Wide home ranges for widely foraging lizards

Space usage by animals may be influenced by a range of factors. In this study we investigate whether foraging behaviour affects the home range size of lizards. Two distinct tactics of foraging have been recognized in predators: sit-and-wait foraging (SW) and active foraging (AF). Foraging activity level of a data set of lizard species, mainly compiled from literature, is compared with their home range sizes. Two opposite predictions can be made about foraging in connection with home range area: on the one hand, SW species may exhibit larger home ranges due to their mating system; on the other hand, AF species have higher metabolic energy and thus food requirements and can be expected to have larger home ranges that have to yield this food. This study shows that percentage of the time moving (as an index of foraging mode) correlates positively with home range, even after correcting for body mass, and these patterns remain when phylogenetic relationships are taken into account. We thus conclude that home range areas parallel activity levels in lizards.     

Spatial distribution and movements of elephants (Loxodonta africana) in the northern Namib Desert region of the Kaokoveld, South West Africa/Namibia

The home ranges, space-use patterns, seasonal distribution, long-distance movements, water-related movements and movement patterns of elephants ( Loxodonta africana ) residing in the northern Namib Desert region of the Kaokoveld, South West Africa/Namibia, are discussed. All movements of the desert-dwelling elephants, including occasional wanderings up to 195 km, were confined to the northern Namib Desert and no evidence could be found of any movements including migrations to and from the Etosha National Park. Home range size varied from 1763 to 2944 km² and the main feature of the spatial distribution of these elephants is the marked fidelity to individual home ranges. Seasonal distribution and space-use patterns within individual home ranges coincide with seasonal climatic changes and the corresponding changes in food and water availability. The elephant's normal movements and migrations were confined to individual home ranges and no mass population migrations took place. Their mobility and their ability to go up to four days without drinking water enabled them to utilize food sources up to 70 km away from waterholes and, together with their intimate knowledge about resource distribution within their home ranges, are regarded as key factors in their survival in the desert. The elephants' spatial distribution and movement patterns indicate a natural population segregation from other elephant populations in the Kaokoveld, clearly demarcating the desert-dwelling elephants as a distinct population with their own movement patterns and geographic range.     

Spatial proximity and asynchronous refuge sharing networks both explain patterns of tick genetic relatedness among lizards,but in different years

A major question for understanding the ecology of parasite infections and diseases in wildlife populations concerns the transmission pathways among hosts. Network models are increasingly used to model the transmission of infections among hosts – however, few studies have integrated host behaviour and genetic relatedness of the parasites transmitted between hosts. In a study of the Australian sleepy lizard Tiliqua rugosa and its three‐host ixodid tick (Bothriocroton hydrosauri), we asked if patterns of genetic relatedness among ticks were best explained by spatial proximity or the host transmission network. Using synchronous GPS locations of over 50 adult lizards at 10 min intervals across the three‐month activity period, over 2 years, we developed two alternative parasite transmission networks. One alternative was based on the spatial proximity of lizards (at the centre of their home ranges), and the other was based on the frequency of asynchronous shared refuge use between pairs of lizards. In each year, adult ticks were removed from lizards and their genotypes were determined at four polymorphic microsatellite loci. Adult ticks collected from the same host were more related to each other than ticks from different hosts. Similarly, adult ticks collected from different lizards had a higher relatedness if those lizards had a shorter path length connecting them on each of the two networks we explored. The predictors of tick relatedness differed between years. In the first year, the asynchronous shared refuges network was the stronger predictor of tick relatedness, whereas in year two, the spatial proximity‐based network was the stronger predictor of tick relatedness. We speculate on how changing environmental conditions might change the relative importance of alternative processes driving the transmission of parasites.     

The effect of body size on the thermoregulation of lizards on hot,dry Mediterranean islands

Body size shapes the overall biology of organisms. We assessed the impact of size on temperature regulation in populations of normal-sized and large-bodied insular Mediterranean lizards (Podarcis gaigeae, Lacertidae). We hypothesized that large lizards would achieve higher body temperatures and thermoregulate more effectively than their smaller kin. Large- and small-bodied lizards share the same thermoregulation pattern, achieving similar body temperatures in the field. Large lizards, however, prefer higher set-point temperatures. Lizards in both populations thermoregulate effectively, but large lizards thermoregulated less effectively than normal-sized lizards. The particular conditions at the islet that harbors the large-bodied population (harsh intraspecific competition) seem to account for this pattern.     

Size evolution in island lizards

Aim  The island rule, small animal gigantism and large animal dwarfism on islands, is a topic of much recent debate. While size evolution of insular lizards has been widely studied, whether or not they follow the island rule has never been investigated. I examined whether lizards show patterns consistent with the island rule.
Location  Islands worldwide.
Methods  I used literature data on the sizes of island–mainland population pairs in 59 species of lizards, spanning the entire size range of the group, and tested whether small insular lizards are larger than their mainland conspecifics and large insular lizards are smaller. I examined the influence of island area, island isolation, and dietary preferences on lizard size evolution.
Results  Using mean snout–vent length as an index of body size, I found that small lizards on islands become smaller than their mainland conspecifics, while large ones become larger still, opposite to predictions of the island rule. This was especially strong in carnivorous lizards; omnivorous and herbivorous species showed a pattern consistent with the island rule but this result was not statistically significant. No trends consistent with the island rule were found when maximum snout–vent length was used. Island area had, at best, a weak effect on body size. Using maximum snout–vent length as an index of body size resulted in most lizard populations appearing to be dwarfed on islands, but no such pattern was revealed when mean snout–vent length was used as a size index.
Main conclusions  I suggest that lizard body size is mostly influenced by resource availability, with large size allowing some lizard populations to exploit resources that are unavailable on the mainland. Lizards do not follow the island rule. Maximum snout–vent length may be biased by sampling effort, which should be taken into account when one uses this size index.     

Population genetic structure, gene flow and sex-biased dispersal in frillneck lizards (Chlamydosaurus kingii)

By using both mitochondrial and nuclear multiloci markers, we explored population genetic structure, gene flow and sex-specific dispersal of frillneck lizards ( Chlamydosaurus kingii ) sampled at three locations, separated by 10 to 50 km, in a homogenous savannah woodland in tropical Australia. Apart from a recombinant lizard, the mitochondrial analyses revealed two nonoverlapping haplotypes/populations, while the nuclear markers showed that the frillneck lizards represented three separate clusters/populations. Due to the small population size of the mtDNA, fixation may occur via founder effects and/or drift. We therefore suggest that either of these two processes, or a combination of the two, are the most likely causes of the discordant results obtained from the mitochondrial and the nuclear markers. In contrast to the nonoverlapping mitochondrial haplotypes, in 12 out of 74 lizards, mixed nuclear genotypes were observed, hence revealing a limited nuclear gene flow. Although gene flow should ultimately result in a blending of the populations, we propose that the distinct nuclear population structure is maintained by frequent fires resulting in local bottlenecks, and concomitant spatial separation of the frillneck lizard populations. Limited mark–recapture data and the difference in distribution of the mitochondrial and nuclear markers suggest that the mixed nuclear genotypes were caused by juvenile male-biased dispersal.     

Spatial ecology of bluetongue lizards (Tiliqua spp.) in the Australian wet–dry tropics

New technologies for quantifying animal locations enable us to document habitat‐selection patterns of cryptic taxa in extraordinary detail. Northern bluetongues (Tiliqua scincoides intermedia) and centralian bluetongues (Tiliqua multifasciata) are large heavy‐bodied scincid lizards that are broadly sympatric in the wet–dry tropics of north‐western Australia. We used data from GPS‐based radiotelemetry (n = 49 lizards, tracked for 2–121 days, total n = 61 640 locations) to examine the size, internal structure and overlap of lizard home ranges. Despite substantial habitat differences at our two study sites (semi‐arid and relatively pristine habitat at Keep River National Park, Northern Territory, vs. highly disturbed and fragmented flood plain habitat in an agricultural area near Kununurra, Western Australia), home ranges were similar between the two areas, and between the two species. Our radio‐tracked lizards continued to disperse into previously unused areas throughout the duration of the study, so that the total areas used by lizards continued to increase. Based on the minimum convex polygon method, total home ranges averaged 4 ha (range 2–12 ha), but only about two‐thirds of each home range was used intensively. Each home range had multiple core areas, and overlap of core as well as peripheral areas (especially with same‐sex conspecifics) was high at the disturbed (Western Australia) site where lizard densities were high. The concentration of lizard activity within small core areas, often used by multiple individuals, suggests that these heavily used sites are critical to lizard conservation. However, the lizards' infrequent long‐distance displacements also make them vulnerable to changes in the wider landscape mosaic. Because GPS‐based radiotelemetry can quantify habitat use at finer spatial and temporal scales than earlier technologies, it can provide a robust base for management of at‐risk fauna.     

中国蜥蜴类生活史和生态学特征数据集

截至2020年底, 中国共有226种蜥蜴类(不包括外来入侵种), 是世界上蜥蜴类多样性最丰富的国家之一。系统整理中国现有蜥蜴类的特征数据在物种起源与进化、形成与灭绝、保护生物学等研究中具有重要意义。但是, 目前还没有关于我国蜥蜴类生活史、生态学和地理分布等物种特征的完整数据库。本文通过系统查阅文献和数据资料, 共收集整理了中国现有226种本土蜥蜴类19个特征数据: 描述年份、中国受威胁等级、全球受威胁等级、是否中国特有种、是否岛屿特有种、平均体长、平均体重、食性、窝卵数、繁殖模式、四肢发育、活动时间、栖息生境、栖息地类型、栖息地宽度、海拔分布范围、地理分布范围、动物地理界和分布省份。在上述特征中, 除了四肢发育、描述年份、是否中国特有种、是否岛屿特有种和分布省份外, 其余特征数据均存在不同程度的缺失, 数据完整度为47.14%-100%。本数据集是目前关于中国蜥蜴类最新和最全的物种特征数据库, 可为我国蜥蜴类生态学、进化生物学、生物地理学和保护生物学等研究领域提供数据支持。     

Size but not relatedness drives the spatial distribution of males within an urban population of Anolis carolinensis lizards

The way that individuals are spatially organized in their environment is a fundamental population characteristic affecting social structure, mating system, and reproductive ecology. However, for many small or cryptic species, the factors driving the spatial distribution of individuals within a population are poorly understood and difficult to quantify. We combined microsatellite data, remote sensing, and mark–recapture techniques to test the relative importance of body size and relatedness in determining the spatial distribution of male Anolis carolinensis individuals within a focal population over a five‐year period. We found that males maintain smaller home ranges than females. We found no relationship between male body size and home range size, nor any substantial impact of relatedness on the geographic proximity. Instead, the main driver of male spatial distribution in this population was differences in body size. We also found no evidence for offspring inheritance of their parent''s territories. Males were never sampled within their father''s territory providing strong support for male‐biased dispersal. This study introduces a novel approach by combining standard mark release capture data with measures of pairwise relatedness, body size, and GPS locations to better understand the factors that drive the spatial distribution of individuals within a population.     

An allometric model of home range formation explains the structuring of animal communities exploiting heterogeneous resources

Understanding and predicting the composition and spatial structure of communities is a central challenge in ecology. An important structural property of animal communities is the distribution of individual home ranges. Home range formation is controlled by resource heterogeneity, the physiology and behaviour of individual animals, and their intra‐ and interspecific interactions. However, a quantitative mechanistic understanding of how home range formation influences community composition is still lacking. To explore the link between home range formation and community composition in heterogeneous landscapes we combine allometric relationships for physiological properties with an algorithm that selects optimal home ranges given locomotion costs, resource depletion and competition in a spatially‐explicit individual‐based modelling framework. From a spatial distribution of resources and an input distribution of animal body mass, our model predicts the size and location of individual home ranges as well as the individual size distribution (ISD) in an animal community. For a broad range of body mass input distributions, including empirical body mass distributions of North American and Australian mammals, our model predictions agree with independent data on the body mass scaling of home range size and individual abundance in terrestrial mammals. Model predictions are also robust against variation in habitat productivity and landscape heterogeneity. The combination of allometric relationships for locomotion costs and resource needs with resource competition in an optimal foraging framework enables us to scale from individual properties to the structure of animal communities in heterogeneous landscapes. The proposed spatially‐explicit modelling concept not only allows for detailed investigation of landscape effects on animal communities, but also provides novel insights into the mechanisms by which resource competition in space shapes animal communities.     

Home ranges of parthenogenetic and bisexual species in a community of Darevskia lizards (Reptilia: Lacertidae)

We analysed the home ranges of a community of Darevskia rock lizards composed of a bisexual species (D. valentini), two parthenogens (D. armeniaca and D. unisexualis), and two backcross forms between bisexual and unisexual forms. We estimated home range areas of ink-marked, GPS-located lizards using Minimum Convex Polygon (MCP) and 95% of the locations for those individuals with five or more sightings. The bisexual D. valentini was the species with the largest home ranges, distances travelled, and the most intersections. No differences between unisexual species and backcrosses were recorded for any comparison. In males, home range size and perimeter were related to morphological characteristics. Contrary to what has been described in allopatry, unisexual species showed smaller home ranges and fewer overlaps than sympatric bisexual species. We tentatively suggest that the presence of potential bisexual partners might increase sexual competition among parthenogenetic females while differences in habitat use should also be considered.     

Seed dispersal by lizards on a continental‐shelf island: predicting interspecific variation in seed rain based on plant distribution and lizard movement patterns

Aim We estimated the patterns of seed deposition provided by the eyed lizard, Timon lepidus, and evaluated whether these patterns can be generalized across plant species with different traits (fruit and seed size) and spatial distributions. Location Monteagudo Island, Atlantic Islands National Park (north‐western Spain). Methods We radio‐tracked seven lizards for 14 days and estimated their home ranges using fixed kernels. We also geo‐referenced all fruit‐bearing individuals of four plant species dispersed by eyed lizards in the study area (Corema album, Osyris alba, Rubus ulmifolius and Tamus communis), measured the passage time of their seeds through the lizard gut, and estimated seed predation in four habitats (bare sand, grassland, shrub and gorse). Seed dispersal kernels were estimated using a combination of these data and were combined with seed predation probability maps to incorporate post‐dispersal seed fate (‘seed survival kernels’). Results Median seed gut‐passage times were around 52–98 h, with maximum values up to 250 h. Lizards achieved maximum displacement in their home ranges within 24–48 h. Seed predation was high (80–100% of seeds in 2 months), particularly under Corema shrub and gorse. Seed dispersal kernels showed a common pattern, with two areas of preferential seed deposition, but the importance of these varied among plant species. Interspecific differences among dispersal kernels were strongly reduced by post‐dispersal seed predation; hence, seed survival kernels of the different plant species showed high auto‐ and pairwise‐correlations at small distances (< 50 m). As a result, survival to post‐dispersal seed predation increased with dispersal distance for O. alba and T. communis, but not for C. album. Main conclusions Seed dispersal by lizards was determined primarily by the interaction between the dispersers’ home ranges and the position of the fruit‐bearing plants. As a result, seed rain shared a common template, but showed considerable variation among species, determined by their specific spatial context. Seed predation increased the spatial coherence of the seed rain of the different species, but also resulted in contrasting relationships between seed survival and dispersal distance, which may be of importance for the demographic and evolutionary processes of the plants.     

Size distributions and sex ratios of colonizing lizards

This paper reports the body size distributions and sex ratios of four species of phyrnosomatid lizard that colonized experimentally created density sinks. The experiments were conducted in western Colorado in 1992, and lizards colonizing the habitats in 1993 were compared to those removed in 1992 and those present in 1991. Lizards colonizing the density sinks were able to disperse from adjacent habitat. For two of the species (Urosaurus ornatus and Sceloporus undulatus), colonizing lizards were significantly smaller than either those removed in 1992 or those present in 1991. Two other species (S. graciosus and Uta stansburiana) showed no difference in the size distribution of colonizing and removed lizards. In addition, sex ratios of colonizing lizards did not differ from those removed in 1992 or present in 1991. The results of the experiments have implications for the dynamics of the target populations, rescue of local populations from extinction, the regional persistence of populations subject to turnover in patchy environments, and priority effects in colonization. Received: 3 February 1997 / Accepted: 20 May 1998     

Global warming and positive fitness response in mountain populations of common lizards Lacerta vivipara

Recent global warming threatens many species and has already caused population‐ and species‐level extinctions. In particular, high risks of extinction are expected for isolated populations of species with low dispersal abilities. These predictions rely on widely used ‘climatic envelope’ models, while individual responses, the ultimate driver of a species response to climate change, have been most often neglected. Here, we report on some changes in life‐history traits of a dispersal‐limited reptile species (a poorly studied taxa) living in isolated populations. Using long‐term data on common lizards collected in southern France, we show that individual body size dramatically increased in all the four populations studied over the past 18 years. This increase in body size in all age classes appeared related to a concomitant increase in temperature experienced during the first month of life (August). Daily maximum temperature in August increased by 2.2°C and yearling snout‐vent‐length increased by about 28%. As a result, adult female body size increased markedly, and, as fecundity is strongly dependent on female body size, clutch size and total reproductive output also increased. For one population where capture–recapture data were available, adult survival was positively related to May temperature. All fitness components investigated therefore responded positively to the increase in temperature, such that it might be concluded that the common lizard has been advantaged by the shift in temperature. We contrast these short‐term results with the long‐term habitat‐based prediction that these populations located close to mountain tops on the southern margin of the species range should be unable to cope with the alteration of their habitat. To achieve a better prediction of a species persistence, one will probably need to combine both habitat and individual‐based approaches.     

Home ranges of introduced rats on Christmas Island: A pilot study

Understanding the spatial ecology of invasive rats (Rattus spp.) is necessary to inform management actions to reduce their impact on native flora and fauna. This study investigates home range sizes of exotic rats around seabird colonies and urban areas on Christmas Island, where rat predation is suspected to be adversely affecting fledgling success among local seabirds. It was hypothesised that rat home range sizes would be smaller in urban areas owing to more consistent food availability. Home ranges of male rats were significantly larger compared with their female counterparts, with male rats maintaining larger home ranges in urban areas compared with seabird colonies. Conversely, female rats had smaller home ranges in urban areas compared with seabird colonies. Our findings suggest a possible correlation between the spatial distribution of food resources and home range size. Additionally, the spatial distribution of breeding females across the landscape had a significant influence on the home ranges of male rats. These findings have important implications for proposed efforts to manage rat populations on Christmas Island, while also providing valuable information regarding the ecology of invasive rats on tropical islands.     

Territorial behavior of the plethodontid salamander Plethodon kentucki: influence of habitat structure and population density

To investigate the possible influence of variation in ecological and demographic factors on the spatial organization of the terrestrial plethodontid salamander Plethodon kentucki, I conducted a 3-year capture-recapture study and determined home-range characteristics and spatial relationships of individuals at two field sites that differed in predominant cover type and population density. Home ranges of adults were fixed and the home ranges of same-sex adult neighbors were mostly exclusive. The spatial arrangement of adult home ranges exhibited overall regularity or regularity within aggregations, whereas the distribution of juvenile home ranges was usually random. Analysis of nearest-neighbor sex indicated a positive intersexual association of adult home ranges. Removal studies provided evidence for defense of adult home ranges only at the high-density site. The distribution of home ranges was influenced by the presence of cover objects, but there was no significant relationship between adult body size and percent of home-range area with cover. Males overlapped the home ranges of gravid females significantly more often than those of non-gravid females, indicating that the distribution of gravid females had a strong influence on the distribution of male home ranges. In laboratory tests, increased male-male aggression during the breeding season suggests that males may compete for access to mates. At the high-density site, larger males may have benefited by having greater reproductive success than smaller males because they were more dominant and their home ranges overlapped a greater number of gravid-female home ranges. My results indicate that habitat structure and population density may influence the spatial organization and mating system of P. kentucki. Received: 26 May 1997 / Accepted: 9 October 1997     

青海高原两种沙蜥mtDNA的渐渗杂交

青海沙蜥和贵德沙蜥在共和沙珠玉地区重叠分布,形态和遗传显著不同。线粒体基因序列分析说明在重叠区两种蜥蜴mtDNA基因相互渗透。重叠区19个青海沙蜥个体中有1个具有贵德沙蜥线粒体基因型,但形态与青海沙蜥相似。6个贵德沙蜥个体中有1个具有青海沙蜥线粒体基因型,形态与贵德沙蜥相似。两种沙蜥在该地区的基因流动可能发生在共和古湖的退缩消失后次级相遇形成的。分子钟推测杂交发生的时间大约在19至16万年前,与古湖的消失时间(15万年前)相符。     

Field studies on a social lizard: Home range and social organization in an Australian skink,Egernia major

Abstract Although most reptiles have polygynous mating systems without long‐term pair bonds, one lineage of large scincid lizards in Australia is exceptional in this respect. Reports of complex sociality in the genus Egernia led us to conduct the first radiotelemetric field study of a species within this group. Land mullets (Egernia major) are large (60 cm total length), viviparous lizards from rainforest habitats in south‐eastern Australia. To document the spatial ecology and social organization of this species, we captured 12 adult lizards in the Barrington Tops area of eastern New South Wales and implanted them with miniature radiotransmitters. The lizards were released at their sites of capture and located daily for the next 6 weeks. All of the radiotracked lizards had discrete home ranges of approximately 10 000 m², based around well‐defined core areas (approximately 2000–3000 m²). Females tended to move further, and to range over wider areas, than did males. All of the radiotracked lizards lived in social groups consisting of one or more adult males and females plus juveniles of all age classes. Subgroups were apparent within one group of five radio‐tagged lizards: individual animals consistently shared their shelter sites and home ranges with one or more specific individuals. Male/female pairings were more frequent than expected under the null hypothesis of random association among individuals. The data in the present study support anecdotal reports of pair bonds in E. major and support suggestions that the social systems of species in this genus are more complex than those of previously studied reptiles.     

What determines variation in home range size across spatiotemporal scales in a large browsing herbivore?

1. Most studies of intraspecific variation in home range size have investigated only a single or a few factors and often at one specific scale. However, considering multiple spatial and temporal scales when defining a home range is important as mechanisms that affect variation in home range size may differ depending on the scale under investigation. 2. We aim to quantify the relative effect of various individual, forage and climatic determinants of variation in home range size across multiple spatiotemporal scales in a large browsing herbivore, the moose (Alces alces), living at the southern limit of its distribution in Norway. 3. Total home range size and core home range areas were estimated for daily to monthly scales in summer and winter using both local convex hull (LoCoH) and fixed kernel home range methods. Variance in home range size was analysed using linear mixed-effects models for repeated measurements. 4. Reproductive status was the most influential individual-level factor explaining variance in moose home range size, with females accompanied by a calf having smaller summer ranges across all scales. Variation in home range size was strongly correlated with spatiotemporal changes in quantity and quality of natural food resources. Home range size decreased with increasing browse density at daily scales, but the relationship changed to positive at longer temporal scales. In contrast, browse quality was consistently negatively correlated with home range size except at the monthly scale during winter when depletion of high-quality forage occurs. Local climate affected total home range size more than core areas. Temperature, precipitation and snow depth influenced home range size directly at short temporal scales. 5. The relative effects of intrinsic and extrinsic determinants of variation in home range size differed with spatiotemporal scale, providing clear evidence that home range size is scale dependent in this large browser. Insight into the behavioural responses of populations to climatic stochasticity and forage variability is essential in view of current and future climate change, especially for populations with thermoregulatory restrictions living at the southern limit of their distribution.     

栖息地荒漠化对草原沙蜥食性的影响

栖息地退化会使环境中蜥蜴的可利用食物资源发生改变。采用剖胃法对内蒙古自治区鄂尔多斯市准格尔旗和达拉特旗地区采集的草原沙蜥样本进行了食性分析。对固定沙丘、半固定沙丘和流动沙丘三类样地中77个实胃样本的652只(套)食物进行鉴定和分析的结果表明,这些食物主要为节肢动物,分为35类,隶属3纲11目32科;也在少部分个体(4.69%)的胃容物中发现少量(1.95%)植物碎片。上述三类栖息地中,草原沙蜥对蚁科、蚜科、瓢虫科、叶蝉科和茧蜂科昆虫均有较大的捕食比例;但相比固定沙区栖息地,草原沙蜥在半固定及流动沙丘中增减的食物种类均比较多,导致草原沙蜥在流动沙区与其他两类样地之间有较大的食物相异性。从固定沙丘、半固定沙丘到流动沙丘,随栖息地荒漠化程度的增加,草原沙蜥捕食食物类群的丰富度逐步减少,辛普森优势度指数相应地下降,但营养生态位宽度、香农威纳多样性指数和皮洛均匀性指数却在流动沙丘栖息地上明显增大。这表明鄂尔多斯地区的草原沙蜥是以捕食昆虫等节肢动物为主、兼食极少量植物的杂食性动物。栖息地荒漠化所致的环境变化显著地影响草原沙蜥的食物组成,使其觅食的食物种类逐步减少。