Nitrogen (N) limitation to net primary production is widespread and influences the responsiveness of ecosystems to many components of global environmental change. Logic and both simple simulation (Vitousek and Fieldin in Biogeochemistry46: 179–202, 1999) and analytical models (Menge in Ecosystems 14:519–532, 2011) demonstrate that the co-occurrence of losses of N in forms that organisms within an ecosystem cannot control and barriers to biological N fixation (BNF) that keep this process from responding to N deficiency are necessary for the development and persistence of N limitation. Models have focused on the continuous process of leaching losses of dissolved organic N in biologically unavailable forms, but here we use a simple simulation model to show that discontinuous losses of ammonium and nitrate, normally forms of N whose losses organisms can control, can be uncontrollable by organisms and can contribute to N limitation under realistic conditions. These discontinuous losses can be caused by temporal variation in precipitation or by ecosystem-level disturbance like harvest, fire, and windthrow. Temporal variation in precipitation is likely to increase and to become increasingly important in causing N losses as anthropogenic climate change proceeds. We also demonstrate that under the conditions simulated here, differentially intense grazing on N- and P-rich symbiotic N fixers is the most important barrier to the responsiveness of BNF to N deficiency.

The role of streams and rivers in the global carbon (C) cycle remains unconstrained, especially in headwater streams where CO2 evasion (FCO2) to the atmosphere is high. Stream C cycling is understudied in the tropics compared to temperate streams, and tropical streams may have among the highest FCO2 due to higher temperatures, continuous organic matter inputs, and high respiration rates both in-stream and in surrounding soils. In this paper, we present paired in-stream O2 and CO2 sensor data from a headwater stream in a lowland rainforest in Costa Rica to explore temporal variability in gas concentrations and ecosystem processes. Further, we estimate groundwater CO2 inputs (GWCO2) from riparian well CO2 measurements. Paired O2–CO2 data reveal stream CO2 supersaturation driven by groundwater CO2 inputs and large in-stream production of CO2. At short time scales, CO2 was diluted during storm events, but increased at longer seasonal scales. Areal fluxes in our study reach show that FCO2 is supported by greater in-stream metabolism compared to GWCO2. Our results underscore the importance of tropical headwater streams as large contributors of carbon dioxide to the atmosphere and show evaded C can be derived from both in-stream and terrestrial sources.