Vessel features of Quercus ilex L. growing under Mediterranean climate have a better climatic signal than tree-ring width

We investigated whether vessel time series of Holm oak (Quercus ilex L.), a diffuse to semi-ring-porous species, can record a climatic signal which differs from the signal encoded in tree-ring width (TRW). The study was conducted in ten Q. ilex trees from a coppice stand in northeast Spain. Chronologies of TRW, mean vessel area (MVA) and maximum vessel area (MAX) were developed and correlated with climate data, for the period 1985–2004 (20 years). Our results indicate that vessel features contain environmental information that is different from that stored in TRW. MAX chronologies correlate better to early spring precipitation (April–May) than TRW chronologies, and so does MVA of the largest 20–25 vessels from the first third of the ring with late spring precipitation (May–June). Also, the combination of MVA and TRW is a better predictor of summer precipitation. This explorative study clearly shows that vessel features can complement the climatic signal of TRW increasing the resolution of the climate reconstructions for the Mediterranean region.     

Adjusting xylem anatomy and growth to inter-annual climate variability in two Fabaceae species (Centrolobium microchaete,Cenostigma pluviosum) from Bolivian dry tropical forests

Climate-growth relationships are strong in seasonally dry tropical forests (SDTFs). To better understand the ecological processes controlling these relationships we need to assess the long-term responses of wood anatomy and radial growth to year-to-year climate variability. We assessed how wood-anatomical traits (mean vessel area –MVA– and vessel density –VD–, percentage of conductive area –CA–, xylem-specific estimated hydraulic conductivity –Ks–) and growth responded to local climate (mean temperature, total precipitation, estimated moisture) variability and teleconnections (Pacific Decadal Oscillation) between 1970 and 2011 in two Fabaceae tree species (Centrolobium microchaete and Cenostigma pluviosum) coexisting in a Bolivian SDTF. We found that C. microchaete produced wider vessels and was more responsive to both local climate conditions and teleconnections than C. pluviosum. In C. microchaete VD positively responded to average temperature in the late-wet season and in the previous dry season, and CA and Ks were higher in years with warmer wet and previous early-dry seasons, as well as in years with higher PDO values. These responses were independent from ring-width variability only for C. microchaete. For C. pluviosum, vessel chronologies were more responsive to local temperature variability, and only MVA and CA positively and negatively responded to moisture in the early-dry and the previous dry seasons, respectively. Our results show that wood hydraulic structure in SDTFs is responsive to climate fluctuations. The combined study of ring width and wood anatomy allows having a more complete picture of the influence of climate on growth, particularly in species as C. microchaete which show low collinearity of tree-ring width and wood plasticity in response to climate variability.     

Influence of climate on tree-ring and earlywood vessel formation in Quercus robur in Latvia

We studied the effects of climatic factors on tree-ring width and vessel lumen area (VLA) in earlywood of English oak (Quercus robur L.) in Latvia. Cores were obtained from healthy canopy oaks in 40 stands located across Latvia. Tree-ring widths and VLA were measured. Principal component analysis was used to arrange the sites along gradients of response of tree-ring width and earlywood to environmental factors. Significant relationships of tree-ring width and mean VLA with climatic factors (mean monthly temperature and precipitation sum) were determined by correlation analysis. Relationships between tree-ring, early- and latewood widths were tested in three sampled stands. The patterns of response of VLA and tree-ring width to environmental factors differed in relation to a west–east gradient of increasing continentality. Three regions of Latvia (western, central and eastern) were distinguished along this gradient. Responses to climate differed between tree-ring width and mean VLA. Occurrence of significant correlations between climatic factors and the proxies differed between regions, likely due to regional differences in temperature and precipitation. Tree-ring width correlated with climatic factors (most commonly with March, May and June temperature and August precipitation of the current growing season and July–August temperatures of the previous growing season); VLA was more strongly related to climatic factors, particularly with temperature in winter and spring months. The proportion of significant correlation coefficients with climatic factors differed between the regions. Among sites, significant correlation of tree-ring width with temperature in spring and summer was more frequent in the western region, while correlation with winter temperature of the previous growing season and precipitation in August was more frequent in the eastern region. For VLA, the frequency of significant correlation coefficients with temperature in winter and spring was higher in the eastern region.     

“Plastic” and “static” behavior of vessel-anatomical features in Oriental beech (Fagus orientalis Lipsky) in view of xylem hydraulic conductivity

Key message

Xylem anatomical traits can be categorized into two groups: plastic properties which show a high inter-annual variability, and static characteristics which vary in a more conservative range.

Abstract

Water conduction in broad-leaved trees depends mainly on the size, number, and arrangement of vessels, which vary from year to year in response to varying exogenous factors, thus contributing to a safe and/or efficient water transport. However, the nature of such compensation is not clear; in particular, it is not obvious which traits act independently and which ones coincidentally. To better understand these inter-relations, tree-ring width (TRW), vessel-related anatomical traits, and the theoretical hydraulic conductivity were measured or modeled in the last 50 growth rings of mature Oriental beech trees growing at different altitudes in northern Iran. The study trees followed similar strategies compensating the effects of external factors by modifying their vessel-anatomical features. TRW and the number of vessels per unit of area were highly but negatively correlated and both were affected by exogenous factors. However, a decrease in vessel frequency (VF) is not a mirror effect of wider tree rings, but trees actively control the number of vessels produced. Principal component analysis revealed that the features VF, TRW and relative total conductivity were more plastic, whereas average vessel-lumen area, tree-ring porosity, and relative specific conductivity behaved more static. Moreover, we suggest that in theoretical approaches, total hydraulic conductivity rather than the specific hydraulic conductivity is a better indicator of a tree’s hydraulic behavior in a given growing season.     

dendroTools: R package for studying linear and nonlinear responses between tree-rings and daily environmental data

We introduce in this paper the dendroTools R package for studying the statistical relationships between tree-ring parameters and daily environmental data. The core function of the package is daily_response(), which works by sliding a moving window through daily environmental data and calculating statistical metrics with one or more tree ring proxies. Possible metrics are correlation coefficient, coefficient of determination and adjusted coefficient of determination. In addition to linear regression, it is possible to use a nonlinear artificial neural network with the Bayesian regularization training algorithm (brnn). dendroTools provides the opportunity to use daily climate data and robust nonlinear functions for the analysis of climate-growth relationships. Models should thus be better adapted to the real (continuous) growth of trees and should gain in predictive capabilities. The dendroTools R package is freely available in the CRAN repository. The functionality of the package is demonstrated on two examples, one using a mean vessel area (MVA) chronology and one a traditional tree-ring width (TRW).     

Impact of extreme drought on tree-ring width and vessel anatomical features of Chukrasia tabularis

Multiple sources of evidence suggest an increasing frequency of extreme climatic events during the past century. In Bangladesh, a country strongly influenced by the South Asian monsoon climate, the years 1999 and 2006 were the most severe droughts among the ten drought events identified over the last four decades. We investigated the impact of these two drought events on radial growth and xylem anatomical features of the brevi-deciduous tree species Chukrasia tabularis in a moist tropical forest in Bangladesh. Tree radial growth declined by 54% during the 1999 and 48.7% during the 2006 droughts, respectively. Among the wood anatomical features, the number of vessels (NV) showed the highest sensitivity to drought, with a 45% decrease in the 1999 drought year, followed by total vessel area (TVA) and mean vessel area (MVA). On the other hand, Vessel density (VD) increased by 13% during the 1999 drought but the increase in VD was very low in the drought year 2006. The decreasing vessel area and increasing vessel density indicate xylem hydraulic adaptation of C. tabularis to minimize drought induced cavitation risk and to avoid hydraulic failure. The significant correlations between the Standardized Precipitation Evapotranspiration Index (SPEI) and time series of tree-ring width and vessel variables imply that decline in radial growth and changes in vessel features in C. tabularis are likely to be caused by drought induced water stress. Our analyses suggest that radial growth and wood anatomical features of C. tabularis are highly sensitive to extreme drought events in South Asian moist tropical forests and can be used to reconstruct past droughts and to model tree response to drought stress under future climate conditions.     

Ring width and vessel features of the mangrove Excoecaria agallocha L. depend on salinity in the Sundarbans,Bangladesh

The Bangladesh Sundarbans is the largest continuous mangrove in the world that providing crucial environmental services, particularly related to coastal protection and livelihoods of millions of people. However, anthropogenic disturbances, diseases infestation and environmental changes including sea level rise (SLR) and fresh-water flux into the delta are threatening the Sundarbans and other mangrove ecosystems worldwide. Protection of mangrove ecosystems requires knowledge on factors that mainly drive growth and vitality of tree species to evaluate which consequences can be expected from, mainly hydrology-related, environmental changes. In this study, we assessed the nature and periodicity of tree rings in Excoecaria agallocha, a wide spread mangrove species in the Bangladesh Sundarbans. We also analysed the influence of climatic factors, such as precipitation, temperature and vapor pressure deficit (VPD), and river discharge, as a proxy of salinity on ring width (RW) and vessel features, such as mean vessel area (MVA) and mean vessel density (MVD). E. agallocha forms distinct tree-ring boundary that characterized by a narrow (2–4 cells wide) band of radially flattened fibres. The RW as well as the MVA and MVD are crossdatable. The RW is mainly driven by salinity which is influenced by freshwater inputs through precipitation during monsoon along with river discharge January to April. The MVA and MVD responded to similar seasons and months as RW, but mostly with opposite signs in MVD. The results suggest that fresh water inputs through precipitation and river discharge positively influence the radial growth of E. agallocha in the Sundarbans. The RW and vessel features can be used as proxies to explore the growth dynamics of this species, especially in relation to global environmental changes.     

Separating the climatic signal from tree-ring width and maximum latewood density records

We propose a technique for separating the climatic signal which is contained in two tree-ring parameters widely used in dendroclimatology. The method is based on the removal of the relationship between tree-ring width (TRW) and maximum latewood density (MXD) observed for narrow tree rings from high latitudes. The new technique is tested on data from three larch stands located along the northern timberline in Eurasia. Correlations were calculated between the temperatures of pentads (five consecutive days), TRW chronologies and MXD chronologies calculated according to the standard and proposed methods. The analysis confirms the great importance of summer temperature for tree radial growth and tree-ring formation. TRW is positively correlated with the temperature of four to eight pentads (depending on the region) at the beginning of the growth season, but MXD as obtained by the standard technique is correlated with temperature over a much longer period. For maximum density series from which the relationship between MXD and TRW has been removed (MXD′), there is a clear correlation with temperatures in the second part of the growing season. These results are consistent with the known dynamics of tree-ring growth in high latitudes and mechanisms of tree-ring formation.     

High-resolution time series of vessel density in Kenyan mangrove trees reveal a link with climate

Tropical trees are often excluded from dendrochronological investigations because of a lack of distinct growth ring boundaries, causing a gap in paleoclimate reconstructions from tropical regions. The potential use of time series of vessel features (density, diameter, surface area and hydraulic conductivity) combined with spectral analysis as a proxy for environmental conditions in the mangrove Rhizophora mucronata was investigated. Intra-annual differences in the vessel features revealed a trade-off between hydraulic efficiency (large vessels) during the rainy season and hydraulic safety (small, more numerous vessels) during the dry season. In addition to the earlywood-latewood variations, a semiannual signal was discovered in the vessel density and diameters after Fourier transformation. The similarity in the Fourier spectra of the vessel features and the climate data, in particular mean relative humidity and precipitation, provides strong evidence for a climatic driving force for the intra-annual variability of the vessel features. The high-resolution approach used in this study, in combination with spectral analysis, may have great potential for the study of climate variability in tropical regions.     

Growing-season precipitation since 1872 in the coastal area of subtropical southeast China reconstructed from tree rings and its relationship with the East Asian summer monsoon system

Tree rings from temperate zones of the world have provided abundant palaeo- ecological and paleo-hydroclimatic information. However, tree rings from subtropical to tropical regions remain relatively scarce, which greatly limit our fully understanding about the climate change issues. In the present work, tree-ring-width (TRW) measurements of Masson pine from Fujian province, the coastal area of subtropical southeast China were successfully crossdated and a TRW STD chronology was developed from 1854 to 2012. Significantly positive correlation was identified between the tree rings and April–November total precipitation (r = 0.71, p < 0.01). The reconstructed April–November precipitation exhibited two comparatively wet (1876–1886 and 1957–1962) and one comparatively dry (1986–2004) periods. An evident drying trend since 1959 was seen and it was mitigated after 1993. Most of the extreme low-precipitation years in the reconstruction were supported by the historical records. As revealed by the spatial correlation patterns, our precipitation reconstruction was also consistent with other hydroclimatic records along the coastal areas of southeast China, proving its ability to capture the large-scale hydrological signal in southeast China (mainly refers to the south of the middle-lower reaches of Yangtze River). The reconstructed precipitation showed significant correlation with the East Asian summer Monsoon (EASM) index. Moreover, it also indicated simultaneous variation with the monsoon precipitation in North China on a decadal scale, implying that growing season precipitation variations in both regions were influenced by the EASM strength. This work highlights the potential of using tree-ring width to reconstruct precipitation in subtropical southeast China, while the relevant issues about precipitation variation in this region is far from resolved.     

High-frequency variation of tree-ring width of some native and alien tree species in Latvia during the period 1965–2009

The plasticity of climate-growth relationships of trees is one of the main factors determining the climate-induced changes in forest productivity and composition. In this study, high-frequency variation of tree-ring width (TRW) of four native and three alien tree species and two hybrids of Populus L. growing in Latvia (hemiboreal zone) was compared using a principal component analysis based on TRW indices for the period 1965–2009. The effect of climatic factors was assessed using a bootstrapped correlation analysis. Influence of common climatic factors related to the length of the vegetation season, winter temperature, and water regime in summer was traced in the TRW of the studied species and hybrids. The combination and effect of the identified factors differed by species (and hybrids), to a certain extent explaining the diversity of TRW patterns. Nevertheless, some similarities among the species were also observed, suggesting the plasticity of growth response. Scots pine was generally sensitive to winter temperatures, but Norway spruce was mainly sensitive to summer water regime, while black alder was sensitive to winter temperatures and precipitation in spring. In contrast, silver birch showed the lowest sensitivity to the tested climatic factors (demonstrating sensitivity to winter precipitation in a few sites), suggesting tolerance to weather fluctuations. The TRW of the alien species was primarily sensitive to climatic factors related to water regime in the summer of the year preceding the formation of tree-ring, implying differences in mechanisms regulating wood increment. Nevertheless, temperature in the dormant period was significant for European larch in a few sites, suggesting sensitivity to cold damage. The variation of TRW of Populus hybrids diverged from others, as their growth was negatively correlated with the temperature in autumn, spring, and summer and positively correlated with water balance. Although the annual water balance in Latvia is positive, the effect of water deficit on tree growth was apparent.     

Tree ring width and wood density as the indicators of climatic factors and insect outbreaks affecting spruce growth

Tree ring width (TRW), maximum (MXD), mean (MED) and minimum (MID) wood density were investigated in samples from the vicinity of the Tuchola Forest Biosphere Reserve (Northern Poland) in an attempt to distinguish the relative importance of climate and insect attack on the growth of Norway spruce. Selected climate parameters were used for a multiple regression to predict tree-ring width during insect outbreaks. This also used AICc for model selection. Additionally, k-means clustering was then used to group the yearly data of TRW, MXD, MID and the data of insect outbreaks. The respective climate data and data on insect outbreaks during the years 1962–1996 revealed a strong influence of May precipitation on TRW and insect outbreaks on MID. Missing tree rings or narrow rings and lower MXD together with higher MID might indicate increased insect activity.     

Climatic impact on tree-ring widths in Abies borisii-regis forests from South-East Albania

The long-term radial growth responses to drought and climatic variability of less-studied species such as Abies borisii-regis (Mattf.) remain poorly understood.We tested the hypothesis that severe short-term drought conditions during summer months will impact the radial growth of A. borisii-regis (Mattf.) trees and such impact will have a more pronounced effect on latewood (LW) than earlywood (EW) width.Correlation analysis was employed to investigate the impact of climatic drivers (temperature, precipitation) and drought, using the Standardized Precipitation Evapotranspiration Index (SPEI) calculated at cumulative time scales (1–12 months), on EW, LW and tree-ring width (TRW) in A. borisii-regis (Mattf.) trees from South-Eastern Albania.We found that EW width was positively correlated with precipitation in July and previous September, while the LW width and TRW was enhanced by the current June–July precipitation. Previous autumn and current summer high temperatures constrained the radial growth in A. borisii-regis (Mattf.) trees, particularly the LW and TRW. All the tree-ring widths components showed the highest significant response to drought at short cumulative time scales (<4 months) mainly during July, August and September. The highest impact of drought was observed for the LW width.Under a future reduction of summer precipitation and temperature increase, the A. borisii-regis (Mattf.) may show a decrease in EW formation, causing a decline of radial growth, leading to a reduction in hydraulic conductivity and carbon uptake in these forests.     

An unstable tree-growth response to climate in two 500 year chronologies,North Eastern Qinghai-Tibetan Plateau

Two new Juniper tree-ring-width (TRW) chronologies spanning more than 500 years were developed in the Yellow River source area, North Eastern Qinghai-Tibetan Plateau (NE-QTP). For the two studied sites, located approximately 50 km apart, split correlation and coherence analysis reveal unstable tree-growth responses to local moisture availability. While significant correlations are obtained with April–June local precipitation, Palmer Drought Severity Index (PDSI) and river flow from 1948/1954 to 1998 and from 1948/1954 to 1970s, these correlations vanish for the time period 1970s-1998. The local instrumental climate data (precipitation, PDSI and river flow) exhibit opposite correlations with large scale modes of variability (El Niño Southern Oscillation, ENSO, and Pacific Decadal Oscillation, PDO) before and after the 1977 PDO shift. One tree-ring chronology is coherent and anti-phased with instrumental ENSO/PDO indices at 5.2-year frequency. On the longer time span, this TRW chronology is compared with PDO reconstructed from historical Chinese data. This comparison also exhibits unstable multi-decadal relationships, notably in the mid 19th century. Altogether, the comparison between our two chronologies, local instrumental climate records, and ENSO/PDO indices suggest a cautious use of local TRW records for paleoclimate reconstructions. Further studies are needed to explore both the spatial coherency of tree-ring records and the temporal stability of their response to local and large scale climate variability.     

多枝柽柳年轮生长速率测量及TRGR指标优势分析

灌木对生境和气候变化具有高度敏感性,其年轮资料在认识区域环境演变过程、全球气候变化和环境保护中具有重要作用。灌木植株在生长过程中受遗传和极端环境的影响,年轮常出现偏心和不规则生长,这使得专业年轮分析软件测量的年轮宽度数据难以准确反映其整体径向生长信息。为探讨适合于寒旱区灌木年轮学研究的年轮测量指标和测量方法,研究以该区域荒漠常见植物多枝柽柳(Tamarix ramosissima Ledeb.)为研究对象,通过U-net深度学习方法,获得年轮提取训练模型,自动获取扫描轮盘各年早材区域。轮盘扫描及语义分割后的图像经GIS配准赋坐标、ENVI图像处理后,借助GIS编辑和测量工具,完成多枝柽柳各年年轮生长速率(Tree-ring growth rate, TRGR)、年轮宽度(Tree-ring width, TRW)和树木基部断面积生长增量(Basal area increment, BAI)的测量;研究基于Timesat Savitzky-Golay(S-G)滤波时间序列拟合,获取点样尺度归一化植被指数(Normalized difference vegetation index, ND...     

The influence of an altered groundwater regime on vessel properties of hybrid poplar

The development of vascular features in stems of Populus x euramericana was studied in two adjacent floodplain forest stands near the Danube in Austria. The construction of a hydropower plant caused one of the stands to loose contact with the groundwater at the age of 12 years, whereas the other stand remained uninfluenced. The investigation estimated the extent of vascular acclimation to altered site hydrology. Vessel lumen areas were separately measured for 26 tree-rings from six trees. Vessel densities (VD) and the percentages of the total vessel-lumen area per tree-ring area (PLA) were determined. Our results suggest that vessel size and density are correlated with circumferential stem growth, which is in turn governed by site water supply. In relation to the control trees, basal area increment (BAI) of the hydrologically altered trees was 66% lower. BAI was positively correlated to vessel size and negatively to VD and PLA. Vessel diameters of the mature control trees were large (with a mean of 95 m and maxima around 160 m), while average VD (42 no. mm^-2) and PLA (31%) were within the normal ranges. In comparison to controls, the hydrologically altered trees reduced their vessel-lumen area by 15% (i.e. 8% in terms of diameter); average VD increased by 43%, and average PLA increased relatively by 19%. Vessel density proved to be the most sensitive indicator of hydrological alterations. Intra-annual development of vessel size indicated a rapid response to the hydrological conditions of the current growing season.     

A lonely dot on the map: Exploring the climate signal in tree-ring density and stable isotopes of clanwilliam cedar,South Africa

Clanwilliam cedar (Widdringtonia cedarbergensis; WICE), a long-lived conifer with distinct tree rings in Cape Province, South Africa, has potential to provide a unique high-resolution climate proxy for southern Africa. However, the climate signal in WICE tree-ring width (TRW) is weak and the dendroclimatic potential of other WICE tree-ring parameters therefore needs to be explored. Here, we investigate the climatic signal in various tree-ring parameters, including TRW, Minimum Density (MND), Maximum Latewood Density (MXD), Maximum Latewood Blue Intensity (MXBI), and stable carbon and oxygen isotopes (δ¹⁸O and δ¹³C) measured in WICE samples collected in 1978. MND was negatively influenced by early spring (October-November) precipitation whereas TRW was positively influenced by spring November-December precipitation. MXD was negatively influenced by autumn (April-May) temperature whereas MXBI was not influenced by temperature. Both MXD and MXBI were negatively influenced by January-March and January-May precipitation respectively. We did not find a significant climate signal in either of the stable isotope time series, which were measured on a limited number of samples. WICE can live to be at least 356 years old and the current TRW chronology extends back to 1564 CE. The development of full-length chronologies of alternative tree-ring parameters, particularly MND, would allow for an annually resolved, multi-century spring precipitation reconstruction for this region in southern Africa, where vulnerability to future climate change is high.     

Winter North Atlantic oscillation effects on the tree rings of the Italian beech (Fagus sylvatica L.)

Climatic signals in beech tree-ring width series from Central Italy have been studied over different periods of time. Prewhitened tree-ring chronologies respond mainly to summer precipitation and they do not correlate in a significant manner with the winter North Atlantic oscillation (NAO) index. In this high-frequency pattern the NAO signs are only found on a small number of rings characterized by being very narrow or wide. By contrast, tree-ring width chronologies in which all the frequency components are conserved were significantly related to the NAO. The significant inverse correlation between actual measurements of ring width and NAO is a consequence of the availability of water in the soil at the beginning of the growing season. In fact, in the Mediterranean area the recharging of soil moisture depends on the amount of winter precipitation, which is inversely correlated with the NAO. Strong signals of winter precipitation and NAO are found in the low-frequency components of tree-ring growth. Received: 18 March 1999 / Revised: 29 February 2000 / Accepted: 1 March 2000     

Moisture‐driven shift in the climate sensitivity of white spruce xylem anatomical traits is coupled to large‐scale oscillation patterns across northern treeline in northwest North America

Tree growth at northern treelines is generally temperature‐limited due to cold and short growing seasons. However, temperature‐induced drought stress was repeatedly reported for certain regions of the boreal forest in northwestern North America, provoked by a significant increase in temperature and possibly reinforced by a regime shift of the pacific decadal oscillation (PDO). The aim of this study is to better understand physiological growth reactions of white spruce, a dominant species of the North American boreal forest, to PDO regime shifts using quantitative wood anatomy and traditional tree‐ring width (TRW) analysis. We investigated white spruce growth at latitudinal treeline across a >1,000 km gradient in northwestern North America. Functionally important xylem anatomical traits (lumen area, cell‐wall thickness, cell number) and TRW were correlated with the drought‐sensitive standardized precipitation–evapotranspiration index of the growing season. Correlations were computed separately for complete phases of the PDO in the 20th century, representing alternating warm/dry (1925–1946), cool/wet (1947–1976) and again warm/dry (1977–1998) climate regimes. Xylem anatomical traits revealed water‐limiting conditions in both warm/dry PDO regimes, while no or spatially contrasting associations were found for the cool/wet regime, indicating a moisture‐driven shift in growth‐limiting factors between PDO periods. TRW reflected only the last shift of 1976/1977, suggesting different climate thresholds and a higher sensitivity to moisture availability of xylem anatomical traits compared to TRW. This high sensitivity of xylem anatomical traits permits to identify first signs of moisture‐driven growth in treeline white spruce at an early stage, suggesting quantitative wood anatomy being a powerful tool to study climate change effects in the northwestern North American treeline ecotone. Projected temperature increase might challenge growth performance of white spruce as a key component of the North American boreal forest biome in the future, when drier conditions are likely to occur with higher frequency and intensity.