Assisted colonization: Integrating conservation strategies in the face of climate change

Global climate change poses an immense challenge for conservation biologists seeking to mitigate impacts to species and ecosystems. Species persistence will depend on geographic range shifts or adaptation in response to warming patterns as novel climates and community assemblages arise. Assisted colonization has been proposed as a method for addressing these challenges. This technique, which consists of transporting species to a new range that is predicted to be favorable for persistence under future climate scenarios, has become the subject of controversy and discussion in the conservation community due to its highly manipulative nature, questions about widespread feasibility, and uncertainty associated with the likelihood of translocated species becoming invasive. We reviewed the discussion and criticism associated with assisted colonization and sought to identify other conservation techniques that also display potential to promote the colonization and adaptation of species in response to climate change. We propose an integrated conservation strategy that includes management for habitat connectivity, conservation genetics, and when necessary, assisted colonization of species that are still unable to shift their ranges even given implementation of the above standard conservation approaches. We argue that this integrated approach will facilitate persistence for a larger proportion of species than is possible by solely using assisted colonization. Furthermore, a multi-faceted approach will likely reduce the uncertainty of conservation outcomes and will become increasingly necessary for conservation of biodiversity in a changing climate.     

Role of climatic niche models in focal-species-based conservation planning: Assessing potential effects of climate change on Northern Spotted Owl in the Pacific Northwest, USA

Although well-studied vertebrates such as the Northern Spotted Owl (NSO) are often used as focal species in regional conservation plans, range shifts associated with climate change may compromise this role. I used the Maxent (maximum entropy) method to develop NSO distribution models from data on NSO locations, forest age, and an ensemble of climate projections. NSO presence was positively associated with the proportion of old and mature forest at two spatial scales. Winter precipitation was the most important climate variable, consistent with previous studies suggesting negative effects on survival and recruitment. Model results suggest that initial niche expansion may be followed by a contraction as climate change intensifies, but this prediction is uncertain due to variability in predicted changes in precipitation between climate projections. Although new reserves created by the US Northwest Forest Plan prioritized areas with greater biological importance for the NSO than did pre-existing reserves, the latter areas, which lie predominantly at higher elevations, increase in importance under climate change. In contrast with previous analyses of the region’s localized old-forest-associated species, vegetation rather than climate dominated NSO distribution models. Rigorous assessment of the implications of climate change for focal species requires development of dynamic vegetation models that incorporate effects of competitor species and altered disturbance regimes. The results suggest that, lacking such data, models that combine climate data with current data on habitat factors such as vegetation can inform conservation planning by providing less-biased estimates of potential range shifts than do niche models based on climate variables alone.     

Adapting global conservation strategies to climate change at the European scale: The otter as a flagship species

Climate change has created the need for new strategies in conservation planning that account for the dynamics of factors threatening endangered species.Here we assessed climate change threat to the European otter, a flagship species for freshwater ecosystems, considering how current conservation areas will perform in preserving the species in a climatically changed future. We used an ensemble forecasting approach considering six modelling techniques applied to eleven subsets of otter occurrences across Europe. We performed a pseudo-independent and an internal evaluation of predictions. Future projections of species distribution were made considering the A2 and B2 scenarios for 2080 across three climate models: CCCMA-CGCM2, CSIRO-MK2 and HCCPR HADCM3. The current and the predicted otter distributions were used to identify priority areas for the conservation of the species, and overlapped to existing network of protected areas.Our projections show that climate change may profoundly reshuffle the otter’s potential distribution in Europe, with important differences between the two scenarios we considered. Overall, the priority areas for conservation of the otter in Europe appear to be unevenly covered by the existing network of protected areas, with the current conservation efforts being insufficient in most cases. For a better conservation, the existing protected areas should be integrated within a more general conservation and management strategy incorporating climate change projections. Due to the important role that the otter plays for freshwater habitats, our study further highlights the potential sensitivity of freshwater habitats in Europe to climate change.     

Identifying priority areas for bioclimatic representation under climate change: a case study for Proteaceae in the Cape Floristic Region, South Africa

Biological reserves are established to protect natural resources and represent the diversity of environments found within a region. Unfortunately, many systems of protected areas do not proportionally capture the range of environmental conditions occupied by species and communities. Combinations of habitat loss and climate change may exacerbate these representational biases, and result in future distributions of environmental conditions that bare little resemblance to historic patterns. New protected areas need to be established to correct existing biases, and create conservation networks that remain representative despite climate change, habitat loss, and changes in species distributions. We demonstrate a new method to identify and prioritize habitat based on its value for improving bioclimatic representation. We assessed representation provided by existing protected areas for 301 Proteaceae species under historic and projected 2050 climate across the Cape Floristic Region in South Africa. The existing reserve system has relatively modest biases with respect to current species distributions and climate. However, if the system is not supplemented, protected areas in 2050 will capture an increasingly skewed sample of climatic conditions occupied by Proteaceae. These biases can be repaired through the systematic establishment of new protected areas, and many of the most valuable areas coincide with high priority ecosystem components and irreplaceable elements identified in the Cape Action for People and the Environmental conservation plan. Protecting these areas achieves nearly the best possible improvement in climatic representation while also meeting biodiversity representation goals.     

Evaluating population persistence of Delmarva fox squirrels and potential impacts of climate change

In addition to the combined effects of forest fragmentation, habitat loss, and population isolation on the long-term persistence of many species including the endangered Delmarva fox squirrel (Sciurus niger cinereus), future changes in climate may make existing habitats less productive and more variable. The Palmer Drought Severity Index (PDSI) for the Delmarva Peninsula of the mid-Atlantic USA, reveals a trend for longer durations of potentially unfavorable conditions for fox squirrel population growth. We used a stochastic population matrix model and available life history information to assess population extinction risk for the Delmarva fox squirrel under a number of scenarios of landscape change and environmental variation, including uncertainties in the future range of climate patterns. Patch size (carrying capacity) was the most important factor influencing persistence of isolated populations. Extinction risk increased markedly across all patch sizes when year to year patterns in environmental variability were autocorrelated to match regional patterns in the PDSI. Increased autocorrelation matching the regional PSDI increased extinction risk, ranging from a factor of 5 to a factor of over 100 in some scenarios when compared to uncorrelated patterns in environmental variability. Increasing the range of variation in survival probabilities was less important to persistence, but its effect also increased when year-to-year changes were autocorrelated in time. Comparing model results with the size and landscape configuration of currently occupied patches on the Delmarva Peninsula showed that many existing populations are above the size threshold identified by these simulations for long-term persistence under current conditions, but these may become vulnerable should climate variability increase and adverse conditions persist for several years at a time.     

Conservation strategies in response to rapid climate change: Australia as a case study

As in all parts of the globe, rapid climate change in Australia will have significant negative impacts on biodiversity. It also will interact with pre-existing stressors such as native vegetation clearing, altered natural disturbance regimes and invasive species - all of which already have major negative effects on biota in Australia. Strategies to reduce climate change impacts on Australian biodiversity include a mixture of mitigation and adaptation actions (sensuMillar et al., 2007) such as: (1) significantly reducing greenhouse gas emissions, (2) ensuring bio-diverse carbon capture, (3) better tackling pre-existing stressors on biodiversity, (4) better preparing for the effects of major natural disturbances, (5) significantly improving off-reserve conservation efforts including fostering appropriate connectivity, and (6) enhancing the existing reserve system by making it more comprehensive, adequate and representative. The first strategy above demands a global response otherwise major mitigation attempts in Australia that are not paralleled elsewhere around the world will have little effect on climate change and, in turn, contribute little to enhanced biodiversity conservation. Strategies 2-6 demand multi-scaled responses, particularly at a regional level, given the major regional differences in direct climate change impacts and their interactions with pre-existing regional stressors. Well developed multi-scaled conservation plans to implement these strategies currently do not exist, nor do appropriate institutional arrangements and capacities. Institutional reforms are urgently needed in Australia to develop the land management, monitoring and regional response capabilities required to conserve biodiversity on a continent already significantly modified.     

The impact of climate change on cherry trees and other species in Japan

Studies from throughout the world have provided evidence that climate change is already affecting the ecology and persistence of species. Japan contributes valuable insights into understanding these changes through long-term records of the timing of cherry blossoms and other phenomena of ecological and cultural interest. The wide latitudinal spread of Japan also provides an opportunity to examine species over a broad range of environmental conditions. In this review, we examine phenological studies that have taken place in Japan and summarize their significance to climate change and conservation research. In Kyoto, records of the timing of celebrations of cherry blossom festivals going back to the 9th century reconstruct the past climate and demonstrate the local increase in temperature associated with global warming and urbanization. This record is probably the longest annual record of phenology from anyplace in the world and shows that cherries are currently flowering earlier than they have at any time during the previous 1200 years. Detailed mapping of cherry tree flowering times in and around Osaka and other cities in Japan show that urbanization causes plants to flower earlier within the city environs than in nearby parks and outlying suburban areas. Flowering records from a large cherry arboretum at Mt. Takao, on the outskirts of Tokyo, show that both among and within species, early flowering is associated with greater responsiveness to temperature variation. Lastly, records of phenology for a wide range of plants and animals recorded at over 100 weather stations throughout Japan show that species vary greatly in their phenological responses to climate change, and highlight ecological relationships that need to be investigated in the field. Together, these studies provide important insights into how species are responding to climate change in Japan. Further research, particularly targeted field observations and experiments, is needed to build on these findings and to improve our understanding of how climate change is altering biological communities and how it will continue to affect them in the future.     

气候变化对黄河中上游地区潜在蒸散影响的估算

采用Penman—Monteith公式计算潜在蒸散,分析1961-2010年黄河中上游地区气候因子的变化趋势,并用偏微分方法定量分析各气候因子变化对潜在蒸散的影响。结果显示：1）近50年来黄河中上游地区气温显著升高（α〈0．01）,风速和日照时间显著减少（α〈0．01）。半干旱区面积不断扩大,气候总体呈暖干变化趋势。2）潜在蒸散对实际水汽压变化最敏感,其次为最高气温、风速和日照时间,对最低气温变化最不敏感。夏季潜在蒸散对日最高气温最敏感,其他季节对实际水汽压最敏感。3）风速对年潜在蒸散变化贡献最大,其次为气温和日照时间,实际水汽压贡献最小。春夏秋冬对潜在蒸散变化贡献最大的气候因子依次为风速、日照时间、气温。温度升高、湿度减少埘潜在蒸散的正影响已被风速、辐射下降的负影响所抵消。综合考虑各气候因子对潜在蒸散的作用有助于加深对气候变化影响机制的认识,并为气候变化适应对策研究提供科学依据。     

Modelling the effect of environmental change on shorebirds: A case study on Poole Harbour, UK

An individuals-based model, MORPH, was used to assess the quality of Poole Harbour, UK, for five overwintering shorebirds: dunlin Calidris alpina, redshank Tringa totanus, black-tailed godwit Limosa limosa, oystercatcher Haematopus ostralegus and curlew Numenius arquata. Site quality, and the effect of environmental change, was measured as predicted overwinter survival. Dunlin had the highest prey biomass densities and were the least likely to be affected by reductions in their food supply, lower temperatures or loss of terrestrial habitats. Black-tailed godwits and curlew had the lowest prey biomass densities and were the most likely to be affected by reductions in their food supply, lower temperatures and loss of terrestrial habitats. All five shorebird species were seriously affected by simulated sea-level rise. Conservation issues identified for the Poole Harbour SPA were the relatively low densities of larger size classes of polychaete worms, the importance of maintaining and managing surrounding terrestrial habitats and the effect of sea-level rise on the length of time for which intertidal food supplies are available.     

Long-distance plant dispersal and habitat fragmentation: identifying conservation targets for spatial landscape planning under climate change

Climate change presents a potentially severe threat to biodiversity. Species will be required to disperse rapidly through fragmented landscapes in order to keep pace with the changing climate. An important challenge for conservation is therefore to manage landscapes so as to assist species in tracking the environmental conditions to which they are adapted. Here we develop a stochastic spatially explicit model to simulate plant dispersal across artificial fragmented landscapes. Based on certain assumptions as to the dispersal mechanism, we assess the impact that varying potential for rare long-distance dispersal (LDD) has on the ability to move over landscapes with differing spatial arrangements of suitable habitat (clumped versus fragmented). Simulations demonstrate how the relative importance of landscape structure in determining migration ability may decrease as the potential for LDD increases. Thus, if LDD is the principal mechanism by which rapid large-scale migrations are achieved, strategically planned networks of protected habitat may have a limited impact on rates of large-scale plant migrations. We relate our results to conventional principles for conservation planning and the geometric design of reserves, and demonstrate how reversal of these principles may maximise the potential for conservation under future climates. In particular, we caution against the justification of large-scale corridors on grounds of climate change since migration along corridors by standard dispersal mechanisms is unlikely to keep pace with projected change for many species. An improved understanding of the dispersal mechanisms by which species achieve rapid migrations, and the way that these processes are affected by patterns of landscape fragmentation, will be important to inform future conservation strategies.     

A review of conservation threats on Gough Island: a case study for terrestrial conservation in the Southern Oceans

Gough Island is a remote Southern Ocean Island that, despite having no permanent human population, is under substantial conservation threat as a result of human activity. A considerable proportion of the flora and fauna has been accidentally introduced, and new data are presented showing that ca. 70% of the free-living pterygote insect species are introductions. We describe how endangered seabirds that breed on the island may suffer from human fisheries activities and present new evidence showing that local temperatures have risen significantly since 1963, threatening to alter the architecture and composition of species communities. These observations are an indication that the terrestrial ecosystems of other remote islands in the Southern Oceans may be under greater conservation threat than previously thought. In light of the threats described, we discuss conservation management priorities for Gough Island. To facilitate conservation of the indigenous biota, and that of similar islands elsewhere, we suggest that a combination of proactive measures designed to maintain the integrity of island communities, and monitoring programs designed rapidly to identify new conservation threats, should be implemented conscientiously.     

Simulating site-specific impacts of climate change on soil erosion and surface hydrology in southern Loess Plateau of China

Proper spatial and temporal treatments of climate change scenarios projected by General Circulation Models (GCMs) are critical to accurate assessment of climatic impacts on natural resources and ecosystems. The objective of this study was to evaluate the site-specific impacts of climate change on soil erosion and surface hydrology at the Changwu station of Shaanxi, China using a new spatiotemporal downscaling method. The Water Erosion Prediction Project (WEPP) model and climate change scenarios projected by the U.K. Hadley Centre's GCM (HadCM3) under the A2, B2, and GGa emissions scenarios were used in this study. The monthly precipitation and temperature projections were downloaded for the periods of 1900–1999 and 2010–2039 for the grid box containing the Changwu station. Univariate transfer functions were derived by matching probability distributions between station-measured and GCM-projected monthly precipitation and temperature for the 1950–1999 period. The derived functions were used to spatially downscale the GCM monthly projections of 2010–2039 in the grid box to the Changwu station. The downscaled monthly data were further disaggregated to daily weather series using a stochastic weather generator (CLIGEN). The HadCM3 projected that average annual precipitation during 2010–2039 would increase by 4 to 18% at Changwu and that frequency and intensity of large storms would also increase. Under the conventional tillage, simulated percent increases during 2010–2039, compared with the present climate, would be 49–112% for runoff and 31–167% for soil loss. However, simulated soil losses under the conservation tillage during 2010–2039 would be reduced by 39–51% compared with those under the conventional tillage in the present climate. The considerable reduction in soil loss in the conservation tillage indicates the importance of adopting conservation tillage in the region to control soil erosion under climate change.     

The potential effects of climate change on autumn soil tillage opportunities in England and Wales

A model was used to estimate the number of autumn machinery work-days in England and Wales for a range of potential climate change sensitivity tests. The sensitivity tests were based on current best estimates of climate change and were referenced to a 30-year baseline climate. The analysis showed that decreases in precipitation have a much greater effect on the average number of work-days than increases in precipitation of a similar magnitude or increases in temperature of between 1 and 2°C. A 2°C increase in temperature was found to be off-set by a 10% increase in precipitation resulting in little change to the number of work-days compared with the baseline. The results suggested that autumn soil tillage opportunities will be improved by global warming unless precipitation increases substantially (by 15% or more). However, there are likely to be strong regional differences in the magnitude of this improvement.     

气候变化背景下安徽省冬小麦气候生产潜力和胁迫风险研究

从气候的资源和灾害双重属性出发,构建了冬小麦气候生产潜力和胁迫风险评价指标,以安徽省为例分析了二者对气候变化的响应特征,综合气候对高产和稳产的影响进行研究区冬小麦种植气候适宜性区划。结果表明:采用逐级订正法结合作物生长动态参数估算安徽省冬小麦气候生产潜力多年平均为12 391kg?hm-2,以沿淮和江淮之间最高;1961—2015年淮北和沿淮东部地区为显著上升趋势,而淮河以南地区则以下降为主。通过考虑在冬小麦生长发育过程中气候条件偏离最适区间而导致的胁迫影响,建立了高温、低温、雨涝、干旱4种气候胁迫的评估指标,并基于气候胁迫的超越概率形成了冬小麦气候风险评价方法。气候变暖使研究区冬小麦高温胁迫显著上升,低温胁迫显著下降,水分胁迫无显著的变化趋势。安徽省冬小麦的气候风险呈现中间低,两头高的分布特征,以沿淮和江淮之间风险最低,淮北北部和江南南部风险较高;淮北地区主要以干旱和低温贡献为主,而淮河以南地区则以雨涝风险为主。融合气候生产潜力和气候胁迫风险形成冬小麦的气候适宜性区划,其空间格局呈南北低、中间高的特征,种植分布格局与气候适宜性的空间匹配程度较高,但有一定的优化调整空间。     

Augmenting freshwater availability in mountain headwater streams: Assessing the sustainability under baseline and future climate change scenarios

Mountain headwater streams are important freshwater sources, but they are mostly intermittent and highly susceptible to climate change. This paper examines the sustainability of augmented freshwater availability in mountain headwater streams for water supply under baseline and future climate change scenarios using an integrated modeling approach. The climate change data in the 2040s (2030–2059), under Representative Concentration Pathway 4.5 and 8.5 scenarios, were downscaled for the impact assessment. In the region, climate change raises the average precipitation by 5–7% and the temperature by 13–15% in the 2040s. SWAT–MODFLOW model, integrating Soil and Water Assessment Tool (SWAT2012) and finite-difference Modular Groundwater Flow (MODFLOW) models in a single package, was used to assess the water balance. Results show that extracting a minimum of 16.2 m³/day from the sand storage and 30 m³/day from the aquifer was possible without affecting the groundwater table and water yield. The average annual catchment recharge was 6% of the precipitation under the baseline simulation. Climate change is projected to reduce the average water yield and groundwater recharge by 26% and 19%, respectively. However, the water supply-demand is significantly small compared to the exploitable rate of water in the area. This study was based on limited data, and therefore the findings need to be interpreted with caution, though the model output was validated using satellite products. Construction of a series of sand dams is suggested to maximize the benefit under the potential climate change and water supply-demand increase.     

气候变化对四川盆地主要粮食作物生产潜力的影响

基于四川盆地1961—2018年63个气象台站的逐日气象资料和1981—2018年46个农业气象观测站的主要粮食作物(水稻、玉米和冬小麦)生育期资料,利用逐级订正的方法计算作物气候生产潜力,分析太阳辐射、气温、降水及气候变化对四川盆地主要粮食作物气候生产潜力的影响,研究旨在为提高区域农业生产力并保障农业可持续发展提供科学依据。结果显示:1961—2018年四川盆地作物多年平均气候生产潜力的分布为水稻由西向东递增,玉米在盆地北部和西南偏高、其他地区偏低,冬小麦南北高、中部低。辐射量减小对3种作物气候生产潜力的影响为负效应;平均气温升高对作物气候生产潜力的影响为正效应;降水量变化是作物气候生产潜力变化出现空间差异的主要原因,降水量增加对作物气候生产潜力的影响为正效应,而降水量减少为负效应。气候变化对水稻气候生产潜力的影响在盆地西南部和北部的部分地区为正效应,其余地区为负效应;气候变化对玉米气候生产潜力的影响在盆地南部和东部的部分地区为正效应,其余地区为负效应;气候变化对冬小麦气候生产潜力的影响在盆地东北部的部分地区为负效应,其余大部地区为正效应。总体来看,气候变化对四川盆地冬小麦气候生产潜力的影响最大,为9.9kg·hm~(-2)·a~(-1),而对玉米和水稻的影响分别为-1.4 kg·hm~(-2)·a~(-1)和0.5 kg·hm~(-2)·a~(-1)。为了适应气候变化,四川盆地应选育光合效率高和抗旱性强的作物品种,并加强农田管理,以提高农业生产水平并保障粮食安全。     

Projected impacts of global warming on cropping calendar and water requirement of maize in a humid climate

In this research, the influence of climate change on maize cultivation was investigated and then, the possible solutions for adopting this natural hazard in the coasts of Caspian Sea in Iran was assessed. Weather data were generated for the 2011–2100 period using a statistical downscaling model under different climatic scenarios. Reference evapotranspiration (ETo) was calculated using a Neuro-fuzzy inference system. Cop water requirement was calculated by multiplying ETo by crop coefficients. Increased cardinal temperatures during 2011–2100 led to shifting in the planting date backward by 10–26 days. In addition, the projected global warming has a considerable effect on the duration of the vegetative growth stage resulting in earlier harvesting. However, the duration of the reproductive stage is less affected. Despite the obvious reduction in the length of the growing season, crop water requirement will increase by 10.6–15.3% in the future due to 1.64–28.4% increase in ETo. However, changing the cultivation time may lead to 11.2–264.5 m³ ha^?1 water saving during the whole cropping cycle through affecting both ETo and the crop growth cycle. This result demonstrates that management of the maize cropping calendar can be an effective way to achieve sustainable agriculture under future climate conditions in the study area.     

Just passing through: Global change and the conservation of biodiversity in protected areas

Climate and land-use changes are expected to cause many species to shift into or beyond the boundaries of protected areas, leading to large turnover in species composition. Here, we tested whether long-established protected areas in Canada were more robust to such climate change impacts than areas with no formal protection by measuring changes in modeled butterfly species distributions (n = 139) within them. We used a recently established distribution modeling technique, Maxent, to model butterfly species’ distributions in two epochs (1900-1930 and 1960-1990). We compared rates of butterfly species richness and composition change within protected areas against distributions of randomly selected, ecologically similar, but non-protected, areas. Change in species richness and composition within protected areas were, for the most part, the same as changes observed among random areas outside protected area boundaries. These results suggest that existing protected area networks in Canada have provided little buffer against the impacts of climate change on butterfly species richness, possibly because land-use change surrounding long-standing protected areas has not been substantial enough to elevate the habitat protection afforded by these protected areas relative to other areas. Although protected areas are unarguably beneficial in conserving biological diversity, their capacity to maintain habitat appears insufficient to prevent broader-scale climate changes from sweeping species beyond their boundaries.     

未来气候变化对四川盆地玉米生育期气候资源及生产潜力的影响

开展气候变化背景下四川盆地玉米生育期气候资源及生产潜力时空变化趋势的预估, 可对未来应对气候变化及玉米生产宏观决策提供重要的理论依据。利用区域气候模式PRECIS输出的未来A2和B2气候情景(2071-2100年)及基准气候条件(1961-1990年)气象要素资料, 分析了四川盆地玉米生育期内主要气候资源(日平均气温≥10 ℃积温、日照时数、降水量、参考作物蒸散量和缺水率)和玉米生产潜力(光合、光温和气候生产潜力)的时空变化特征。结果表明, 与基准气候条件相比, 在A2和B2两种气候情景下, 2071-2100年四川盆地玉米生育期内≥10 ℃积温、日照时数和参考作物蒸散量都呈增加趋势; 两种气候情景下, 日平均气温≥10 ℃积温的增量分别为460~641 ℃·d和376~492 ℃·d, 在盆地西部增量最大; 日照时数的增量分别为15~225 h和33~202 h, 雅安增加最多; 参考作物蒸散量的增量分别为76~144 mm和73~123 mm, 雅安增加最多。降水量在大部分地区呈减少趋势, 变幅分别为 87~56 mm和 73~47 mm, 雅安减少最多。玉米缺水率分别增加2%~18%和5%~16%, 雅安增幅最大, 未来四川盆地玉米受干旱灾害的风险可能加大。在A2和B2情景下, 2071-2100年玉米光合生产潜力分别增加228~3 277 kg·hm^-2和485~2 960 kg·hm^-2, 雅安和川北部分地区增量最大; 光温生产潜力也呈增加趋势, 分别增加2 923~5 874 kg·hm^-2和2 697~4 909 kg·hm^-2, 雅安的增量最大; 气候生产潜力同样呈增加趋势, 分别增加984~2 975 kg·hm^-2和293~2 090 kg·hm^-2, 盆地西部增加较多。未来四川盆地气候资源变化对玉米的生产有利, 产量存在提升空间。