Nutrient availability controls the impact of mammalian herbivores on soil carbon and nitrogen pools in grasslands

Grasslands are subject to considerable alteration due to human activities globally, including widespread changes in populations and composition of large mammalian herbivores and elevated supply of nutrients. Grassland soils remain important reservoirs of carbon (C) and nitrogen (N). Herbivores may affect both C and N pools and these changes likely interact with increases in soil nutrient availability. Given the scale of grassland soil fluxes, such changes can have striking consequences for atmospheric C concentrations and the climate. Here, we use the Nutrient Network experiment to examine the responses of soil C and N pools to mammalian herbivore exclusion across 22 grasslands, under ambient and elevated nutrient availabilities (fertilized with NPK + micronutrients). We show that the impact of herbivore exclusion on soil C and N pools depends on fertilization. Under ambient nutrient conditions, we observed no effect of herbivore exclusion, but under elevated nutrient supply, pools are smaller upon herbivore exclusion. The highest mean soil C and N pools were found in grazed and fertilized plots. The decrease in soil C and N upon herbivore exclusion in combination with fertilization correlated with a decrease in aboveground plant biomass and microbial activity, indicating a reduced storage of organic matter and microbial residues as soil C and N. The response of soil C and N pools to herbivore exclusion was contingent on temperature – herbivores likely cause losses of C and N in colder sites and increases in warmer sites. Additionally, grasslands that contain mammalian herbivores have the potential to sequester more N under increased temperature variability and nutrient enrichment than ungrazed grasslands. Our study highlights the importance of conserving mammalian herbivore populations in grasslands worldwide. We need to incorporate local‐scale herbivory, and its interaction with nutrient enrichment and climate, within global‐scale models to better predict land–atmosphere interactions under future climate change.     

Plant Diversity Impacts Decomposition and Herbivory via Changes in Aboveground Arthropods

Loss of plant diversity influences essential ecosystem processes as aboveground productivity, and can have cascading effects on the arthropod communities in adjacent trophic levels. However, few studies have examined how those changes in arthropod communities can have additional impacts on ecosystem processes caused by them (e.g. pollination, bioturbation, predation, decomposition, herbivory). Therefore, including arthropod effects in predictions of the impact of plant diversity loss on such ecosystem processes is an important but little studied piece of information. In a grassland biodiversity experiment, we addressed this gap by assessing aboveground decomposer and herbivore communities and linking their abundance and diversity to rates of decomposition and herbivory. Path analyses showed that increasing plant diversity led to higher abundance and diversity of decomposing arthropods through higher plant biomass. Higher species richness of decomposers, in turn, enhanced decomposition. Similarly, species-rich plant communities hosted a higher abundance and diversity of herbivores through elevated plant biomass and C:N ratio, leading to higher herbivory rates. Integrating trophic interactions into the study of biodiversity effects is required to understand the multiple pathways by which biodiversity affects ecosystem functioning.     

Developing X-ray Computed Tomography to non-invasively image 3-D root systems architecture in soil

Background

The positive relationship between biodiversity and ecosystem functioning (BEF) is due mainly to complementarity between species. Most BEF studies primarily focused on plant interactions; however, plants are embedded in a dense network of multitrophic interactions above and below the ground, which are likely to play a crucial role in BEF relationships.

Scope

In the present review I point out the relevance of aboveground–belowground interactions as a source of complementarity effects in grassland biodiversity experiments. A review of the current knowledge on the role of decomposers, arbuscular mycorrhizal fungi, rhizobia, plant growth promoting rhizobacteria, invertebrate ecosystem engineers, herbivores, pathogens and predators in biodiversity experiments, indicates that soil biota can drive both positive and negative complementarity between plant species via a multitude of mechanisms.

Conclusions

I pose four main processes by which aboveground–belowground interactions determine positive complementarity effects: enlarging biotope space, mediating legume effects, increasing plant community resistance, and maintaining plant diversity. By contrast, soil biota may also reinforce negative complementarity effects by competing with plants for nutrients or by exerting herbivore or pathogen pressure, thereby reducing community productivity. Thus, considering aboveground–belowground interactions as well as interactions between antagonistic and mutualistic consumers may improve the mechanistic understanding of complementarity effects in plant diversity–ecosystem functioning experiments and should inspire future research.     

Plant–herbivore interactions and ecological stoichiometry: when do herbivores determine plant nutrient limitation?

Recent studies on plant–herbivore indirect interactions via nutrient recycling have led to the hypothesis that herbivores with a low nitrogen: phosphorus ratio, feeding on plants with a higher nitrogen: phosphorus ratio, recycle relatively more nitrogen, driving plants into phosphorus limitation. We demonstrate in this paper that such a hypothesis is valid only under restricted conditions, i.e. the nitrogen: phosphorus ratio of inorganic nutrients supplied to the system must be neither too high nor too low compared with the nitrogen: phosphorus ratio of the whole plant + herbivore biomass. If plants have a greater affinity for phosphorus than for nitrogen, low herbivore nitrogen: phosphorus ratio can even promote nitrogen limitation. These results are qualitatively robust, whether grazing functions are donor-controlled or recipient-controlled. We present a graphical analysis of these conditions based on the Zero Net Growth Isocline method.     

气候因子通过土壤微生物生物量氮促进青藏高原高寒草地地上生态系统功能

近年来, 在人类活动和气候变化的影响下, 物种多样性丧失趋势不断加剧, 对生态系统功能带来严重后果。目前, 关于生态系统功能的研究, 忽略了土壤和微生物碳氮养分循环过程对地上生态系统功能(AEF)的重要驱动作用, 而土壤碳氮要素和微生物的任何变化都有可能改变地下群落对生态系统功能的维持作用。该研究旨在探究高寒草地AEF的主要控制因子, 以及其关键要素对AEF的作用机理。2015年7-8月, 对青藏高原地区115个样点进行了草地群落和土壤属性等要素样带调查; 综合植物地上生物量, 叶片碳、氮和磷含量等参数计算AEF值, 分析地下土壤有机碳含量、全氮含量、生物量等关键要素对AEF值的影响。结合取样点年降水量和年平均气温, 深入探讨影响AEF的主要控制因子和作用机理。结果表明降水对AEF有较大影响, 而气温影响相对较低。年降水量、土壤微生物生物量碳含量和干旱指数对AEF值的相对重要性贡献较高(重要值分别为21.1%、10.9%和10.1%), 控制青藏高原高寒草地AEF值的关键是土壤因子。在气候因子对土壤养分和微生物的作用下, 土壤微生物生物量氮含量在调控高寒草地AEF值方面发挥重要作用。     

Functional diversity of leaf nitrogen concentrations drives grassland carbon fluxes

Little is known about the role of plant functional diversity for ecosystem‐level carbon (C) fluxes. To fill this knowledge gap, we translocated monoliths hosting communities with four and 16 sown species from a long‐term grassland biodiversity experiment (‘The Jena Experiment’) into a controlled environment facility for ecosystem research (Ecotron). This allowed quantifying the effects of plant diversity on ecosystem C fluxes as well as three parameters of C uptake efficiency (water and nitrogen use efficiencies and apparent quantum yield). By combining data on ecosystem C fluxes with vegetation structure and functional trait‐based predictors, we found that increasing plant species and functional diversity led to higher gross and net ecosystem C uptake rates. Path analyses and light response curves unravelled the diversity of leaf nitrogen concentration in the canopy as a key functional predictor of C fluxes, either directly or indirectly via LAI and aboveground biomass.     

Trophic and non‐trophic interactions influence the mechanisms underlying biodiversity–ecosystem functioning relationships under different abiotic conditions

Plant diversity effects on ecosystem functioning usually have been studied from a plant perspective. However, the mechanisms underlying biodiversity–ecosystem functioning relationships may also depend on positive or negative interactions between plants and other biotic and abiotic factors, which remain poorly understood. Here we assessed whether plant–herbivore and/or plant–detritivore interactions modify the biodiversity–ecosystem functioning relationship and the mechanisms underlying biodiversity effects, including complementarity and selection effects, biomass allocation, vertical distribution of roots, and plant survival using a microcosm experiment. We also evaluated to what extent trophic and non‐trophic interactions are affected by abiotic conditions by studying drought effects. Our results show that biotic and abiotic conditions influence the shape of the biodiversity–ecosystem function relationship, varying from hump‐shaped to linear. For instance, total biomass increased linearly with plant richness in the presence of detritivores, but not in the absence of detritivores. Moreover, detritivore effects on belowground plant productivity were highly context dependent, varying in the presence of herbivores. Plant interactions with soil biota, especially with herbivores, influenced the mechanisms underlying diversity effects. Herbivores increased plant complementarity and modified biomass allocation and vertical distribution of roots. Furthermore, biotic–abiotic interactions influenced plant productivity differently across plant functional groups. Our findings emphasize the importance of complex biotic interactions underlying biodiversity effects, and that these biotic interactions may change with abiotic conditions. Despite minor changes in productivity in the short‐term, soil biota‐induced changes in plant–plant interactions and plant survival are likely to have significant long‐term consequences for ecosystem functioning. Considering the context‐dependency of multichannel interactions may contribute to reconciling differences among observed patterns in biodiversity studies. Further, abiotic conditions modified the effects of biotic interactions, suggesting that changes in environmental conditions may not only affect ecosystems directly, but also change the biotic composition of and dynamics within ecosystems.     

Invertebrate,not small vertebrate,herbivory interacts with nutrient availability to impact tallgrass prairie community composition and forb biomass

The effects of herbivores and their interactions with nutrient availability on primary production and plant community composition in grassland systems is expected to vary with herbivore type. We examined the effects of invertebrate and small vertebrate herbivores and their interactions with nutrient availability on grassland plant community composition and aboveground biomass in a tallgrass prairie ecosystem. The abundance of forbs relative to grasses increased with invertebrate herbivore removals. This increase in forb abundance led to a shift in community composition, where invertebrate removals resulted in greater plant species evenness as well as a divergence in composition among plots. In contrast, vertebrate herbivore removals did not affect plant community composition or aboveground biomass. Nutrient additions alone resulted in a decrease in plant species richness and an increase in the abundance of the dominant grass, but the dominant grass species did not greatly increase in abundance when nutrient additions were combined with invertebrate removals. Rather, several subdominant forbs came to dominate the plant community. Additionally, the combined nutrient addition and invertebrate herbivore removal treatment increased forb biomass, suggesting that invertebrate herbivores suppress the responses of forb species to chronic nutrient additions. Overall, the release of forbs from invertebrate herbivore pressure may result in large shifts in species composition, with consequences for aboveground biomass and forage quality due to altered grass:forb ratios in grassland systems.     

Plant controls on Late Quaternary whole ecosystem structure and function

Plants and animals influence biomass production and nutrient cycling in terrestrial ecosystems; however, their relative importance remains unclear. We assessed the extent to which mega‐herbivore species controlled plant community composition and nutrient cycling, relative to other factors during and after the Late Quaternary extinction event in Britain and Ireland, when two‐thirds of the region's mega‐herbivore species went extinct. Warmer temperatures, plant–soil and plant–plant interactions, and reduced burning contributed to the expansion of woody plants and declining nitrogen availability in our five study ecosystems. Shrub biomass was consistently one of the strongest predictors of ecosystem change, equalling or exceeding the effects of other biotic and abiotic factors. In contrast, there was relatively little evidence for mega‐herbivore control on plant community composition and nitrogen availability. The ability of plants to determine the fate of terrestrial ecosystems during periods of global environmental change may therefore be greater than previously thought.     

氮添加对昆仑山高山草地土壤、微生物和植物生态化学计量特征的影响

全球范围内的氮沉降增加改变了生态系统氮(N)素循环过程,由此带来的生态学效应已成为当前研究的热点。以昆仑山高山草地生态系统2种优势植物黄花棘豆(Oxytropis ochrocephala)和针茅(Stipa capillata)为研究对象,开展人工氮肥添加试验,研究土壤-微生物-植物系统各组分生态化学计量特征对氮添加的响应特征。结果表明:①氮添加显著提高了土壤NH4^+-N和土壤NO3^--N含量(P<0.05),土壤全N、全磷(P)、速效P含量没有明显变化。②氮添加条件下针茅叶片N含量增加,P含量降低,而黄花棘豆N和P含量无明显变化。③土壤微生物量碳(MBC)和微生物量氮(MBN)随着施氮量的增加呈现出先增加后降低的趋势,当施氮量为6N·m^-2·y^-1时呈现出最高值。土壤NH4+-N含量与土壤微生物量N含量有显著的正相关关系(P<0.01)。综合分析表明,短期氮添加有利于土壤养分和微生物量的积累,促进植物和微生物养分吸收利用。2种优势植物的生态化学计量特征对氮沉降的响应不同,过量的氮输入将会造成植物生长受到P限制,氮沉降会改变昆仑山高山草地生态系统的生物地球化学循环过程。     

Herbivory makes major contributions to ecosystem carbon and nutrient cycling in tropical forests

The functional role of herbivores in tropical rainforests remains poorly understood. We quantified the magnitude of, and underlying controls on, carbon, nitrogen and phosphorus cycled by invertebrate herbivory along a 2800 m elevational gradient in the tropical Andes spanning 12°C mean annual temperature. We find, firstly, that leaf area loss is greater at warmer sites with lower foliar phosphorus, and secondly, that the estimated herbivore‐mediated flux of foliar nitrogen and phosphorus from plants to soil via leaf area loss is similar to, or greater than, other major sources of these nutrients in tropical forests. Finally, we estimate that herbivores consume a significant portion of plant carbon, potentially causing major shifts in the pattern of plant and soil carbon cycling. We conclude that future shifts in herbivore abundance and activity as a result of environmental change could have major impacts on soil fertility and ecosystem carbon sequestration in tropical forests.     

Nitrogen fertilization affects bacteria utilizing plant-derived carbon in the rhizosphere of beech seedlings

Background and aims

Variation in fire intensity within an ecosystem is likely to moderate fire effects on plant and soil properties. We tested the effect of fire intensity on grassland biomass, soil microbial biomass, and soil nutrients. Additional tests determined plant-microbe, plant-nutrient, and microbe-nutrient associations.

Methods

A replicated field experiment produced a fire intensity gradient. We measured plant and soil microbial biomasses at peak plant productivity the first growing season after fire. We concurrently measured flux in 11 soil nutrients and soil moisture.

Results

Fire intensity positively affected soil nitrogen, phosphorus (P), and zinc but did not appreciably affect plant biomass, microbial biomass, and other soil nutrients. Plant biomass was seemingly (co-)limited by boron, manganese, and P. Microbial biomass was (co-)limited mainly by P and also iron.

Conclusions

In the Northern Great Plains, plant and soil microbial biomasses were limited mainly by P and some micronutrients. Fire intensity affected soil nutrients, however, pulsed P (due to fire) did not result in appreciable fire intensity effects on plant and microbial biomasses. Variable responses in plant productivity to fire are common and indicate the complexity of factors that regulate plant production after fire.

     

Small mammal herbivores mediate the effects of soil nitrogen and invertebrate herbivores on grassland diversity

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Responses of belowground communities to large aboveground herbivores: Meta‐analysis reveals biome‐dependent patterns and critical research gaps

The importance of herbivore–plant and soil biota–plant interactions in terrestrial ecosystems is amply recognized, but the effects of aboveground herbivores on soil biota remain challenging to predict. To find global patterns in belowground responses to vertebrate herbivores, we performed a meta‐analysis of studies that had measured abundance or activity of soil organisms inside and outside field exclosures (areas that excluded herbivores). Responses were often controlled by climate, ecosystem type, and dominant herbivore identity. Soil microfauna and especially root‐feeding nematodes were negatively affected by herbivores in subarctic sites. In arid ecosystems, herbivore presence tended to reduce microbial biomass and nitrogen mineralization. Herbivores decreased soil respiration in subarctic ecosystems and increased it in temperate ecosystems, but had no net effect on microbial biomass or nitrogen mineralization in those ecosystems. Responses of soil fauna, microbial biomass, and nitrogen mineralization shifted from neutral to negative with increasing herbivore body size. Responses of animal decomposers tended to switch from negative to positive with increasing precipitation, but also differed among taxa, for instance Oribatida responded negatively to herbivores, whereas Collembola did not. Our findings imply that losses and gains of aboveground herbivores will interact with climate and land use changes, inducing functional shifts in soil communities. To conceptualize the mechanisms behind our findings and link them with previous theoretical frameworks, we propose two complementary approaches to predict soil biological responses to vertebrate herbivores, one focused on an herbivore body size gradient, and the other on a climate severity gradient. Major research gaps were revealed, with tropical biomes, protists, and soil macrofauna being especially overlooked.     

Climate change disproportionately increases herbivore over plant or parasitoid biomass

All living organisms are linked through trophic relationships with resources and consumers, the balance of which determines overall ecosystem stability and functioning. Ecological research has identified a multitude of mechanisms that contribute to this balance, but ecologists are now challenged with predicting responses to global environmental changes. Despite a wealth of studies highlighting likely outcomes for specific mechanisms and subsets of a system (e.g., plants, plant-herbivore or predator-prey interactions), studies comparing overall effects of changes at multiple trophic levels are rare. We used a combination of experiments in a grassland system to test how biomass at the plant, herbivore and natural enemy (parasitoid) levels responds to the interactive effects of two key global change drivers: warming and nitrogen deposition. We found that higher temperatures and elevated nitrogen generated a multitrophic community that was increasingly dominated by herbivores. Moreover, we found synergistic effects of the drivers on biomass, which differed across trophic levels. Both absolute and relative biomass of herbivores increased disproportionately to that of plants and, in particular, parasitoids, which did not show any significant response to the treatments. Reduced parasitism rates mirrored the profound biomass changes in the system. These findings carry important implications for the response of biota to environmental changes; reduced top-down regulation is likely to coincide with an increase in herbivory, which in turn is likely to cascade to other fundamental ecosystem processes. Our findings also provide multitrophic data to support the general concern of increasing herbivore pest outbreaks in a warmer world.     

Insects affect relationships between plant species richness and ecosystem processes

This study examined whether insects can alter relationships between plant species diversity and ecosystem function in grassland communities, by (i) altering biomass across a plant diversity gradient, (ii) altering relative abundances of plant species, or (iii) altering ecosystem function directly. We measured herbivore damage on seminatural grassland plots planted with 1, 2, 4, 8, or 12 plant species, and compared plant biomass in a subset of these plots with replicates in which insect levels were reduced. Plant biomass and herbivore damage increased with species richness. Reducing insect populations resulted in greater evenness of relative plant species abundances and revealed a strong positive relationship between plant species richness and above-ground biomass. Reducing insects also changed the relationship between plant species richness and decomposition. Plant species mixtures and their relative abundances partially explained plant biomass results, but not decomposition results. These results suggest that insects can alter relationships between plant diversity and ecosystem processes through all three mechanisms.     

Distinct impacts of different mammalian herbivore assemblages on arctic tundra CO2 exchange during the peak of the growing season

Herbivores play a key role in the carbon (C) cycle of arctic ecosystems, but these effects are currently poorly represented within models predicting land–atmosphere interactions under future climate change. Although some studies have examined the influence of various individual species of herbivores on tundra C sequestration, few studies have directly compared the effects of different herbivore assemblages. We measured peak growing season instantaneous ecosystem carbon dioxide (CO₂) exchange (photosynthesis, respiration and net ecosystem exchange) on replicated plots in arctic tundra which, for 14 years, have excluded different portions of the herbivore population (grazed controls, large mammals excluded, both small and large mammals excluded). Herbivory suppressed photosynthetic CO₂ uptake, but caused little change in ecosystem respiration. Despite evidence that small mammals consume a greater portion of plant biomass in these ecosystems, the effect of excluding only large herbivores was indistinguishable from that of excluding both large and small mammals. The herbivory‐induced decline in photosynthesis was not entirely attributable to a decline in leaf area but also likely reflects shifts in plant community composition and/or species physiology. One shrub species – Betula nana – accounted for only around 13% of total aboveground vascular plant biomass but played a central role in controlling ecosystem CO₂ uptake and release, and was suppressed by herbivory. We conclude that herbivores can have large effects on ecosystem C cycling due to shifts in plant aboveground biomass and community composition. An improved understanding of the mechanisms underlying the distinct ecosystem impacts of different herbivore groups will help to more accurately predict the net impacts of diverse herbivore communities on arctic C fluxes.     

Effects of different fencing regimes on community structure of degraded desert grasslands on Mu Us desert,China

Grazing is one of the major anthropogenic driving factors influencing community structure and ecological function of grasslands. Fencing has been proved to be one of the main measures for rehabilitating degraded grasslands in northwestern China. However, data from combined empirical studies on the effects of different management regimes in desert grasslands are lacking. So we selected long‐term fencing (fenced since 1991), mid‐term fencing and seasonal fencing (fenced since 2002), and adjacent free‐grazing grasslands to investigate vegetation and soil properties on southwest Mu Us desert. Our results showed that fencing increased plant cover, height, aboveground biomass (AGB) of different plant life‐form groups, Shannon–Wiener diversity index, Evenness index, Simpson index, total soil nitrogen, total soil phosphorus, and soil organic matter, but decreased plant density, species richness, Richness index, soil bulk density, water content, and pH. However, 22–24 years of long‐term complete fencing might cause redegradation of vegetation and soil nutrients, characterized by the reduction of some vegetation properties, biodiversity, total AGB, and some soil properties. Seasonal fencing with 11–13 year was more beneficial to vegetation restoration than that with completely fencing measures. Our study suggests that appropriate artificial disturbances, such as seasonal fencing (winter grazing and summer fencing), should be used after long‐term fencing in order to maintain grassland productivity and biodiversity. These findings will help to provide theoretical support for vegetation restoration and sustainable management in grassland under grazing prohibition at Mu Us desert.     

内蒙古草甸草原土壤理化性质和微生物学特性对刈割与氮添加的响应

频繁的刈割和氮输入增加是导致草地生态系统退化的重要原因.土壤微生物学特性作为评估土壤质量的重要生物学指标,对草地刈割和氮输入增加的响应规律仍不十分明确.本研究依托内蒙古呼伦贝尔草原刈割复合氮添加野外实验平台,分析了土壤理化性质、土壤微生物生物量、土壤呼吸和土壤酶对刈割、氮添加的响应及其生长季动态变化.结果表明: 刈割显著降低了土壤微生物生物量碳、氮、磷和土壤呼吸(基础呼吸和底物诱导呼吸),与刈割后导致的水分限制及碳限制有关.刈割显著降低了氮磷获取酶(N-乙酰-β-D-葡萄糖苷酶和酸性磷酸单酯酶)的活性,符合“资源分配假说”.氮添加显著降低土壤pH值,但土壤微生物生物量对氮添加和pH降低均无显著响应,表明氮输入增加引起的土壤酸化不是影响微生物生物量的主要因素.氮添加对土壤呼吸和酶活性也无显著影响,与以往在典型草原的大多数研究结果不一致.刈割和氮添加复合处理显著降低了土壤微生物生物量磷,但提高了土壤中有效磷含量,降低了酸性磷酸酶活性.微生物生物量碳、氮、磷和土壤呼吸等的相关参数均在7月最高,这与夏季高温多雨有关.土壤酶活性在春夏季较高,生长季末期较低.这表明在该草甸草原,刈割将导致土壤碳氮磷养分失衡,从而加剧草原退化;而氮添加在短期内并未对土壤微生物生物量和活性产生显著影响.     

Warming alters coupled carbon and nutrient cycles in experimental streams

Although much effort has been devoted to quantifying how warming alters carbon cycling across diverse ecosystems, less is known about how these changes are linked to the cycling of bioavailable nitrogen and phosphorus. In freshwater ecosystems, benthic biofilms (i.e. thin films of algae, bacteria, fungi, and detrital matter) act as biogeochemical hotspots by controlling important fluxes of energy and material. Understanding how biofilms respond to warming is thus critical for predicting responses of coupled elemental cycles in freshwater systems. We developed biofilm communities in experimental streamside channels along a gradient of mean water temperatures (7.5–23.6 °C), while closely maintaining natural diel and seasonal temperature variation with a common water and propagule source. Both structural (i.e. biomass, stoichiometry, assemblage structure) and functional (i.e. metabolism, N₂‐fixation, nutrient uptake) attributes of biofilms were measured on multiple dates to link changes in carbon flow explicitly to the dynamics of nitrogen and phosphorus. Temperature had strong positive effects on biofilm biomass (2.8‐ to 24‐fold variation) and net ecosystem productivity (44‐ to 317‐fold variation), despite extremely low concentrations of limiting dissolved nitrogen. Temperature had surprisingly minimal effects on biofilm stoichiometry: carbon:nitrogen (C:N) ratios were temperature‐invariant, while carbon:phosphorus (C:P) ratios declined slightly with increasing temperature. Biofilm communities were dominated by cyanobacteria at all temperatures (>91% of total biovolume) and N₂‐fixation rates increased up to 120‐fold between the coldest and warmest treatments. Although ammonium‐N uptake increased with temperature (2.8‐ to 6.8‐fold variation), the much higher N₂‐fixation rates supplied the majority of N to the ecosystem at higher temperatures. Our results demonstrate that temperature can alter how carbon is cycled and coupled to nitrogen and phosphorus. The uncoupling of C fixation from dissolved inorganic nitrogen supply produced large unexpected changes in biofilm development, elemental cycling, and likely downstream exports of nutrients and organic matter.