmentation, and should therefore be prohibited. High-quality habitat for the monkeys is difficult to restore from clear-cutting.     

House sparrow (Passer domesticus) habitat use in urbanized landscapes

The house sparrow (Passer domesticus) is showing population declines in many parts of Europe, with recent declines being particularly severe in urban areas. To date, relatively little is known about the species’ habitat associations within urbanized landscapes. We report here an investigation of the habitat associations of house sparrows using a survey of 1223 stratified randomly selected 500 × 500-m squares within urbanized landscapes of the UK, defined as at least 25% ‘human cover’. The densities of chirping male house sparrows and of all house sparrows were analysed separately to obtain insights into breeding habitat requirements and general habitat associations, respectively. Multi-model inference showed that residential areas (houses, flats), allotments (areas used for small-scale horticulture) and farm buildings were key predictors of house sparrow density and chirping male density. Separate analyses on landscapes of differing human cover showed similar results. Within residential areas, the increase of house sparrow density with habitat area (on a log scale) was approximately threefold greater when private gardens were present than when they were absent. The model predicted a rapid decline in house sparrow abundance when only a small area of private gardens is converted to continuous housing. Allotments and residential areas with gardens are likely to be under pressure due to increased demand for housing, specifically from the infilling of green space within urban areas. It would seem to be imperative that any action plan to protect urban house sparrow populations should include specific protection of such key habitats. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Forest Fragment and Breeding Habitat Characteristics Explain Frog Diversity and Abundance in Singapore

Habitat loss and fragmentation can have severe negative and irreversible effects on biodiversity. We investigated the effects of forest fragmentation on frog diversity in Singapore because of its high rates of deforestation and the demonstration that frogs are some of the most sensitive species to habitat degradation. We surveyed frog species in 12 forest fragments varying from 11 to 935 ha. We compared differences in species richness, abundance, and Shannon's index in relation to forest fragment size, connectivity (distance between fragments), and breeding habitat heterogeneity. A total of 20 species from 12 genera and five families were encountered in 12 fragments. Larger fragments and those closer to larger fragments had higher species richness. Abundance, however, was not correlated with forest area or connectivity, but we found fewer individual frogs in the larger fragments. We also found that breeding habitat heterogeneity best explained frog species diversity and abundance in forest fragments. Fragments with a high diversity of breeding habitats had more species. We found no evidence to suggest that abundance and diversity are strongly correlated, particularly in disturbed areas, but that breeding habitat heterogeneity is an under-appreciated factor that should be considered when prioritizing areas for anuran conservation. Enriching breeding habitat heterogeneity, creating corridors between fragments, and reforesting degraded areas are some of the most beneficial strategies for preserving urban frog biodiversity.     

The Use of an Invasive Species Habitat by a Small Folivorous Primate: Implications for Lemur Conservation in Madagascar

The lemurs of Madagascar are among the most threatened mammalian taxa in the world, with habitat loss due to shifting cultivation and timber harvest heavily contributing to their precarious state. Deforestation often leads to fragmentation, resulting in mixed-habitat matrices throughout a landscape where disturbed areas are prone to invasion by exotic plants. Our study site, the Mandena littoral forest (southeast Madagascar), is a matrix of littoral forest, littoral swamp, and Melaleuca swamp habitats. Here, Melaleuca quinquenervia has invaded the wetland ecosystem, creating a mono-dominant habitat that currently provides the only potential habitat corridor between forest fragments. We sought to understand the role of this invasive Melaleuca swamp on the behavioral ecology of a threatened, small-bodied folivore, the southern bamboo lemur (Hapalemur meridionalis). We collected botanical and behavioral data on four groups of H. meridionalis between January and December 2013. Our results confirm Melaleuca swamp as an important part of their home range: while lemurs seasonally limited activities to certain habitats, all groups were capable of utilizing this invasive habitat for feeding and resting. Furthermore, the fact that Hapalemur use an invasive plant species as a dispersal corridor increases our knowledge of their ecological flexibility, and may be useful in the conservation management of remaining threatened populations.     

The hunting habitats of Goshawks Accipiter gentilis in boreal forests of central Sweden

Radio-tracking was used to study the habitat utilization of hunting Goshawks Accipiter gentilis in central Swedish boreal forest. Data were collected during September-June, in 1977–1981. There was no major difference between autumn, winter or spring in any aspect of habitat preference, despite seasonal differences in prey composition. Despite the strong reversed sexual size-dimorphism, no habitat segregation between the sexes was found. It is argued that this is due to the lack of prey segregation between the sexes. Goshawks showed a strong preference for mature forest, which was chosen twice as often as would be expected from its prevalence in the environment, whereas the younger successional stages of the forest were under-used. Goshawks preferred large habitat patches, and showed no major preference with respect to tree species composition of the forest. Kills made by radio-tagged Goshawks showed the same habitat- and patch-size distribution as did Goshawk locations in general. It is argued that the profitability of different hunting habitats, and thus habitat preferences, is not only determined by prey density in the habitats, but also by other habitat features influencing the Goshawk's ability to hunt there. Winter home-ranges averaged 5700 ha, considerably larger than Goshawk ranges recorded in areas with richer prey supply.     

Are urban habitats ecological traps for a native songbird? Season‐long productivity,apparent survival,and site fidelity in urban and rural habitats

The northern mockingbird Mimus polyglottos is a native species that is more abundant in urban than non‐urban habitats (i.e. an urban‐positive species). Abundance alone, however, is not an accurate index of habitat quality because urban habitats could represent ecological traps (attractive sink habitat) for urban‐positive species. We compared mockingbird nesting productivity, apparent survival, and decision rules governing site fidelity in urban and rural habitats. If the higher abundance of mockingbirds in urban habitats is driven by higher quality urban habitat, then we predicted that productivity of urban mockingbirds would exceed the estimated source‐sink threshold and productivity of non‐urban mockingbirds. If, on the other hand, urban habitats act as ecological traps, productivity would be lower in urban habitats and would fall below the estimated source‐sink threshold. Productivity of urban pairs exceeded that of non‐urban pairs and more than offset estimated adult mortality, which makes urban habitat a likely source habitat. Apparent adult survival was higher in urban habitats than in non‐urban habitats, although this could be driven by dispersal more than mortality. Decision rules also appeared to differ between urban and non‐urban populations. Females in urban habitats with successful nests were more likely to return than those with unsuccessful nests, whereas return rates of females in nonurban habitats were unrelated to nesting success and may be more related to nesting habitat availability. We conclude that urban habitats do not act as ecological traps that lure mockingbirds into sink habitat and that increased breeding productivity contributes to their success in urban habitats.     

Diurnal activity patterns of coypu in an urban habitat

The ability of species to adapt to changing environmental conditions is a crucial need for surviving in human influenced landscapes. Especially non-indigenous species, which are flexible in their behaviour, may successfully enter and establish in new habitats. We compared the activity patterns of the invasive coypuMyocastor coypus (Molina, 1782) in an urban habitat in Germany with patterns observed in its native distribution range in South America as well as in areas, where it was introduced. In summer 1998 and winter 1999 a radio tracking study was carried out on six adult coypus, supplemented by count data on untagged animals. Whereas main activity in non-urban populations was recorded at night including dusk and dawn, coypus in the urban habitat were active during the day and early evening. We never observed animals between midnight and dawn. This reversal of activity patterns is attributable to deposition of human food in the city area during the daytime.     

Shifts from native to invasive small mammals across gradients from tropical forest to urban habitat in Borneo

Urbanization has paved the way for the spread of commensal rodents at global scale. However, it is largely unknown how these species use tropical anthropogenic landscapes originally covered with forests and inhabited by diverse small mammal assemblages. We surveyed non-flying small mammals in various urban and suburban habitat types and adjacent forest in the tropical town of Kota Kinabalu in Borneo. We used occupancy and polynomial regression models to determine variation in species occurrences along gradients of land-use intensity. Müller’s sundamys (Sundamys muelleri) was the only native small mammal species found in urban and suburban landscapes with a continuous decrease in occurrence probability from forests to urban habitats. The invasive Asian black rat (Rattus rattus species complex) and the invasive Asian house shrew (Suncus murinus) had the highest occurrence probabilities in habitats of intermediate land-use intensity, but Asian black rats are also likely to occasionally invade forested habitats and occupied urban habitats in sympatry with the Norway rat (Rattus norvegicus). In urban and suburban habitats, fallow land possibly favoured the occurrence of S. muelleri and S. murinus. Other native small mammal species (Muridae, Sciuridae, Tupaiidae) were found only in forested areas. Our study shows that native small mammals found in forest are largely replaced by invasive species in urban and suburban habitats. Due to their occurrence in habitats of various land use intensities, S. muelleri and R. rattus comprise central links between forest wildlife and urban species, an association that is important to consider in studies of parasite and disease transmission dynamics.     

Habitat degradation and introduction of exotic plants favor persistence of invasive species and population growth of native polyphagous fruit fly pests in a Northwestern Argentinean mosaic

Expansion of agricultural land is one of the most significant human alterations to the global environment because it entails not only native habitat loss but also introduction of exotic species. These alterations affect habitat structure and arthropod dynamics, such as those among host plants, tephritid fruit flies, and their natural enemies. We compared abundance and dynamics of pest and non-pest tephritids and their natural enemies over a mosaic of habitats differing in structure, diversity and disturbance history on the Sierra de San Javier in Tucuman, Argentina. Our prediction was that conserved habitats would be more resistant to the establishment and spread of invasive tephritid species due in part to a greater abundance of natural enemies, a greater diversity of native species in the same family and trophic level, and a greater wealth of biotic interactions. We further predicted that native species with broad host ranges should be more sensitive to habitat loss yet more competitive in less disturbed habitats than generalist native and exotic species. We found that environmental degradation, and introduction and spread of exotic host plants strongly affected distribution patterns, abundance, and phenology of native and exotic tephritids. Monophagous tephritid species and several specialized parasitoids were more sensitive to habitat loss than polyphagous species and parasitoids exhibiting a wide host range. In contrast, native monophagous species and native parasitoids appeared to exclude the invasive Mediterranean fruit fly from conserved patches of native vegetation. Nevertheless, the Mediterranean fruit fly persisted in uncontested exotic host plants and thrived in highly degradeted urban landscapes.     

Responses of kukupa (

The kukupa or New Zealand pigeon (Hemiphaga novaeseelandiae) is gradually declining on the New Zealand mainland, due mostly to predation by introduced pest mammals including ship rats (Rattus rattus) and brushtail possums (Trichosurus vulpecula). We report on a co-operative project between Maori landowners, the Department of Conservation, and Manaaki Whenua–Landcare Research researchers to restore a Northland kukupa population and to examine kukupa nesting success in relation to pest abundance. Ship rats and possums were targeted by trapping and poisoning throughout Motatau Forest (350 ha) from 1997 to 1999; only possums were targeted in 2000. All 13 kukupa nests located before pest control started in late 1997 failed at the egg stage, but all seven nests located in 1998–99 successfully fledged young when trapping and tracking indices of possums and ship rats were less than 4%. After pest control, counts of kukupa and some other bird species increased at Motatau compared with counts in a nearby non-treatment block, suggesting numbers of adult kukupa can be increased in small forest areas by intensive pest control. This increase is due at least partly to increased nest success. Evidence from time-lapse video cameras, sign remaining at nests, and nest success rates under different pest control regimes suggest both ship rats and possums are important predators at kukupa nests.     

Identifying eradication units in an invasive mammalian pest species

Eradication of invasive species is an important component for species conservation and ecosystem restoration. Success of eradications is dependent on knowledge of population connectivity in order to determine reinvasion pathways, and hence populations requiring simultaneous eradication (eradication units). The common brushtail possum (Trichosurus vulpecula) was introduced into New Zealand from Australia and Tasmania, and now occupies a wide range of habitats across the majority of New Zealand. Possums are one of the most destructive invaders within New Zealand, with extensive control operations occurring throughout the country. Understanding the population connectivity of possums on mainland New Zealand (North and South Islands) will enhance the success of planned eradications. We examined the genetic population structure of invasive possums to identify gene flow, thus reinvasion pathways, between seven populations around Dunedin and on the Otago Peninsula where eradication of possums is occurring. Genetic variation at 12 microsatellite loci was comparable between all sampled populations and exhibited a significant isolation by distance pattern. Bayesian clustering methods supported the existence of two population clusters, indicating the presence of a reinvasion pathway onto the Otago Peninsula from urban areas at the Southern end of the Peninsula. To avoid recolonisation, all possums on the Otago Peninsula should be eradicated simultaneously, with the implementation and ongoing maintenance and monitoring of an urban buffer zone. We recommend pre-eradication genetic analyses be adopted by all pest managers to define appropriate eradication units, thereby maximising eradication success and avoiding costly failures.     

Forest-edge utilization by carnivores in relation to local and landscape habitat characteristics in central European farmland

Rapid changes in agricultural landscape structure and composition affect many different farmland biotas, including carnivores, which are a key element of ecosystem stability, yet little is known about their distribution and habitat use. In this study, we evaluated how habitat characteristics on two different spatial scales (local and landscape scale) affected the forest-edge utilization by small and medium-sized carnivores in fragmented central European farmland. Based on an indirect method for detecting carnivores (scent stations), we sampled 212 forest fragments of different sizes (1–7864 ha) during April to May from 2006 to 2009. Our results indicate that carnivore utilization of forest-edge habitats was driven by landscape rather than local characteristics even though the overall extent of explained variation was small. The most important factors that determined response of the carnivore community were the area of farmland and that of urban land on a landscape scale. The corridor connectivity between small forest fragments and other spatial elements played a crucial role in the occurrence of red fox. Our results suggest that comprehensive studies on multi-species carnivore assemblage using scent station might be useful in evaluating species-specific response to habitat characteristics, especially if large numbers of stations visited by carnivores are available.     

Birds at the urban fringe: avian community shifts in different peri-urban ecotones of a megacity

Cities can be regionalized in intra-urban and peri-urban areas. The space between urban areas and adjacent systems represents an ecological transition that often acts as a semi-permeable biological filter. In this study, we assessed changes in avian community species richness, density, and composition at different peri-urban ecotones (i.e., urban-croplands, urban-grasslands, urban-shrublands) of northeastern Mexico City. Species richness was lower in the urban component of urban-grassland and urban-shrubland ecotones, while bird densities were higher in the urban components of the urban-grassland and urban-shrubland peri-urban ecotones, mainly due to the high number of urban exploiter species. However, the urban-cropland peri-urban ecotone exhibited a different pattern, with similar low bird species richness and density values between both components (urban and non-urban). A species composition analysis revealed that urban bird communities were not influenced by adjacent non-urban habitats, since the urban components of peri-urban ecotones were more similar among them than in relation to the rest of non-urban components. In summary, results of this study show that urbanization can represent an important biological filter for birds, often reducing species richness and homogenizing avian communities at local scales. As the environmental variables determining ecological processes related to the semi-permeable filter effect that urban areas pose to biodiversity might depend on urban habitats, regions, and spatial scales, further studies are needed to fully understand this phenomenon.