3种林木凋落物分解特征及其对赤红壤酸度及养分含量的影响

在实验室条件下对黧蒴锥[Castanopsis fissa(Champ.ex Benth.)Rehd.et Wils.]、湿地松(Pinus elliottii Engelm.)和尾叶桉(Eucalyptus urophylla S.T.Blakely)凋落物的分解特征及其对华南赤红壤的pH值、有机质和碱解氮、速效磷和速效钾含量的影响进行了研究.结果表明:经过1 a的分解后,黧蒴锥、湿地松和尾叶桉凋落物的失重率分别为26.48％、13.80％和28.15％;3种凋落物中全氮均为净释放模式,全磷为富集模式,黧蒴锥和尾叶桉凋落物中全钾和有机碳为净释放模式,湿地松凋落物中全钾和有机碳为富集模式.随凋落物分解时间的延长,土壤pH值呈现逐渐上升至第8个月达到峰值、之后略有下降的变化趋势;有机质含量呈波动的变化趋势;碱解氮含量呈先升高后降低的变化趋势;速效磷含量呈略有波动但总体下降的趋势;速效钾含量呈“逐渐上升—急剧下降—急剧上升—趋于平稳”的变化趋势,且均在第8个月最低.与实验初期相比,实验结束时对照组及各处理组的土壤pH值和碱解氮含量均不同程度增加、有机质含量不同程度降低、速效磷含量显著下降,对照组速效钾含量下降而各处理组速效钾含量明显提高.与对照相比,实验结束时3种凋落物均导致土壤酸度减弱、速效钾含量增加、但有机质和速效磷含量没有明显提高.3种凋落物对土壤pH值及土壤养分含量的影响有明显差异,总体上看,黧蒴锥凋落物覆盖可明显减弱土壤酸度,增加土壤有机质、碱解氮和速效钾含量,对土壤的改良效果最好.研究结果揭示:不同凋落物对土壤pH值及养分含量的影响与凋落物的成分及生物学特性有关.     

鄂东南低丘地区枫香林下枫香幼苗更新限制因子

深入理解幼苗更新的生态过程及其限制因子是植物种群保育和森林管理的基础。在鄂东南的低丘地区, 采用控制试验研究了限制枫香(Liquidambar formosana)幼苗在其自身林下更新的生态因子。由于枫香幼苗在马尾松(Pinus massoniana)林下偶尔更新良好, 在枫香林下几乎没有更新, 以马尾松林为对照, 设计了4个试验来检验: (1)低温和土壤湿度对枫香种子储藏过程中寿命的影响, (2)两种林型的土壤、凋落物对枫香种子越冬及存活的影响, (3)凋落物对枫香种子萌发是否具有化感作用, (4)林下的光强变化是否影响幼苗的存活。结果表明, 枫香种子在很低的温度下仍可以立即萌发, 这表明枫香种子没有休眠特性是影响枫香幼苗更新的重要因子; 土壤、凋落物和凋落物滤液对种子越冬后的存活没有影响, 不能解释“枫香林下没有枫香苗”的问题; 两种林型的主要凋落物滤液对枫香种子萌发没有抑制作用或促进作用; 一年生枫香幼苗遮阴试验的结果表明, 3%透光率下幼苗的死亡率接近90%。因此枫香林下弱的光环境和种子的低温萌发特性是影响枫香幼苗更新的重要因子。     

啮齿动物及凋落物存留对天山野果林野生樱桃李种子命运的影响

2010年9月至2013年4月通过设置人工土壤种子库,研究了野生樱桃李土壤种子库的动态及啮齿动物和凋落物覆盖对种子库中种子命运的影响.结果表明: 在有取食动物扰动下,48.3%的种子萌发输出为幼苗,50%的种子被动物搬运或当场取食,4%的种子腐烂.在排除了取食动物干扰的条件下,樱桃李种子形成了短期持久的土壤种子库,3年后依然有部分种子萌发并输出为幼苗.凋落物覆盖和对照处理中,被搬运和当场取食种子的比例均显著低于清除凋落物的裸地.地表凋落物存留能减少动物搬运、取食,但不足以导致新生幼苗的大量出现,而啮齿类动物的搬运或取食是影响野生樱桃李种子命运和种子库动态的主要因素.
     

三峡地区柏木种子萌发和幼苗更新的研究

对三峡地区柏木(Cupressus funebris)林柏木种子的萌发、幼苗存活、幼苗生长进行了研究.结果表明,(1)柏木种子的萌动期约为10 d,播种35-45 d后进入萌发高峰期,柏木种子的平均萌发率为47.7%.土壤基质对种子萌发的影响是极显著的,光照对萌发则没有显著的影响.(2)幼苗平均死亡率为52.6%.土壤类型和光照强度对幼苗的存活都有显著影响,在林下土和80%光照条件下幼苗的存活率最高.(3)柏木幼苗生长初期茎的生长动态呈"S"型,而叶片数量的增长动态近于直线型.幼苗生长初期生物量形成主要受光照条件的影响,土壤类型的影响不显著.土壤和光照条件对柏木林柏木幼苗的更新都有显著影响,但是土壤条件主要影响种子萌发和幼苗的存活,而光照条件则主要对幼苗的生长和存活产生影响.     

格氏栲幼苗叶绿素荧光和非结构性碳水化合物积累对种子散布位置的响应

植物种子从母树掉落形成土壤种子库时,凋落物或土壤是其最初接触的物理环境,种子所处位置(种子在凋落物上层、土壤表层或凋落物下层)影响了幼苗天然更新进程。模拟格氏栲种子在凋落物上层(种子下层铺垫2和4 cm凋落物)、土壤表层(无凋落物)及凋落物下层(种子上层覆盖2、4、6和8 cm凋落物)等3种不同散布位置,探讨种子散布位置对幼苗叶绿素荧光特性、非结构性碳水化合物、比叶面积、叶干物质含量和养分含量的影响。结果表明:不同散布位置的幼苗单位面积的叶氮含量与可溶性糖、非结构性碳水化合物含量呈显著正相关,与比叶面积呈显著负相关。适宜凋落物覆盖(2和4 cm)的幼苗通过提高叶绿素相对含量、可溶性糖含量、非结构性碳水化合物含量、叶干物质含量和单位面积的叶氮含量和叶磷含量,降低比叶面积等的资源获取策略来实现自身快速生长需求。无凋落物和深层凋落物覆盖(6和8 cm)的幼苗采取高单位重量的叶氮含量和比叶面积,低叶干物质含量和非结构性碳水化合物含量的资源保守型策略以截获更多有效光资源,进而弥补深层凋落物带来的郁闭环境,降低幼苗因“碳饥饿”而死亡的几率。下层铺垫凋落物的幼苗通过在叶片储藏淀粉,降低叶片光合组织消耗能量(低PSⅡ最大光化学效率)等维持幼苗生长。熵值法综合分析表明,浅层凋落物覆盖(2 cm)对格氏栲幼苗生长的促进作用最为显著,未来可通过调节天然林凋落物层厚度以促进格氏栲幼苗生长与更新。     

凋落物物理阻隔对格氏栲种子萌发及胚根生长的影响

为了探讨凋落物物理阻隔对格氏栲天然林自然更新状况的影响,通过模拟野外凋落物覆盖,设置格氏栲种子上层覆盖厚0 cm(CK)、2 cm(D2)、4 cm(D4)、6 cm(D6)、8 cm(D8)及种子下层铺垫厚2 cm(U2)、4 cm(U4)凋落物等7个处理,分析凋落物覆盖方式及厚度对格氏栲种子萌发及胚根生长的影响。结果表明:(1)凋落物覆盖方式及厚度对种子萌发进程存在显著影响。CK萌发持续时间最长,上层覆盖处理(D)次之,下层铺垫处理(U)的种子起始萌发时间显著滞后。(2)CK种子萌发率最高,其次D6处理发芽速度较快,发芽整齐;U处理较D处理的发芽率及发芽势均显著降低,且萌发抑制率显著增加。(3)D处理的种子胚根生长速度快,胚根长度大于CK;U处理的种子胚根生长速度呈先慢后快趋势。可见,凋落物是影响格氏栲种子萌发及胚根生长的重要因素,主要通过阻碍种子与土壤接触而抑制萌发,影响格氏栲林更新。     

水分条件对干热河谷锥连栎种子萌发及幼苗早期生长的影响

对锥连栎(Quercus franchetii Skan)在4种施水量下的种子萌发、幼苗早期生长形态及生物量分配的响应特征进行了研究。结果表明,施水量对锥连栎的种子萌发过程具有显著影响。随着施水量增加,其种子萌发率、萌发指数以及萌发时间和萌发持续时间均呈明显的上升趋势,而萌发时滞和萌发速率系数呈下降趋势。锥连栎幼苗早期生长对施水量的适应幅度较大,每次施水4～12 mm时,幼苗均能存活,但生长状况存在明显差异,随着施水量的减少,幼苗生长和生物量的积累速度均出现显著减缓。同时,施水量对锥连栎幼苗生物量分配具有显著影响,随着施水量增加,幼苗根冠比呈显著下降趋势,表明在生物量分配上锥连栎幼苗具有极大的可塑性,施水量减少时,光合产物将向地下部分迁移,增加根冠比。     

森林凋落物对种子萌发与幼苗生长的影响

森林更新是森林生态系统维持自身稳定的重要环节,其早期阶段(种子萌发与幼苗生长)对气候变化十分敏感,同时显著地受到森林凋落物的影响。对这种影响机制的深入研究,有助于进一步认识和了解森林更新动态和群落演替方向。本文简要概述了近年来国内外森林凋落物研究动态,从森林凋落物影响种子萌发与幼苗生长的物理作用机制(凋落物对种子与幼苗产生的物理障碍与机械损伤,微环境的温度、湿度以及光照条件的变化等)、生物化学作用机制(产生化感物质、降解释放营养物质)以及通过改变土壤理化性质间接对其产生影响等3个方面进行了系统综述,并且就气候变化与森林凋落物的协同作用通过多种方式直接或间接地对树木种子萌发与幼苗生长产生影响的机理进行了初步探讨。在此基础上提出了这一领域未来研究趋势和需要加以重视的问题。     

凋落物和土壤覆盖对动物取食和搬运辽东栎种子的影响

在六盘山区的华北落叶松人工林,研究了清除凋落物、凋落物覆盖和土壤覆盖(以不清除凋落物直接将种子投放于森林地表为对照)等处理对动物取食和搬运辽东栎种子的影响.结果表明:种子释放3d后,凋落物和土壤覆盖处理种子均具有较高的留存率(分别为10.7％和7.0％);释放14 d后,土壤覆盖处理种子的留存率仍最高(0.7％),但凋落物覆盖处理种子的留存率为0.凋落物和土壤覆盖处理种子的就地取食率很高(分别为45.9％和41.5％);清除凋落物处理种子的就地取食率最低(27.0％),但其被搬运后的取食率最高(49.8％);凋落物覆盖、清除凋落物和土壤覆盖处理种子被动物搬运后的埋藏率均显著高于对照(P＜0.01).种子被动物搬运后集中分布于5 m以内,尤其在＜1 m和l～2m两个距离组的分布频率更高;种子被搬运后取食的平均距离大干埋藏的平均距离,以土壤覆盖和凋落物覆盖处理最大,分别为2.38 m±0.55 m和1.44m±0.26 m.     

种子散布位置对格氏栲幼苗根系形态及生长特征的影响

种子从母株掉落于地面萌发后,其根系在不同散布位置(凋落物上层、土壤表层和凋落物下层)的生长形态影响幼苗定居及建成,而目前对其根系形态及生长特征的了解并不充分,限制了对幼苗根系在不同散布下适应策略的理解。为此,以格氏栲(Castanopsis kawakamii)为研究对象,通过模拟种子在凋落物中位置,设置凋落物上层(种子下层铺垫2 cm和4 cm凋落物,U2和U4处理)、土壤表层与凋落物下层(种子上层覆盖0、2、4、6 cm和8 cm凋落物,CK、D2、D4、D6和D8处理)等3种散布,探讨不同散布位置对格氏栲幼苗根系9个生长指标的影响。结果表明：(1)种子散布位置对幼苗根干物质质量具有显著影响,D2处理达最大值。(2)D2处理的幼苗根长、根表面积、根尖数、分枝数和比根长高于其它处理;根系平均直径在D6处理达最大值。(3)相关分析表明根长、根表面积、根尖数、分枝数和比根长与根系平均直径呈显著负相关关系。(4)对根系9个生长指标提取主成分后聚类为4个类群,D2与D4处理各划分一类;U2与U4处理划分一类,其余三个处理划分一类。综上所述,凋落物浅层覆盖(D2处理)适宜格氏栲根系生长;凋落物...     

Decoupling litter barrier and soil moisture influences on the establishment of an invasive grass

Background and aims

Through recruitment, plants establish in novel environments. Recruitment also is the stage where plants undergo the highest mortality. We investigate the recruitment niche for Microstegium vimineum, an annual grass from East Asia spreading throughout eastern North American forests.

Methods

Current observational and greenhouse research indicates that M. vimineum recruitment may be inhibited by leaf litter and promoted by soil moisture; we use field studies to experimentally test how these factors influence M. vimineum germination, seedling survival and reproduction. Specifically, we introduce M. vimineum seeds into forest microhabitats with experimentally varied levels of soil moisture and leaf litter.

Results

Soil moisture increases M. vimineum germination regardless of leaf litter thickness and ameliorates seedling mortality in deep leaf litter. Seed production per m² increases with watering, reflecting higher germination and survival, whereas per capita seed production increases with leaf litter thickness, reflecting density-dependent limits on seed production.

Conclusions

The interactive effects of varied levels of soil moisture and leaf litter thickness on key M. vimineum life history stages highlight the need to consider multiple drivers, such as rainfall and local forest disturbance, when assessing how soil properties influence the establishment of invasive plants.     

冻融过程对荒漠短命植物种子萌发的影响

荒漠区冻融交替显著改变土壤温度和水分条件,并进一步影响荒漠植物种子萌发。为解析荒漠土壤冻融过程对植物种子萌发的影响,本研究以古尔班通古特沙漠4种典型短命植物[东方旱麦草(Eremopyrum orientale)、卵果鹤虱(Lappula patula)、尖喙牻牛儿苗(Erodium oxyrrhynchum)和条叶庭荠(Alyssum linifollum)]为对象,通过模拟不同温度和土壤水分及冻融循环次数对种子进行处理,对比分析冻融结束后4种植物种子萌发参数(发芽率、发芽势、发芽指数、种子活力)。结果表明,温度对4种植物种子萌发参数均无显著影响,而土壤水分和循环次数均有显著影响(水分仅对条叶庭荠无影响),且存在显著交互作用。冻融循环次数增加可促进冻融后的种子活力和种子萌发,尤其是在低水分条件下表现得更为显著。冻融期高土壤水分显著降低东方旱麦草的种子发芽率,对其他物种的影响受冻融次数和白天温度状况的调节。在相同土壤水分条件下,冻融期白天高温可促进种子萌发。总体来看,冻融对荒漠短命植物种子萌发有显著影响并存在一定的种间差异性,这将造成冻融期结束后不同物种的种子萌发和幼苗生长存在差异,进而影响荒漠区植物群落结构。     

Fire season and intensity affect shrub recruitment in temperate sclerophyllous woodlands

The season in which a fire occurs may regulate plant seedling recruitment because of: (1) the interaction of season and intensity of fire and the temperature requirements for seed release, germination and growth; (2) post-fire rainfall and temperature patterns affecting germination; (3) the interaction of post-fire germination conditions and competition from surrounding vegetation; and (4) the interaction of post-fire germination conditions and seed predators and/or seedling herbivores. This study examined the effects of different fire intensities and fire seasons on the emergence and survival of shrubs representing a range of fire response syndromes from a summer rainfall cool climate region. Replicated experimental burns were conducted in two seasons (spring and autumn) in 2 consecutive years and fuel loads were increased to examine the effects of fire intensity (low intensity and moderate intensity). Post-fire watering treatments partitioned the effects of seasonal temperature from soil moisture. Higher intensity fires resulted in enhanced seedling emergence for hard-seeded species but rarely influenced survival. Spring fires enhanced seedling emergence across all functional groups. Reduced autumn recruitment was related to seasonal temperature inhibiting germination rather than a lack of soil moisture or competition. In Mediterranean-type climate regions, seedling emergence has been related to post-fire rainfall and exposure of seeds to seed predators. We think a similar model may operate in temperate summer rainfall regions where cold-induced dormancy over winter exposes seeds to predators for a longer time and subsequently results in recruitment failure. Our results support the theory that the effect of fire season is more predictable where there are strong seasonal patterns in climate. In this study seasonal temperature rather than rainfall appears to be more influential.     

The Influence of Variable Rainfall Frequency on Germination and Early Growth of Shade-Tolerant Dipterocarp Seedlings in Borneo

Climate change induced alterations to rainfall patterns have the potential to affect the regeneration dynamics of plant species, especially in historically everwet tropical rainforest. Differential species response to infrequent rainfall may influence seed germination and seedling establishment in turn affecting species distributions. We tested the role of watering frequency intervals (from daily to six-day watering) on the germination and the early growth of Dipterocarpaceae seedlings in Borneo. We used seeds that ranged in size from 500 to 20,000 mg in order to test the role of seed mass in mediating the effects of infrequent watering. With frequent rainfall, germination and seedling development traits bore no relationship to seed mass, but all metrics of seedling growth increased with increasing seed mass. Cumulative germination declined by 39.4% on average for all species when plants were watered at six-day intervals, and days to germination increased by 76.5% on average for all species from daily to six-day intervals. Final height and biomass declined on average in the six-day interval by 16% and 30%, respectively, but the percentage decrease in final size was greater for large-seeded species. Rooting depth per leaf area also significantly declined with seed mass indicating large-seeded species allocate relatively more biomass for leaf production. This difference in allocation provided an establishment advantage to large-seeded species when water was non-limiting but inhibited their growth under infrequent rainfall. The observed reduction in the growth of large-seeded species under infrequent rainfall would likely restrict their establishment in drier microsites associated with coarse sandy soils and ridge tops. In total, these species differences in germination and initial seedling growth indicates a possible niche axis that may help explain both current species distributions and future responses to climate change.     

Experimental evidence for a phylogenetic Janzen-Connell effect in a subtropical forest

Observational evidence increasingly suggests that the Janzen-Connell effect extends beyond the species boundary. However, this has not been confirmed experimentally. Herein, we present both observational and experimental evidence for a phylogenetic Janzen-Connell effect. In a subtropical forest in Guangdong province, China, we observed that co-occurring tree species are less phylogenetically related than expected. The inhibition effects of neighbouring trees on seedling survival decreased with increasing phylogenetic distance between them. In a shade-house experiment, we studied seedling survival of eight species on soil collected close to Castanopsis fissa relative to their survival on soil close to their own adult trees, and found that this relative survival rate increased with phylogenetic distance from C. fissa. This phylogenetic signal disappeared when seedlings were planted in fungicide-treated soil. Our results clearly support negative effects of phylogenetically similar neighbouring trees on seedling survival and suggest that these effects are caused by associated host-specific fungal pathogens.     

森林凋落物对林地微生境的影响

评述了凋落物改变林地微生境的效应．凋落物减弱了林地地表及表层土壤(2mm厚)中的光照,改变了光谱特性,降低了PPFD和Pfr．缩小了日温差,提高了土壤湿度和肥力,增加了毒素和土壤动物的活动等,但凋落物改变生境的效应强度与凋落物组成、厚度、分解状态和周围生境(如植被、气候、地形和地貌等特性)等有关,在自然森林生态系统中,凋落物对林地微生境的效应因凋落物、植被、气候、地形和地貌的多样化变得更为复杂,而凋落物对植被天然更新的影响也因其对生境改变效应的复杂性而具有多样性和综合性,最后,对凋落物改变生境效应的复杂性和有关研究进行了展望。     

Effects of light,moisture, temperature,and litter on the regeneration of five tree species in the tropical montane wet forest of Puerto Rico

Field experiments were conducted to investigate the effects of light, moisture, temperature, and litter on the regeneration of two early-, one mid-, and two late-successional tropical tree species. High light and litter seem to be universally good cues for regeneration, increasing seed/seedling survival for all species except for Cecropia (an early-successional species) whose small seeds may not be able to penetrate the litter layer. In addition, the high temperature environment in both artificially shaded and nonshaded areas of a natural gap exhibits less seed loss, an increase in the percent and rate of germination, and an increase in seedling survival for Dacryodes (a late-successional species), than the lower temperature environment under an intact canopy. Low soil water is also a good cue for Dacryodes germination as it is for Prestoea and Cecropia. Finally, the lower temperature environment found under the forest canopy (compared to the natural gap) leads to less seed loss and more germination for Guarea (a mid-successional species). Our results suggest that a good patch for regeneration of many species in this forest, early- as well as late-successional species, would have high light and a litter layer that moderates temperature and moisture extremes. The substantial variation in suitability among regeneration filters and species could: (1) contribute to low establishment success, i.e., most dispersed propagules do not become trees, (2) make it difficult to group species into germination strategies, and (3) make it hard to generalize about a net effect of any specific environmental variable on establishment. We suggest that tropical disturbances should be viewed in terms of their impact on a variety of environmental cues, which may signal germination and impact subsequent growth and survival.     

Scratching beneath the surface: Bandicoot bioturbation contributes to ecosystem processes

Animals that forage for food or dig burrows by biopedturbation can alter the biotic and abiotic characteristics of their habitat. The digging activities of such ecosystem engineers, although small at a local scale, may be important for broader scale landscape processes by influencing soil and litter properties, trapping organic matter and seeds, and subsequently altering seedling recruitment. We examined environmental characteristics (soil moisture content, hydrophobicity and litter composition) of foraging pits created by the southern brown bandicoot (Isoodon obesulus; Peramelidae), a digging Australian marsupial, over a 6‐month period. Fresh diggings typically contained a higher moisture content and lower hydrophobicity than undisturbed soil. A month later, foraging pits contained greater amounts of fine litter and lower amounts of coarse litter than adjacent undug surfaces, indicating that foraging pits may provide a conducive microhabitat for litter decomposition, potentially reducing litter loads and enhancing nutrient decomposition. We tested whether diggings might affect seedling recruitment (seed removal by seed harvesters and seed germination rates) by artificially mimicking diggings. Although there were no differences in the removal of seeds, seedling recruitment for three native plant species (Acacia saligna, Kennedia prostrata and Eucalyptus gomphocephala) was higher in plots containing artificial diggings compared with undug sites. The digging actions of bandicoots influenced soil moisture and hydrophobicity, the size distribution of litter and seedling recruitment at a local scale. The majority of Australian digging mammals are threatened, with many suffering substantial population and range contraction. However, their persistence in landscapes plays an important role in maintaining the health and function of ecosystems.     

西鄂尔多斯珍稀濒危植物长叶红砂种子萌发特征

珍稀濒危灌木长叶红砂是我国西北干旱地区荒漠植被中的古老残遗物种,对维持荒漠植被稳定具有重要作用.本文研究了长叶红砂种子特性,及其在不同光照、温度、土壤水分和沙埋等环境因子中的萌发策略.结果表明:长叶红砂种子具有高活力、高萌发率、耐贮存的特点,在光照和黑暗条件下均能很好地萌发.种子萌发的适宜温度为恒温20 ℃～25 ℃或变温15 ℃/25 ℃,萌发率高达93%.土壤含水量为2%时,种子开始萌发;土壤含水量为12%时,其萌发率最高,达89%.长叶红砂种子适宜的沙埋深度为1 cm,＞5 cm沙埋不出苗.沙埋深度对出苗率和幼苗生长高度的影响显著,而对幼苗质量的影响不显著.水分条件和沙埋深度是制约长叶红砂种子萌发和出苗的主要因素,而种子的高萌发率增加了幼苗生存的风险,不利于其应对荒漠极端环境变化.这种特性是长叶红砂濒危的重要原因之一.     

Germination,survival, and growth of grass and forb seedlings: Effects of soil moisture variability

Seed germination and seedling growth, survivorship, and final biomass and their responses to watering interval were studied in two grass and six forb species to assess germination and seedling growth responses to increased soil moisture variability as might occur with future increases in precipitation variability. Seeds were planted in prairie soil and watered at 1, 2, 4, or 7 d intervals (I). Seed germination peaked at I = 4 d whereas leaf growth in grasses and forbs, and final biomass in grasses peaked at I = 7 d, suggesting that growth and biomass were favored at greater soil moisture variability than seed germination. Biomass responses to I were stronger than the germination responses, suggesting that soil moisture variability more strongly influenced post germination growth. Individual species responses to I fell into three groups; those with responses to I for: (1) seed germination and seedling survival, (2) biomass, or (3) both germination and biomass production. These species groups may be more useful than life form (i.e., grass/forb) for understanding seed germination and seedling dynamics in grasslands during periods of soil moisture variability. Seed germination and early growth may assume more importance in grassland plant community dynamics under more variable precipitation patterns.