Are hippos Africa's most influential megaherbivore? A review of ecosystem engineering by the semi-aquatic common hippopotamus

Megaherbivores perform vital ecosystem engineering roles, and have their last remaining stronghold in Africa. Of Africa's remaining megaherbivores, the common hippopotamus (Hippopotamus amphibius) has received the least scientific and conservation attention, despite how influential their ecosystem engineering activities appear to be. Given the potentially crucial ecosystem engineering influence of hippos, as well as mounting conservation concerns threatening their long-term persistence, a review of the evidence for hippos being ecosystem engineers, and the effects of their engineering, is both timely and necessary. In this review, we assess, (i) aspects of hippo biology that underlie their unique ecosystem engineering potential; (ii) evaluate hippo ecological impacts in terrestrial and aquatic environments; (iii) compare the ecosystem engineering influence of hippos to other extant African megaherbivores; (iv) evaluate factors most critical to hippo conservation and ecosystem engineering; and (v) highlight future research directions and challenges that may yield new insights into the ecological role of hippos, and of megaherbivores more broadly. We find that a variety of key life-history traits determine the hippo's unique influence, including their semi-aquatic lifestyle, large body size, specialised gut anatomy, muzzle structure, small and partially webbed feet, and highly gregarious nature. On land, hippos create grazing lawns that contain distinct plant communities and alter fire spatial extent, which shapes woody plant demographics and might assist in maintaining fire-sensitive riverine vegetation. In water, hippos deposit nutrient-rich dung, stimulating aquatic food chains and altering water chemistry and quality, impacting a host of different organisms. Hippo trampling and wallowing alters geomorphological processes, widening riverbanks, creating new river channels, and forming gullies along well-utilised hippo paths. Taken together, we propose that these myriad impacts combine to make hippos Africa's most influential megaherbivore, specifically because of the high diversity and intensity of their ecological impacts compared with other megaherbivores, and because of their unique capacity to transfer nutrients across ecosystem boundaries, enriching both terrestrial and aquatic ecosystems. Nonetheless, water pollution and extraction for agriculture and industry, erratic rainfall patterns and human–hippo conflict, threaten hippo ecosystem engineering and persistence. Therefore, we encourage greater consideration of the unique role of hippos as ecosystem engineers when considering the functional importance of megafauna in African ecosystems, and increased attention to declining hippo habitat and populations, which if unchecked could change the way in which many African ecosystems function.     

Habitat utilization by hippopotamus in the Mara River*

The results are presented of a 2-month study of the Mara River hippopotamus population in the Serengeti National Park and the Maasai Mara Game Reserve. A method for correcting aerial counts of hippos in turbid rivers and based on herd behaviour characteristics is described. There were an estimated 1927 hippos at an average density of 16-1/km in the area censused. The population became more dispersed and the mean group size decreased from thirty to fifteen after a rise in water level. The value of comparisons between populations in various terms of expression of density, and mean group sizes, are discussed. The amount of day living space, defined as areas preferred or utilizable by hippo during the day, and productivity of the food plants are seen to be very important factors in hippo habitat utilization, and as constraints on population increase. Hippo use of the habitat and the effects of hippo on the vegetation at various distances from the river were quantified, and a cycle of vegetational changes suggested which may be associated with the effects of hippo grazing and fire in other parts of Africa. Groups were found to consist of nearly equal numbers of males and females. There was no evidence of territorial behaviour either on land or in the water. A number of observations on social interactions and defaecating behaviour are described and indicate that olfactory communication is very important in interactions between hippos. Hippo society is unstable due to climatic influences, but a dominance hierarchy including males and females exists in the groups. Males appear less firmly attached to groups than females, but a male is normally the dominant animal in a group.     

2001-2018年中国-老挝交通走廊核心区植被稳定性对极端干旱的响应

中国-老挝交通走廊核心区干旱事件的频繁发生威胁着该区域生态系统的稳定性。基于Palmer干旱指数（PDSI）和增强型植被指数（EVI）数据量化了中老交通走廊核心区不同植被类型的稳定性（年均EVI与其标准差之比）及其对干旱的抵抗力（干旱期间植被结构和功能保持其原始水平的能力）和恢复力（植被恢复到干旱前水平的速度）。结果表明：（1）2001-2018年间,中老交通走廊核心区共发生5次极端干旱事件,出现极端干旱的年份有2005年、2010年、2015年、2016年和2017年,以上年份极端干旱面积占总面积的比例分别为13.37%、47.46%、10.41%、12.00%和3.05%;（2）不同植被类型对极端干旱的响应不同,中老交通走廊核心区内常绿阔叶林的稳定性显著高于其他植被类型,和其他植被类型相比,常绿阔叶林虽然暴露在干旱环境中的时间更长,但其具有更高的稳定性,在维持中老交通走廊核心区的生态系统稳定性上发挥着重要作用;（3）常绿阔叶林和木本稀树草原对极端干旱的抵抗力和恢复力显著高于混交林、草地和农田,研究区内各植被类型对极端干旱的抵抗力与温度和降水呈显著正相关关系。     

Intake, ingesta retention, particle size distribution and digestibility in the hippopotamidae

Although several aspects of the digestive physiology of the hippopotamidae-non-ruminating foregut fermenters-have been described, ingesta kinetics and passage characteristics of these species are not well understood. The most outstanding feature of the hippo digestive physiology reported so far is the very long mean ingesta retention times (MRTs) measured by Foose [Foose, T., 1982. Trophic strategies of ruminant versus nonruminant ungulates. PhD dissertation, University of Chicago, Chicago.]. Since those data had been investigated with animals without water access, we intended to measure MRT in hippos which were allowed to enter water pools during the night. MRT parameters as well as dry matter (DM) digestibility were determined in four common (Hippopotamus amphibius) and four pygmy hippos (Hexaprotodon liberiensis) on two different diets each using cobalt ethylendiamintetraacetate (Co-EDTA) as a fluid, chromium (Cr)-mordanted fibre (<2 mm) as a particle and acid detergent lignin (ADL) as an internal digestibility marker. Four of the animals additionally received cerium (Ce)-mordanted fibres (2-10 mm) as particle markers. Total MRTs for fluids and particles ranged between 20-35 and 48-106 h in the common and between 13-39 and 32-107 h in the pygmy hippos. The difference between fluid and particle retention was greater than usually reported in ruminants. Excretion patterns of the markers differed from those usually observed in ruminants but resembled those reported for macropods (kangaroos), indicating a plug-flow reactor-like physiology in the hippo forestomach (FRST). This finding complements other described similarities between the macropod and the hippo forestomach. The measurements of larger particle retention profiles suggest that in the hippo, larger particles might be excreted either faster or at the same rate as smaller particles, indicating a general difference between ruminants and hippos with respect to differential particle retention. The digestive physiology of hippos is characterised by a generally low food intake, long ingesta retention times and dry matter digestibilities lower than reported in ruminants. Moderate digestibilities in spite of long retention times might be the result of the generally high average ingesta particle size in hippos. The comparatively easy management of pygmy hippos, together with the significant correlations between food intake, MRT and digestibility in the pygmy hippos of this study, recommends this species for further studies on the interplay of these parameters in herbivore digestive physiology.     

Amazon drought and forest response: Largely reduced forest photosynthesis but slightly increased canopy greenness during the extreme drought of 2015/2016

Amazon droughts have impacted regional ecosystem functioning as well as global carbon cycling. The severe dry‐season droughts in 2005 and 2010, driven by Atlantic sea surface temperature (SST) anomaly, have been widely investigated in terms of drought severity and impacts on ecosystems. Although the influence of Pacific SST anomaly on wet‐season precipitation has been well recognized, it remains uncertain to what extent the droughts driven by Pacific SST anomaly could affect forest greenness and photosynthesis in the Amazon. Here, we examined the monthly and annual dynamics of forest greenness and photosynthetic capacity when Amazon ecosystems experienced an extreme drought in 2015/2016 driven by a strong El Niño event. We found that the drought during August 2015–July 2016 was one of the two most severe meteorological droughts since 1901. Due to the enhanced solar radiation during this drought, overall forest greenness showed a small increase, and 21.6% of forests even greened up (greenness index anomaly ≥1 standard deviation). In contrast, solar‐induced chlorophyll fluorescence (SIF), an indicator of vegetation photosynthetic capacity, showed a significant decrease. Responses of forest greenness and photosynthesis decoupled during this drought, indicating that forest photosynthesis could still be suppressed regardless of the variation in canopy greenness. If future El Niño frequency increases as projected by earth system models, droughts would result in persistent reduction in Amazon forest productivity, substantial changes in tree composition, and considerable carbon emissions from Amazon.     

Effects of a multi-year drought on a drought-adapted shrub, <Emphasis Type="Italic">Artemisia tridentata</Emphasis>

Models of climate change predict more variable precipitation for much of western North America, including more severe multi-year droughts. Droughts are known to increase mortality to trees although less is known about effects on shrubs from arid environments and about effects on reproduction. In this study, we followed a cohort of young sagebrush plants from 2010 to 2016, a period that included a severe drought from 2012 to 2015. Plants experienced little mortality preceding and during the drought. However, in the year following the drought, 14% of individuals died and 33% of branches on living plants died. There was little flowering in the years preceding the drought and flowering increased in each successive year from 2014 to 2016. Plants that produced more flowers in 2015 had more dead branches in 2016. Larger plants had fewer branches that died. Contrary to expectations, afternoon shade was not associated with greater survival or flowering, perhaps because shaded plants were in proximity to large trees which likely competed for water. Plants of the two common chemotypes had similar rates of survival and flowering. Experimental watering during the summer of 2015 did not affect survival and may have increased flowering in 2016. If multi-year droughts become more common in the future, even drought-adapted shrubs may be expected to suffer high rates of mortality.     

High N,dry: Experimental nitrogen deposition exacerbates native shrub loss and nonnative plant invasion during extreme drought

Hotter, longer, and more frequent global change‐type drought events may profoundly impact terrestrial ecosystems by triggering widespread vegetation mortality. However, severe drought is only one component of global change, and ecological effects of drought may be compounded by other drivers, such as anthropogenic nitrogen (N) deposition and nonnative plant invasion. Elevated N deposition, for example, may reduce drought tolerance through increased plant productivity, thereby contributing to drought‐induced mortality. High N availability also often favors invasive, nonnative plant species, and the loss of woody vegetation due to drought may create a window of opportunity for these invaders. We investigated the effects of multiple levels of simulated N deposition on a Mediterranean‐type shrubland plant community in southern California from 2011 to 2016, a period coinciding with an extreme, multiyear drought in the region. We hypothesized that N addition would increase native shrub productivity, but that this would increase susceptibility to drought and result in increased shrub loss over time. We also predicted that N addition would favor nonnatives, especially annual grasses, leading to higher biomass and cover of these species. Consistent with these hypotheses, we found that high N availability increased native shrub canopy loss and mortality, likely due to the higher productivity and leaf area and reduced water‐use efficiency we observed in shrubs subject to N addition. As native shrub cover declined, we also observed a concomitant increase in cover and biomass of nonnative annuals, particularly under high levels of experimental N deposition. Together, these results suggest that the impacts of extended drought on shrubland ecosystems may be more severe under elevated N deposition, potentially contributing to the widespread loss of native woody species and vegetation‐type conversion.     

Stable isotope ecology of fossil hippopotamids from the Lake Turkana Basin of East Africa

The diet of African hippopotamids can be documented through δ ¹³C analyses of enamel and other tissues. Analysis of a 10-million-year sequence of hippopotamids in and near the Lake Turkana Basin of northern Kenya shows that hippos have included a substantial fraction of C₃ vegetation in their diets since the late Miocene when C₄ vegetation first appears in hippo diet as a measurable fraction. The C₄ component of vegetation becomes dominant (>50%) by Upper Burgi time ( c . 2 million years ago) but does not reach 100% for all individuals. It is therefore not unexpected that the δ ¹³C values of modern hippopotamids show a higher fraction of dietary C₃ biomass than has been estimated from traditional observations. Analysis of δ ¹⁸O of hippos from different stratigraphic levels shows no systematic trend over time; the average value for fossil hippos over the last 10 million years is similar to that of modern hippos from the Omo River system.     

Woody vegetation damage by the African elephant during severe drought at Pongola Game Reserve,South Africa

Elephants (Loxodonta africana) significantly alter ecosystem structure and composition through browsing (e.g. pollarding, debarking and toppling). Such browsing is predicted to intensify during severe drought which may become more common with climate change. Here, we make use of an elephant impact survey from 2012 to 2015 and during the El Nino drought of 2015–2016 at Pongola Game Reserve (107 km²), KwaZulu-Natal, to investigate how severe drought influenced damage severity of different tree heights and species by elephants in this small reserve. Contrary to expectations, damage to common species did not change with severe drought. Crown damage had the highest predicted probability across heights (29%–90%) and species (46%–75%) regardless of drought. However, we found severe drought increased the predicted probabilities of crown damage to smaller trees <4 m, mortality >6 m and severe damage at 4–6 m. Consequently, elephant damage during severe drought may alter vegetation structure by severely damaging or killing large trees (>4 m) and extensively damaging the crowns of trees <4 m. Long-term monitoring of elephant effects on woody vegetation is essential to enable science-based management in response to future drought and elephant damage (e.g. range expansion, beehive deterrents) to protect elephants and conserve woody vegetation.     

Legacies in Switchgrass Resistance to and Recovery from Drought Suggest That Good Years Can Sustain Plants Through Bad Years

The warm-season perennial switchgrass (Panicum virgatum) is a candidate bioenergy crop. To be successful, switchgrass production must be maintained on low-quality landscapes with minimal inputs while facing future climates that are expected to be more extreme and more variable. We propose that antecedent rainfall constrains how plants respond to drought, as well as subsequently recover from drought. To test this idea, we examined how six switchgrass genotypes responded to a 1-year severe drought and then recovered under normal rainfall in the following year. These plants had previously grown for 3 years under a range of dry to wet rainfall levels in a shallow-soil common garden with no fertilizer. Plants previously exposed to drought produced less biomass, and basal area after the severe drought was relieved compared to previously well-watered plants. In addition, there were legacy effects caused by plant size: plants that were larger pre-drought were more likely to survive the severe drought, and plants that were larger during the severe drought recovered more biomass, basal area, and tillers post-drought. Although genotypes differed somewhat in their responses, the size constraint was consistent across genotypes. These findings suggest that we can establish more drought-resilient switchgrass stands by, for example, planning for initial irrigation or planting during a wet year to allow plants to grow larger prior to experiencing drought. Additional studies are needed to understand whether these rainfall and size legacies persist or are transient.     

The effects of drought on the<Emphasis Type="Italic"> Solidago altissima</Emphasis>-<Emphasis Type="Italic">Eurosta solidaginis</Emphasis>-natural enemy complex: population dynamics,local extirpations,and measures of selection intensity on gall size

Environmental catastrophes, such as severe drought, can reduce host-plant quality and/or abundance, which in turn decrease levels of herbivore populations. Such changes in herbivore populations affect populations of their natural enemies. As part of a long-term field experiment (1983–1991), galls of Eurosta solidaginis from 16 fields in central Pennsylvania were systematically collected from goldenrod ramets. Galls were dissected to compare the occurrence of E. solidaginis mortality caused by its natural enemies in 2 drought years (1988, 1991) with 5 pre-drought years (1983–1987) and 2 post-drought years (1989–1990). Gall diameters were significantly smaller in both drought years and early larval death significantly decreased E. solidaginis survivorship in the first drought year. Of the natural enemies, the parasitoid wasp Eurytoma gigantea caused significant selection for larger gall size in all pre-drought years, the 1991 drought, and both post-drought years, due to its differential attack of smaller galls. In spite of drought-induced small gall size in 1988, there was negligible selection on gall size by natural enemies. However, populations of E. solidaginis did suffer local extirpations at nine of the 16 fields during the first drought year and population recoveries of the gall inducer and natural enemies varied among fields in the post-drought years. As a consequence of reduced herbivore abundance in drought and post-drought years, some natural-enemy populations were absent. Drought therefore drastically reduced the abundance of E. solidaginis and natural enemies resulting in slow recoveries to pre-drought numbers. Received: 16 April 1998 / Accepted: 4 August 1998     

Carnivory in the common hippopotamus Hippopotamus amphibius: implications for the ecology and epidemiology of anthrax in African landscapes

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Oxygen cycle in a hippo pool, Serengeti National Park, Tanzania

The vertical distribution of temperature and dissolved oxygen was sampled over 24 h in a pond of the Seronera River inhabited by hippos. The waters were very turbid (visibility < 2 cm) because of the high turbidity from animals trampling sediment and a permanent surface algal bloom sustained by faecal matter. Direct solar heating was restricted to the top few centimetres. This resulted in a strong thermal and density stratification inhibiting aeration of the water column. Waters at mid-depth were aerated only when hippos stirred the water. Anoxic conditions were common in bottom water, these were occasionally ventilated in daytime by mixing due to bottom heating from decaying organic matter and at night by convective cooling. Poor water quality in hippo pools may affect wildlife.     

Drought induces lagged tree mortality in a subalpine forest in the Rocky Mountains

Extreme climatic events are key factors in initiating gradual or sudden changes in forest ecosystems through the promotion of severe, tree-killing disturbances such as fire, blowdown, and widespread insect outbreaks. In contrast to these climatically-incited disturbances, little is known about the more direct effect of drought on tree mortality, especially in high-elevation forests. Therefore projections of drought-induced mortality under future climatic conditions remain uncertain. For a subalpine forest landscape in the Rocky Mountains of northern Colorado (USA), we quantified lag effects of drought on mortality of Engelmann spruce Picea engelmannii , subalpine fir Abies lasiocarpa , and lodgepole pine Pinus contorta . For the period 1910–2004, we related death dates of 164 crossdated dead trees to early-season and late-season droughts. Following early-season droughts, spruce mortality increased over five years and fir mortality increased sharply over 11 years. Following late-season droughts, spruce showed a small increase in mortality within one year, whereas fir showed a consistent period of increased mortality over two years. Pine mortality was not affected by drought. Low pre-drought radial growth rates predisposed spruce and fir to drought-related mortality. Spruce and fir trees that died during a recent drought (2000–2004) had significantly lower pre-drought growth rates than live neighbour trees. Overall, we found large interspecific differences in drought-related mortality with fir showing the strongest effect followed by spruce and pine. This direct influence of climatic variability on differential tree mortality has the potential for driving large-scale changes in subalpine forests of the Rocky Mountains.     

Missing teeth: Discordances in the trade of hippo ivory between Africa and Hong Kong

As the global epicentre of wildlife trade, Hong Kong plays an important role in the preservation or demise of biodiversity, including species found continents away. If mismanaged, legal trade in threatened species can lead to unsustainable exploitation. Inaccurate and incomplete trade records from the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES) undermine the regulation of this trade. We examine the trade of hippo (Hippopotamus amphibius) teeth to illustrate the extent of mismatched data between key trading nations. More than 90% of global hippo teeth trade is imported to, and re‐exported from, Hong Kong. Of that imported, over 75% originated in Tanzania or Uganda, but there are notable disparities in declared trade volumes. In most transactions, Hong Kong declares more volume imported than the volume declared exported by Uganda. Overall, Hong Kong has reported the import of 3,176 kg more hippo teeth than declared exported by Tanzania. This indicates that actual trade levels may exceed internationally agreed quotas. In total, over 14,000 kg of hippo teeth is unaccounted for between Uganda and Hong Kong, representing more than 2,700 individual hippos—2% of the global population. This gross discordance in trade data undermines regulatory measures and challenges the persistence of hippo populations in Africa.     

Rapid, landscape scale responses in riparian tundra vegetation to exclusion of small and large mammalian herbivores

Productive tundra plant communities composed of a variety of fast growing herbaceous and woody plants are likely to attract mammalian herbivores. Such vegetation is likely to respond to different-sized herbivores more rapidly than currently acknowledged from the tundra. Accentuated by currently changing populations of arctic mammals there is a need to understand impacts of different-sized herbivores on the dynamics of productive tundra plant communities. Here we assess the differential effects of ungulate (reindeer) and small rodent herbivores (voles and lemmings) on high productive tundra vegetation. A spatially extensive exclosure experiment was run for three years on river sediment plains along two river catchments in low-arctic Norway. The river catchments were similar in species pools but differed in species abundance composition of both plants and vertebrate herbivores. Biomass of forbs, deciduous shrubs and silica-poor grasses increased by 40–50% in response to release from herbivory, whereas biomass of silica-rich grasses decreased by 50–75%. Hence both additive and compensatory effects of small rodents and reindeer exclusion caused these significant changes in abundance composition of the plant communities. Changes were also rapid, evident after only one growing season, and are among the fastest and strongest ever documented in Arctic vegetation. The rate of changes indicates a tight link between the dynamics of productive tundra vegetation and both small and large herbivores. Responses were however not spatially consistent, being highly different between the catchments. We conclude that despite similar species pools, variation in plant species abundance and herbivore species dynamics give different prerequisites for change.     

Carbon and Nitrogen Decoupling Under an 11-Year Drought in the Shortgrass Steppe

The frequency and magnitude of drought is expected to increase in the US Great Plains under future climate regimes. Although semiarid systems are considered highly resistant to water limitation, novel drought events could alter linkages among biogeochemical processes, and result in new feedbacks that influence the timescale of ecosystem recovery. We examined changes in carbon and nitrogen cycling in the last 2 years of an 11-year drought manipulation in the shortgrass steppe, and under the first 2 years of recovery from drought. We measured plant production, plant tissue chemistry, soil trace gas flux, and soil inorganic nitrogen dynamics to test the extent that this magnitude of drought altered carbon and nitrogen fluxes and how these changes affected post-drought dynamics. We found that soil inorganic nitrogen was up to five times higher under severe drought than under control conditions, but that this nitrogen may not have been accessible to plants and microbial communities during drought due to diffusion limitations. Drought plots had higher N₂O flux when they received equal rainfall pulses, showing that this accumulated N may be vulnerable to loss. In addition, plants in drought plots had higher tissue nitrogen for 2 years following drought. These results show that decadal-length droughts that may occur under future precipitation regimes are likely to alter ecosystem properties through interactions among precipitation, vegetation, and N cycling. Shifts in plant N, vulnerability of nitrogen to loss, and rainfall use efficiency that we observed are likely to affect the recovery time of semiarid systems subject to droughts of this magnitude.     

Hippopotamus and livestock grazing: influences on riparian vegetation and facilitation of other herbivores in the Mara Region of Kenya

Riparian savanna habitats grazed by hippopotamus or livestock experience seasonal ecological stresses through the depletion of herbaceous vegetation, and are often points of contacts and conflicts between herbivores, humans and their livestock. We investigated how hippopotamus and livestock grazing influence vegetation structure and cover and facilitate other wild herbivores in the Mara region of Kenya. We used 5 km-long transects, each with 13 plots measuring 10 × 10 m², and which radiate from rivers in the Masai Mara National Reserve and adjoining community pastoral ranches. For each plot, we measured the height and visually estimated the percent cover of grasses, forbs, shrubs and bare ground, herbivore abundance and species richness. Our results showed that grass height was shortest closest to rivers in both landscapes, increased with increasing distance from rivers in the reserve, but was uniformly short in the pastoral ranches. Shifting mosaics of short grass lawns interspersed with patches of medium to tall grasses occurred within 2.5 km of the rivers in the reserve in areas grazed habitually by hippos. Hence, hippo grazing enhanced the structural heterogeneity of vegetation but livestock grazing had a homogenizing effect in the pastoral ranches. The distribution of biomass and the species richness of other ungulates with distance from rivers followed a quadratic pattern in the reserve, suggesting that hippopotamus grazing attracted more herbivores to the vegetation patches at intermediate distances from rivers in the reserve. However, the distribution of biomass and the species richness of other ungulates followed a linear pattern in the pastoral ranches, implying that herbivores avoided areas grazed heavily by livestock in the pastoral ranches, especially near rivers.     

Forest productivity recovery or collapse? Model-data integration insights on drought-induced tipping points

More frequent and severe droughts are driving increased forest mortality around the globe. We urgently need to describe and predict how drought affects forest carbon cycling and identify thresholds of environmental stress that trigger ecosystem collapse. Quantifying the effects of drought at an ecosystem level is complex because dynamic climate–plant relationships can cause rapid and/or prolonged shifts in carbon balance. We employ the CARbon DAta MOdel fraMework (CARDAMOM) to investigate legacy effects of drought on forest carbon pools and fluxes. Our Bayesian model-data fusion approach uses tower observed meteorological forcing and carbon fluxes to determine the response and sensitivity of aboveground and belowground ecological processes associated with the 2012–2015 California drought. Our study area is a mid-montane mixed conifer forest in the Southern Sierras. CARDAMOM constrained with gross primary productivity (GPP) estimates covering 2011–2017 show a ~75% reduction in GPP, compared to negligible GPP change when constrained with 2011 only. Precipitation across 2012–2015 was 45% (474 mm) lower than the historical average and drove a cascading depletion in soil moisture and carbon pools (foliar, labile, roots, and litter). Adding 157 mm during an especially stressful year (2014, annual rainfall = 293 mm) led to a smaller depletion of water and carbon pools, steering the ecosystem away from a state of GPP tipping-point collapse to recovery. We present novel process-driven insights that demonstrate the sensitivity of GPP collapse to ecosystem foliar carbon and soil moisture states—showing that the full extent of GPP response takes several years to arise. Thus, long-term changes in soil moisture and carbon pools can provide a mechanistic link between drought and forest mortality. Our study provides an example for how key precipitation threshold ranges can influence forest productivity, making them useful for monitoring and predicting forest mortality events.     

Effects of decadal experimental drought and climate extremes on vegetation growth in Mediterranean forests and shrublands

Increased drought combined with extreme episodes of heatwaves is triggering severe impacts on vegetation growth, particularly for plant communities in arid and semiarid ecosystems. Although there is an abundance of short‐term field drought experiments in natural ecosystems, remaining knowledge gaps limit the understanding and prediction of vegetation growth to ongoing and future climate scenarios. Here, we assessed the impacts of long‐term (1999–2016) experimental drought (ca. ?30% rainfall) on the vegetation growth of a Mediterranean high (H) and low (L)‐canopy forests and an early‐successional shrubland, as indicated by above‐ground biomass increment (ABI) and standing density, respectively. We found habitat context (impact of historical climate change, soil depth and successional status) of the study sites significantly affected the magnitude of climate impacts; there were synergistic effects of experimental drought and meteorological drought (Standardised Precipitation–Evapotranspiration Index, SPEI) as well as extreme dry years on vegetation growth. Long‐term experimental drought decreased the ABI for the two forest canopy types and the standing density for the shrubland. Water availabilities in winter–spring (SPEIs) were positively correlated with the ABI and standing density. Moreover, experimental drought decreased the vegetation growth in extreme dry years for the shrubland. We propose that future work not only study the vegetation dynamics with physiological, phenological and demographical changes in long‐term processes and across climate gradients, but also should explore the changes of multiple functions simultaneously (e.g. multifunctionality) under long‐term processes and extremes. This type of analysis of long‐term data is essential to understand and predict biodiversity loss, composition shifts, declines in ecosystem function and carbon budgets at temporal and spatial scales, to enable policy makers to design and implement strategies for the maintenance of sustainable ecosystem function under future climate change scenarios.