Effects of location in territory and distance from neighbours on the use of song repertoires by Carolina wrens

By capturing territorial Carolina wrens (Thryothorus ludovicianus) and housing them in portable cages on their territories, I could control each one's location in its territory and distance from neighbours in experiments on the use of song repertoires. Experiment 1 demonstrated that these wrens sang more songs in the centre of their territories than at the edge, but that they did not use more song types or different song types at the centre than at the edge. In experiment 2, in which I played tape-recorded songs at two distances from wrens caged in the centres of their territories, birds responded more strongly to songs at 25m, simulating an intruder, than to songs at 165 m, simulating a territorial neighbour. Birds also switched more frequently between song types and sang more song types per 100 songs in response to the nearer playback. Experiment 3 compared captive wrens 140 m, 80 m, 20 m, or 0 m apart on adjacent territories. As the distance between neighbours decreased, birds sang less, but also switched more frequently between song types, used more song types per 100 songs, and matched songs with neighbours more frequently. There were no differences in the kinds of song types sung at different distances from neighbours. A comparison of the results from experiments 1 and 3 confirms that the use of song repertoires is influenced by distance from conspecifics and not by location in the territory.     

Factors in habitat partitioning in Pseudotropheus aurora (Pisces: Cichlidae), an introduced species to a species-rich community of Lake Malawi

The territorial behaviour of a small rock-dwelling cichlid, Pseudotropheus aurora , was studied at Thumbi Island West at the southern end of Lake Malawi. P. aurora is one of several species that have been introduced to an already diverse fish community. Many of the endemic and introduced species are highly territorial. The coexistence of an introduced territorial species within a speciesrich community of other territorial species is examined. Territories from which P. aurora individuals were removed were invariably re-occupied by conspecifics. Similarly, when all territories of a certain area were emptied of all territory owners of any existing species, the former P. aurora territories were re-occupied by conspecifics.
The possible roles of site specificity, competitive exclusion and behaviourally transmitted traditions regarding territories are discussed.     

Too close for comfort: spatial patterns in acorn barnacle populations

Spatial patterns in aggregations form as a result of the interplay between costs and benefits experienced by individuals. Such self-organisation of aggregations can be explained using a zonal model in which a short-range zone of repulsion and longer-range zone of attraction surrounding individuals leads to emergent pattern properties. The signal of these processes can be detected using spatial pattern analyses. Furthermore, in sessile organisms, post-settlement mortality reveals the relative costs and benefits of positions within the aggregation. Acorn barnacles are known to require contact with conspecifics for reproduction and are therefore believed to aggregate for this purpose; isolated individuals may also be more susceptible to abiotic stress and predation. At short distances, however, competition for space and resources is likely to occur. In this study spatial patterns of barnacles (Semibalanus balanoides L.) were analysed using pair-correlation functions. Individuals were dispersed at distances below 0.30 cm, but peak relative density occurred at a distance of 0.36 cm from conspecifics. This is much closer than required for reproductive access, implying a strong aggregative drive, up to the point of physical contact with neighbours. Nevertheless, analysis of dead barnacles illustrated that such proximity carries a cost as barnacles with many neighbours were more likely to have died. The inferences obtained from these patterns are that barnacles aggregate as closely as they can, and that local neighbourhood competition is a powerful determinant of mortality. These processes give rise to the observed pattern properties.     

Intraspecific interactions affect the spatial pattern of a dominant shrub in a semiarid shrubland: A prospective approach

Dispersal, physical conditions and biotic interactions contribute to determine the spatial distribution of individuals in plant populations. Much of what we know has been learned from studies that retrospectively posit mechanisms presumed to have generated the observed spatial patterns. Here we present a prospective approach. We start by measuring spatial demographic effects and evaluate if they can generate observed spatial patterns. We evaluated the influence of interactions among conspecifics on vital rates, demography and spatial distribution of Croton aff. wagneri, a dominant shrub in dry Andean ecosystems. Recruitment, survival and growth varied in relation with distance to conspecifics neighbours and with their summed cover. We built a spatial individual-based model and simulated its population dynamics in 30 × 30 m plots for a 30-year period. We compared the predicted spatial pattern from these demographic models with that observed among plants in 16 independent plots with the same area. Simulated populations mimicked observed spatial patterns, although in plots at high elevations the simulated populations did not reproduce the observed inhibition at small scales. Observed and simulated patterns indicated differences between elevations in maximum aggregation and location of the distances with higher aggregation. We discuss how consideration of critical seed and juvenile stages and interspecific interactions could further improve our understanding of spatial pattern and recommend that these factors be considered in future models.     

Defence, intrusion and the evolutionary stability of territoriality

Territorial behaviour can only be adaptive if its costs are outweighed by its benefits. Territorial individuals incur costs by defending their territories against intruders. Usually these intruders are assumed to be non-territorial floaters attempting to take over the whole territory or neighbours trying to extend the borders of their own territory. We instead investigate how costs and benefits of territorial behaviour are affected by neighbours which invade to steal resources on a territory.We show analytically that in the absence of defence intrusion into neighbouring territories always pays and that even if territories are defended intrusion levels can still be high. Using a more detailed simulation model we find that territory defence usually disappears from the population even if owners have a strong advantage over intruders in terms of fighting costs or foraging efficiency. Defence and thus territoriality can only be evolutionarily stable if fighting costs for the intruder relative to the productivity of the territory are very high or if crossing the borders between territories carries additional costs.Our results show that stealing of resources by neighbours can have a considerable effect on the evolutionary stability of territory defence and thus territoriality itself. A more mechanistic model of territorial behaviour is needed to incorporate these kinds of mechanisms into a general theory on the evolution of territoriality.     

Spatial Distribution of Oak Mistletoe as It Relates to Habits of Oak Woodland Frugivores

This study addresses the underlying spatial distribution of oak mistletoe, Phoradendron villosum, a hemi-parasitic plant that provides a continuous supply of berries for frugivorous birds overwintering the oak savanna habitat of California''s outer coast range. As the winter community of birds consuming oak mistletoe varies from group-living territorial species to birds that roam in flocks, we asked if mistletoe volume was spatially autocorrelated at the scale of persistent territories or whether the patterns predicted by long-term territory use by western bluebirds are overcome by seed dispersal by more mobile bird species. The abundance of mistletoe was mapped on trees within a 700 ha study site in Carmel Valley, California. Spatial autocorrelation of mistletoe volume was analyzed using the variogram method and spatial distribution of oak mistletoe trees was analyzed using Ripley''s K and O-ring statistics. On a separate set of 45 trees, mistletoe volume was highly correlated with the volume of female, fruit-bearing plants, indicating that overall mistletoe volume is a good predictor of fruit availability. Variogram analysis showed that mistletoe volume was spatially autocorrelated up to approximately 250 m, a distance consistent with persistent territoriality of western bluebirds and philopatry of sons, which often breed next door to their parents and are more likely to remain home when their parents have abundant mistletoe. Using Ripley''s K and O-ring analyses, we showed that mistletoe trees were aggregated for distances up to 558 m, but for distances between 558 to 724 m the O-ring analysis deviated from Ripley''s K in showing repulsion rather than aggregation. While trees with mistletoe were aggregated at larger distances, mistletoe was spatially correlated at a smaller distance, consistent with what is expected based on persistent group territoriality of western bluebirds in winter and the extreme philopatry of their sons.     

Song Trait Similarity in Great Tits Varies with Social Structure

For many animals, long-range signalling is essential to maintain contact with conspecifics. In territorial species, individuals often have to balance signalling towards unfamiliar potential competitors (to solely broadcast territory ownership) with signalling towards familiar immediate neighbours (to also maintain so-called “dear enemy” relations). Hence, to understand how signals evolve due to these multilevel relationships, it is important to understand how general signal traits vary in relation to the overall social environment. For many territorial songbirds dawn is a key signalling period, with several neighbouring individuals singing simultaneously without immediate conflict. In this study we tested whether sharing a territory boundary, rather than spatial proximity, is related to similarity in dawn song traits between territorial great tits (Parus major) in a wild personality-typed population. We collected a large dataset of automatized dawn song recordings from 72 unique male great tits, during the fertile period of their mate, and compared specific song traits between neighbours and non-neighbours. We show here that both song rate and start time of dawn song were repeatable song traits. Moreover, neighbours were significantly more dissimilar in song rate compared to non-neighbours, while there was no effect of proximity on song rate similarity. Additionally, similarity in start time of dawn song was unrelated to sharing a territory boundary, but birds were significantly more similar in start time of dawn song when they were breeding in close proximity of each other. We suggest that the dissimilarity in dawn song rate between neighbours is either the result of neighbouring great tits actively avoiding similar song rates to possibly prevent interference, or a passive consequence of territory settlement preferences relative to the types of neighbours. Neighbourhood structuring is therefore likely to be a relevant selection pressure shaping variation in territorial birdsong.     

The Influence of Relatedness, Neighbourhood and Overall Distance on Colony Mate Recognition in a Polydomous Ant

The ‘dear enemy phenomenon’ predicts that territorial animals respond less aggressively towards more familiar neighbours than towards unfamiliar conspecifics if potential losses to strangers are more costly than potential losses to neighbours. Conversely, territorial animals should respond more aggressively to neighbours, if potential losses to them are more costly than potential losses to strangers. In social insects the question of how colony members distinguish neighbours from strangers, however, is intertwined with the more general question of how colony members discriminate themselves from non‐colony members; both genetic and spatial distance can correlate with levels of inter‐colonial aggression. In this paper I disentangle the role of experience, genetic and spatial distance on inter‐colonial aggression in a polydomous population of Iridomyrmex purpureus. In I. purpureus, aggression is related to the spatial distance between colonies irrespective of genetic similarity. Spatial distance affected aggression in two different ways. First, workers were more likely to exhibit aggression towards alien conspecifics of adjoining rather than non‐adjoining territories, suggesting the opposite of the dear enemy phenomenon. Second, workers were more often aggressive towards conspecifics of more distant colonies, implying that environmental cues play a role in the recognition system of I. purpureus.     

Avian Mobbing Response is Restricted by Territory Boundaries: Experimental Evidence from Two Species of Forest Warblers

Predator mobbing has been viewed as an adaptation to reduce the risk of predation, however, factors influencing mobbing behaviour are still debated. We report on the results of an experiment with Dendroica caerulescens and Dendroica virens designed to determine (1) whether mobbing response by forest songbirds during the breeding season is restricted by territory boundaries, (2) the distance songbirds will move in response to anti‐predator mobbing calls, and (3) whether reproductive status, age, and time of the breeding season determine the distance moved to mob. We did not detect an effect of reproductive status, age, or time of breeding season on the distance moved by birds to mob. All birds responded to the mobbing playback within their territory (defined by territorial defence in relation to specific song playbacks). The maximum distance moved within a territory to engage in mobbing ranged from 25 to 175 m ( = 72 ± 6 m). Three of 37 birds responded to playbacks outside their territory boundaries. In all three cases, maximum movement distances outside territories were short (25 m). Thus, for two species of warblers, mobbing is highly constrained by territory boundaries during the breeding season. This finding is congruent with arguments that mobbing is primarily a selfish behaviour, at least with respect to conspecifics. Our results also provide support for the ‘move‐on’ hypothesis.     

Aggregation as an antipredator strategy in the rock-paper-scissors model

We study a nonhierarchical tritrophic system, whose predator-prey interactions are described by the rock-paper-scissors game rules. In our stochastic simulations, individuals may move strategically towards the direction with more conspecifics to form clumps instead of moving aimlessly on the lattice. Considering that the conditioning to move gregariously depends on the organism's physical and cognitive abilities, we introduce a maximum distance an individual can perceive the environment and a minimum conditioning level to perform the gregarious movement. We investigate the pattern formation and compute the average size of the single-species spatial domains emerging from the grouping behaviour. The results reveal that the defence tactic reduces the predation risk significantly, being more profitable if individuals perceive further distances, thus creating bigger groups. Our outcomes show that the species with more conditioned organisms dominate the cyclic spatial game, controlling most of the territory. On the other hand, the species with fewer individuals ready to perform aggregation strategy gives its predator the chance to fill the more significant fraction of the grid. The spatial interactions assumed in our numerical experiments constitute a data set that may help biologists and data scientists understand how local interactions influence ecosystem dynamics.     

Prospectors combine social and environmental information to improve habitat selection and breeding success in the subsequent year

1.?Because habitats have profound effects on individual fitness, there is strong selection for improving the choice of breeding habitat. One possible mechanism is for individuals to use public information when prospecting future breeding sites; however, to our knowledge, no study has shown prospecting behaviour to be directly linked to subsequent choice of breeding site and future reproductive success. 2.?We collected long-term data on territory-specific prospecting behaviour and subsequent breeding in the short-lived northern wheatear (Oenanthe oenanthe). Non-breeders established prospecting territories (<2 ha) that overlapped the breeding territories of conspecifics. We tested whether: (i) prospectors used social and environmental cues that predicted territory-specific breeding success in the following year, and (ii) the prospecting territory was tightly linked to the subsequent breeding territory of the prospector, and whether this link would be weakened by intraspecific competition with original territory owners if they also survived. 3.?As expected, prospectors were attracted to a combination of site-specific cues that predicted future breeding success, i.e. short ground vegetation, a successfully breeding focal pair and successful close neighbours. 4.?Prospecting behaviour was directly linked to the choice of the following year's breeding territory: 79% of surviving prospectors established a breeding territory at their prospecting site in the following year, with their breeding success being higher than other individuals of the same age. As predicted, fidelity to the prospected site was strongly dependent on whether the original territory owner of the same sex had died or moved. 5.?Our findings suggest that the use of multiple cues reduces the negative impact of stochasticity on the reliability of social cues at small spatial scales (e.g. territories) and hence increases the probability of breeding success in the next year. Also, the use of conspecific attraction (i.e. the preference for breeding aggregations) is selectively advantageous because individuals are more likely to find a vacancy in an aggregation as compared to a solitary site. By extension, we hypothesize that species life-history traits may influence the spatial scale of prospecting behaviour and habitat selection strategies.     

海南岛霸王岭热带低地雨林树木的空间格局

树木空间格局及其形成过程是物种共存及生物多样性维持机制研究的一个重要方面。该文以海南岛两个1 hm ²的典型热带低地雨林老龄林森林动态样地为基础, 通过4个点格局模型(均质Poisson过程、异质Poisson过程、均质Thomas过程和异质Thomas过程)模拟扩散限制和生境异质性作用对树木空间分布格局的影响, 并分析不同空间尺度下(< 2 m, 2-5 m, 5-10 m, 10-15 m, 15-20 m和20-25 m)不同作用的相对重要性。结果表明: 热带低地雨林的所有树木总体上呈现聚集分布的空间格局, 随着尺度的增大, 聚集强度逐渐减小。树种在模拟空间分布格局最优模型中的比例由高到低分别是: 均质Thomas过程, 均质Poisson过程、异质Thomas过程和异质Poisson过程。扩散限制作用是形成热带低地雨林树木空间分布格局最重要的生态过程, 其次是完全随机作用以及生境异质性和扩散限制的联合作用, 而生境异质性的作用最小。不同空间尺度上模拟各树种空间分布格局的最优模型比例差异显著, 扩散限制作用能够在多数空间尺度上模拟多个树种的空间分布格局, 其次为随机作用; 生境异质性和扩散限制的联合作用主要在小尺度(0-5 m)影响树种分布, 而生境异质性在较大尺度(15-25 m)上影响树种的空间分布格局。     

Territorial intrusion risk and antipredator behaviour: a mathematical model

In territorial animals that hide to avoid predators, a predatory attack creates a conflict because a hiding animal cannot defend its territory from conspecific intruders. When intruders are persistent, a past conspecific intrusion informs a territorial resident that future intrusions by the same animal are likely. Using a mathematical model, I examine the effects that past territorial intrusions can have on antipredator behaviour. Past territorial intrusions rarely affect a resident animal's time to hide (the optimal behaviour is to hide as soon as the predator initiates its attack). In contrast, past intrusions should shorten the length of time during which territory holders remain in hiding, with the magnitude of this effect depending on the time of the predator's attack, the re-intruder's pattern of return, and the intrusion rates of other conspecifics. The results of the model show that we need more information on patterns of re-intruders' behaviour, and emphasize that a similar functional explanation could underlie other behavioural changes following territorial and/or aggressive encounters (such as winner/loser effects or changes in display frequency and territorial vigilance). Differences between my findings and those from previous studies suggest that the trade-off between antipredator behaviour and territorial defence can involve different costs from the trade-off between antipredator behaviour and foraging.     

Activity and space patterns of Pyrenean desman (Galemys pyrenaicus) suggest non-aggressive and non-territorial behaviour

The Pyrenean desman is considered a flagship species for biodiversity and evolution. However, its scientific knowledge is still under development and currently on debate, particularly in relation to its behavioural ecology and social organization. Based on the previous hypothesis of individual desmans being solitary and territorial, activity and space patterns were considered to be arranged to avoid conspecifics. However, recent insights into the species’ social behaviour revealed non-interspecific avoidance. With this study, we provide novel insights into the activity and space patterns of the desman, and their relation to its social behaviour. A total of 30 individuals were trapped, 18 of which provided informative radiotracking data to study (1) activity behaviour, (2) the proportion of the home range utilized and the distances travelled, and (3) the dominant movement directionality. Activity and space use patterns were affected by daylight and seasonality, but not by sex, age or number of other conspecifics sharing the home range. In contrast to the previous observations, individuals did not show a pattern of directionality in their movements. Noticeably, we observed encounters between individuals without evidence of aggressive behaviour. Our results suggest that desmans do not alter their activity or their spatial behaviour to avoid encounters with conspecifics, as previously suggested. These novel findings provide more evidence of a social structure and organization with social interactions and non-aggressive behaviour. This is of relevance for management actions and for the conservation of this endemic mammal.     

Interactions between Males of the Lizard Liolaemus tenuis: Roles of Familiarity and Memory

For territorial organisms, recognition of familiar individuals can reduce the frequency and intensity of aggressive encounters (‘dear enemy’ phenomenon), stabilize social systems, and reduce the cost of territory maintenance. Here, we investigated the behavioural events displayed during contests between familiar and unfamiliar individuals in the lizard Liolaemus tenuis (Liolaemidae), a species in which males are territorial. The behaviours recorded were attack, warning, evasion, and submission, and the latencies to the first aggressive (attack or warning) behaviour. Additionally, we assessed the ability of individuals to remember a familiar conspecific after a period without social interaction. Individual males reduced and delayed aggressive behaviour directed towards socially familiar individuals compared with unfamiliar ones. These results suggest that males distinguished between familiar and unfamiliar conspecific males and are in agreement with the ‘dear enemy’ phenomenon. Other behaviours were similar in the contests between familiar and unfamiliar individuals. Recognition of familiar conspecifics was lost after 20 d without social interactions. This may be relevant for interactions with floater males or with neighbours that lose their territory and subsequently attempt to fight for their ex‐neighbour's territory.     

The social ecology of the Pyrenean desman (Galemys pyrenaicus) (Insectivora: Talpidae), as revealed by radiotelemetry

The social organization, spatial utilization and movement patterns of free-ranging desmans ( Galemys pyrenaicus G.) were studied by radiotracking individuals in the French Pyrenees. During the period of study, May to July, single adult male and female desmans formed a pair and defended their shared territory from neighbouring conspecifics by vigilance and scent-marking behaviour. Other adults did not form pair bonds and remained solitary. In such pairs, the territory of the adult male was always larger than, and completely enclosed, that of the paired female. Neighbouring pairs occupied contiguous territories of similar length. The territories of neighbouring males overlapped to a small extent, but the territory of one male and that of a neighbouring female were never seen to overlap. Juveniles were observed to utilize the territory of resident pairs, prior to their dispersal. Paired adults and juveniles exploited their ranges on a regular daily basis, whilst those of solitary adults, being larger than those of the former, were utilized on a 48–hour basis. Paired males appeared to invest more time in defending the borders of their shared territory than did females which, in contrast, spent most of their time within the centre of the territory. Despite some degree of territorial overlap between neighbouring conspecifics, their mutual avoidance, achieved through a system of temporal range utilization, resulted in few agonistic encounters.     

Are Unfamiliar Neighbours Considered to Be Dear-Enemies?

Background

Discriminating threatening individuals from non-threatening ones allow territory owners to modulate their territorial responses according to the threat posed by each intruder. This ability reduces costs associated with territorial defence. Reduced aggression towards familiar adjacent neighbours, termed the dear-enemy effect, has been shown in numerous species. An important question that has never been investigated is whether territory owners perceive distant neighbours established in the same group as strangers because of their unfamiliarity, or as dear-enemies because of their group membership.

Methodology/Principal Findings

To investigate this question, we played back to male skylarks (Alauda arvensis) songs of adjacent neighbours, distant neighbours established a few territories away in the same microdialect area and strangers. Additionally, we carried out a propagation experiment to investigate how far skylark songs are propagated in their natural habitat and we estimated repertoire similarity between adjacent neighbours, distant neighbours and strangers. We show that skylarks, in the field, respond less aggressively to songs of their distant and likely unfamiliar neighbours, as shown by the propagation experiment, compared to stranger songs. The song analysis revealed that individuals share a high amount of syllables and sequences with both their adjacent and distant neighbours, but only few syllables and no sequences with strangers.

Conclusions

The observed reduction of aggression between distant neighbours thus probably results from their familiarity with the vocal group signature shared by all members of the neighbourhood. Therefore, in skylarks, dear-enemy-like relationships can be established between unfamiliar individuals who share a common acoustic code.     

Territory Establishment and Antler Cycle in Male Roe Deer

Territory establishment and antler cycle in male roe deer, Capreolus capreolus, were studied in two areas in south-eastern Sweden during 1989–1993. Initial territorial behaviour was observed in early Mar., when overlap between male ranges decreased from 66 % to 0 % over a 3–6-wk period, as determined from telemetry data. The decrease was associated with an increased number of agonistic interactions. Most territories were established at the end of Mar. to early Apr., with a tendency for males re-establishing old territories to be first. In all years, territories were taken up 1–3 wk before males shed velvet from their antlers. The distance between geometric centres of neighbouring males in winter and after territory establishment increased from 164 to 364 m. Former territory holders moved the geometric centres of their winter range and subsequent territory a shorter distance than first-year territorials, 129 and 267 m respectively. Territory defence ceased after the rut in late Aug., and the spatial relations between neighbours reverted to the situation before the territorial season. Neither velvet shedding nor antler casting was correlated with male age (yearlings excluded), which contradicts earlier statements. Neither were there any associations between timing of antler casting and subsequent velvet shedding, nor vice versa. Hence, a male casting antlers early one year derives no advantage by early shedding the next season. Furthermore, to be in hard antlers is not a prerequisite for territory take-up, whereas antlers probably are necessary for maintaining the area throughout the season. Functional aspects on roe buck territoriality are also discussed.     

Individual distance in crustacea II. the mantis shrimp Gonodactylus oerstedii

The distances of separation at which individuals of the territorial shrimp Gonodactylus oerstedii showed agonistic behaviors were measured in the laboratory. Sizes of interacting animals had no effect on distances at which behavior patterns were shown. Different acts occurred at significantly different distances.

In the case of territorial G. oerstedii, most agonistic acts occurred just as an introduced animal crossed the gravel line marking the boundary of the territory. Actual distance separating animals was less important than location of an introduced shrimp in determining the probability of agonistic interactions.

Overall, there was an inverse correlation between degree of site attachment in a species and the variance in its behavior‐distance values.     

IKONOS影像景观格局特征尺度的小波与半方差分析

建立不同方向和尺度下景观格局的小波分析方法,并以IKONOS PAN 1m 影像为例,对城市景观和农田景观格局进行了实例研究,并与半方差函数的分析结果进行比较分析。试验结果发现试验区的城市景观图像表现出不同层次和方向的结构特征,主要是阴影－单个楼房－楼房与道路的组合3个尺度的特征,分别出现在2m,16m和128m以及256m分析尺度上,农田景观主要是树木行－农田2个层次和方向尺度特征组合,分别为2m,128m和256m。半方差分析结果对试验区景观结构的多层次和方向不敏感,但可发现景观结构的总体平均特征。城市景观总体结构在空间上的变程为135m,反映的是楼房与道路组合的总体平均特征;农田景观总体结构在空间上的变程为152m。通过改变半方差函数的分析范围和计算间距可以克服半方差对多尺度分析的不足,如该试验通过缩小城市景观和农田分析范围,也发现城市景观有一个在空间结构上15m的变程,与小波分析的特征尺度16m非常接近,而农田景观出现空穴效应的周期一半长度大约为2m,正好是小波分析出种植树木的行距结构特征尺度的大小。这要分析者对研究有一定的先验知识才能够做到选择合适的分析范围,而这恰是景观格局特征尺度分析服务的目的。通过试验区研究证明小波分析是一非常有应用前景的景观格局研究方法,与半方差分析相比,对在解释多尺度结构和不同方向的结构具有明显的优势,且不受分析数据统计平稳假设的约束,但由于对图像分解是基于2进制,导致分析尺度以2的幂函数变化,不像半方差分析可通过理论模型拟合,对特征尺度的估计可以是连续的。