The alpine–subalpine Loch Vale watershed (LVW) of Colorado, USA, has relatively high natural lithogenic P5+ fluxes to surface waters. For 1992–2018, the largest number of stream samples with P5+ concentrations ([P5+]) above detection limits occurred in 2008, corresponding with the highest frost-cracking intensity (FCI). Therefore, relatively cold winters and warm summers with a comparatively low mean annual temperature partly influence stream [P5+]. Sediment cores were collected from The Loch, an outlet lake of the LVW. Iron-, Al-, and Mn-oxide-bound phosphorus (adsorbed and authigenic phosphates; NP) serves as a proxy measurement for paleolake [P5+]. The highest NP in the core occurred during the cold and dry Allerød interstade. The lowest NP concentrations in the core occurred during climatically very wet periods in the Late Pleistocene and Early Holocene. Therefore, [P5+] are highest with relatively cold winters followed by relatively warm summers, relatively low mean annual temperatures, and relatively little rainfall and/or cryospheric melting. Currently the LVW is experiencing warming and melting of the permanent cryosphere with a rapidly declining FCI since 2008. This has the potential to dramatically decrease [P5+] in surface water ecosystems of the LVW, reducing biological productivity, enhancing P-limitation, and increasing ecosystem reliance on aeolian P5+.

We investigated whether changes in nutrient availability affected N, P, S and polyphenol concentrations in different leaf-development stages of three brevideciduous and three evergreen dominant woody species in a nutrient-limited savanna woodland in Central Brazil. Treatments included eight years of annual fertilization with 100 kg ha−1 of N, P, N plus P and control, each replicated in four randomized 15 × 15-m plots. All species increased S concentrations (minimum 28%) in young and mature leaves in fertilized plots. Dalbergia miscolobium decreased total phenol concentrations with P (−34.3%, −23.7%) and NP fertilization (−28.2%, −17.1%). Blepharocalyx salicifolius increased total phenol (27.6%, 18.8%) and tannin (46.3%; 43.5%) in P fertilized and increased total phenol (33.9%) and tannin (27.8%, 43.5%) in NP fertilized plots. Total phenol concentration decreased with leaf age in Ouratea hexasperma, Styrax ferrugineus and Blepharocalyx salicifolius, which also decreased tannin concentration with leaf age. For all treatments, brevideciduous species had higher N, P, total phenols and tannin concentrations and lower S concentration than evergreens. These differences between phenological groups suggest that tropical ecosystems responses to environmental changes are more complex than anticipated by global vegetation models, with consequences for predictions in ecosystem functions and resilience.

Tropical tropical montane cloud forests (TMCFs) cover a small portion of the Earth, yet they are significant biodiversity hotspots and centres of endemism, and they provide important hydrological and biogeochemical functions that affect human livelihoods. Given their fundamental sensitivity to climate, TMCFs also serve as an early warning system for climate change impacts. This paper outlines a new international initiative, CloudNet, that aims to promote integrated research across TMCFs, and introduces a special issue that reviews emerging themes and topics in the ecology of TMCFs, highlighting knowledge gaps and suggesting new directions for research. CloudNet is helping coordinate several new research projects and protocols: (1) a global repository of TMCF data and meta-analyses across multiple sites; (2) a multi-site study of plant functional traits across TMCFs; (3) a multi-site study of decomposition processes across TMCFs; (4) a protocol for standardizing climate data collection across TMCFs. These studies are intended to evaluate the extent to which general patterns emerge, accounting for biogeographic, phylogenetic and environmental differences among sites. Common data collection across TMCFs should also allow better integration across disciplines, such as linking nutrient limitation, seed production and propagule recruitment, and enable cross-site comparisons of how TMCFs respond to drivers of global change, including rising cloud bases, increasing temperatures, altered disturbance regimes, biological invasions and extinction, and changing human land use.

Tropical montane forests (TMF) are associated with a widely observed suite of characteristics encompassing forest structure, plant traits and biogeochemistry. With respect to nutrient relations, montane forests are characterized by slow decomposition of organic matter, high investment in below-ground biomass and poor litter quality, relative to tropical lowland forests. However, within TMF there is considerable variation in substrate age, parent material, disturbance and species composition. Here we emphasize that many TMFs are likely to be co-limited by multiple nutrients, and that feedback among soil properties, species traits, microbial communities and environmental conditions drive forest productivity and soil carbon storage. To date, studies of the biogeochemistry of montane forests have been restricted to a few, mostly neotropical, sites and focused mainly on trees while ignoring mycorrhizas, epiphytes and microbial community structure. Incorporating the geographic, environmental and biotic variability in TMF will lead to a greater recognition of plant–soil feedbacks that are critical to understanding constraints on productivity, both under present conditions and under future climate, nitrogen-deposition and land-use scenarios.

In order to understand the stoichiometric balance between foliar nitrogen (N) and phosphorus (P) on P-poor soils, we investigated how foliar N and P attributes (i.e. N and P concentrations in green and senesced leaves, N and P resorption efficiencies) of 30 tropical tree species co-vary along a gradient of soil P availability across three forests on Mount Kinabalu, Borneo. We found strong and positive correlations between foliar N and P in the concentrations and resorption efficiencies within each forest and across the three forests. Slopes of standardized major axis between foliar N and P concentrations for both green and senesced leaves were not different among the three forests, although the values of the scaling exponent in the relationships of foliar N to P across the three forests were significantly lower than 1. We suggest that down-regulation of N concentrations in green leaves on P-poor soils is one of several possible mechanisms explaining why N concentrations decrease with decreasing P concentrations in both green and senesced leaves toward a lower P availability in soils. On the other hand, the physiological and ecological reasons why N and P resorption efficiencies are positively correlated with each other across tree species remain unclear.

Ecosystem pool of phosphorus (P) was determined as the sum of above-ground vegetation, roots, necromass and soils to 1 m deep in a tropical rain forest in Sabah, Malaysia. Relationships among soil P fractions, acid phosphatase activity and fine-root biomass across soil horizons were also determined to understand P availability. Ecosystem pools of P, and of simultaneously quantified nitrogen (N) and carbon (C) were 3.4, 12 and 370 Mg ha−1, respectively. Only 2.6% of the total ecosystem P was in the above-ground vegetation, contrasting to C (60%) and N (16%). Canopy foliage of dominant tree species showed an extremely high N to P ratio of 31.5, which implied the excessively short supply of P compared with ample N. Soil P primarily consisted of recalcitrant occluded fractions (78–91%) and only 4% was labile. Approximately three-quarters of labile soil P was an organic fraction (Po). The concentration of labile Po did not differ between soil horizons, while both phosphatase activity and fine-root density were the greatest in the topsoil (top 5 cm) and dramatically decreased with depth. This suggests that trees depend on the acquisition of P from the labile Po in the topsoil, despite a greater amount of labile P in the subsoil. Trees with a high foliar N/P ratio may invest N to acquire P from the topsoil by secreting phosphatase that consists of proteins, rather than investing C to extending roots to scavenge P in the subsoil.

Almost 50% of the estimated 2500 species in the Bromeliaceae grow epiphytically in a remarkably wide range of habitats from inhospitable deserts to tropical rain forests (Benzing 2000). The degree of dependence on the host varies (Benzing 1990, Laube & Zotz 2006), and in some cases, epiphytic bromeliads may dispense completely with living hosts, and may thrive on artificial substrates such as electrical cables. This is not entirely surprising because this family provides examples of particularly remarkable adaptations to the epiphytic habitat like water-impounding leaf bases and water- and nutrient-absorbing scales (Benzing 2000). The so-called atmospheric forms in this family (e.g. Tillandsia recurvata) possess a dense covering of these scales, and use roots entirely as holdfasts, while leaves perform photosynthesis and take up water and nutrients. Although quite regularly mentioned in the literature (Benzing 1990, 2000; Brighigna et al. 1997, Lüttge 1989), we are not aware of any study investigating the vigour of plants growing on cables compared with plants growing on trees. The only functional aspects that have already been studied are related to nutrients, i.e. presence of nitrogen-fixing endophytes in Tillandsia recurvata plants from natural hosts and from electrical cables in Mexico (Puente & Bashan 1994) and differences in chemical composition of plant tissues and nutrient retention by leaves in Tillandsia capillaris growing on cables and trees in Argentina (Abril & Bucher 2009). Intermittent water supply, but also low nutrient supply, play an important role in epiphyte ecology (Zotz & Hietz 2001), which leads to the following alternative hypotheses: (1) Plants on electrical cables are usually close to roads.

The mass of fine litterfall and nutrient circulation through litterfall were determined in four Melrosideros polymorpha/Cibotium spp.-dominated rainforests that differed in substrate age, parent material texture and annual precipitation on Kilauea and Mauna Loa volcanoes on the island of Hawaii. Three of the sites had rates of litterfall of 5.2 Mg ha−1 y−1; the fourth, which was on the most fertile soil, produced 7.0 Mg ha−1 y−1 of litterfall with higher concentrations of nitrogen and phosphorus. Tree ferns of the genus Cibotium cycled relatively large amounts of nitrogen, phosphorus and potassium through litterfall; their contribution to nutrient circulation was disproportionate to their mass in the forest, or in litterfall. The forest on the youngest substrate, which also had the lowest concentrations of nitrogen in litterfall, was fertilized with complete factorial combinations of nitrogen, phosphorus and a treatment consisting of all other plant nutrients. Additions of nitrogen increased the quantity and nitrogen concentration in litterfall during the second year following the initiation of fertilization, while no other treatment had a significant effect. Additions of nitrogen had no effect on litterfall mass or nutrient concentrations in the most nutrient-rich site.

Vascular epiphytes can be quite abundant in vegetation close to the ocean surf, where they are exposed to a more or less continuous input of salt spray. The ecophysiological basis of their occurrence, i.e. salt tolerance or avoidance, is unresolved, because all previous studies were observational and conclusions thus circumstantial. Here, the effect of varying concentrations of salt water on germination, and growth and survival of seedlings and established plants was investigated in a growth cabinet study under controlled conditions. Seeds (1500), seedlings (750) and small tank plants (336) were from four populations of Werauhia sanguinolenta that were growing either close to the sea or inland in Panama. Changes of Na+ and K+ concentrations in plant tissue were also determined. No differences in the sensitivity to salt were found among populations, nor among life stages. External concentrations (Cext) of up to 15% sea water (c. 0.5% Na+) allowed complete germination as well as positive growth and survival in both seedlings and established plants over short periods (8–10 wk). After longer exposure (12 wk) of established plants visible damage and increased mortality were observed at lower Cext, but critical tissue Na+ levels were similar: c. 50 mg gDW−1. It is concluded that this common epiphyte does not meet the definition of a halophyte, but still possesses a rather high tolerance to sodium.

Differentiation of limiting nutrients within small spatial scales has been observed in coastal mangrove forests, but research on other tropical peatlands suggests it is a more widespread phenomenon. In the Changuinola mire of coastal Panama, oligotrophy was hypothesized to increase along a gradient of peat development (peat doming). Nutrient and carbon concentration of leaf tissue, soil, and soil porewater were characterised over a successive sequence of plant communities along the gradient. Soil phosphorus (P) and nitrogen (N) concentrations decreased from 1200 μg P g−1 and 27 mg N g−1 to 377 μg P g−1 and 22 mg N g−1 within 2.7 km into the mire interior. These changes coincided with an increase in soil and average leaf N:P molar ratios from 52–128 and 24–41, respectively. Soil P was strongly related to leaf P and soil N:P to foliar N:P. There was a wide range in δ15N values for canopy (4.0 to −9.4‰), Campnosperma panamense (4.0 to −7.8‰) and understorey (4.8 to −3.1‰) species. Foliar δ15N values of canopy species were strongly related to soil N:P, soil P and leaf P. The depleted foliar δ15N values appeared to be an effect of both the N atmospheric source and P limitation. Here, P limitation is likely associated with ombrotrophic conditions that developed as hydrologic inputs became dominated by precipitation.

The effects of N, P, NP and NPKS application on plant growth, above-ground primary production and N and P accumulation in the above-ground biomass of a flooded savanna were assessed over two growth cycles. Application of N or P did not improve plant growth and primary production, addition of NP had a noticeable effect, whereas NPKS fertilization led to a three-fold increase in plant production. The four most important grasses in the analysed ecosystem responded to nutrient application with increasing growth, indicating that they all are constrained by nutrient availability, but each species exhibited a different seasonal growth pattern suggesting a temporal division of nutrient resources. N accumulated in the harvested biomass during the growth season was 50% above that of control plants when NP was applied and showed a four-fold increase when NPKS was added. With NP addition, in 2 mo plants accumulated three times more P in the harvested biomass than the control plants. The maximum N accumulated during the growth season in the NPKS treatment represented half the fertilizer N, and probably all the N added (15 g m−2) may have been used if the whole biomass had been considered. With the rather small amounts of P added to the soil in the P treatment (2 g m−2), the maximum amount of P accumulated after 2 mo scarcely represented 13% of the added P, suggesting that most of the fertilizer P became rapidly immobilized or sorbed, not being available to plants in the short term. However, the increasing growth rates of grasses during the following 7 mo indicate that some P fluxed from these pools to the soil solution and to plants.

Competitive interactions among pioneer species may have a significant impact on the course of secondary succession in tropical rain forests. We predicted that the outcome of competitive interactions in early succession would vary with soil nutrient availability. To test this hypothesis we grew seven pioneer tree species alone and in dense competitive mixtures, with four nutrient treatments: no nutrient addition, and N, P, and N plus P addition. Performance of plants grown alone and in mixtures was strongly nutrient limited. However, contrary to expectation, the competitive hierarchy among the seven species was almost identical among the four nutrient treatments. The dominant species, Melastoma malabathricum, accounted for > 70% of total stand biomass in all nutrient treatments. Seedlings of this species had higher rates of gas exchange and initial growth, and lower root allocation than the other species. Profiles of light availability within the competitive stands indicated that light levels at ground level were well above levels at which pioneer species can successfully survive and grow, yet seedlings of species other than Melastoma remained stunted. Leaf N concentrations in all stands were 25–55% reduced by competitive interactions, and N addition increased relative competitive performance in only Melastoma suggesting that Melastoma was particularly effective at N acquisition, limiting nutrient uptake by the other species. Toward the end of the experiment individuals of Melastoma began to reproduce, suggesting that the competitive hierarchy would have changed in a longer-duration experiment.

The fate of ice biota released via meltwater into pools of seawater trapped between melting ice floes (crack pools) was followed in late January in the southern Weddell Sea. Low salinity crack pools shared the following features: nitrate exhaustion, high pH and POC/PON ratios, high bacterial biomass composed of large cells, and a dense algal assemblage dominated to over 90% by only two diatom species. It is suggested that this “climax stage” evolved from a nutrient rich, moderate biomass situation prevailing in high salinity crack pools, and is representative of summer succession of sea ice biota. “Overflow” production following nitrate exhaustion by the algae resulted in internal (lipid) and external (presumably mucus) carbon pools. The latter must fuel bacterial biomass build-up, as algal mortality appeared to be low. The large algal and bacterial stocks point to low grazing pressure exerted by phagotrophic protists, presumably due to poor food quality (e.g. high C/N ratios) and/or excessive mucus production. It is concluded that environmental selection of the abundant ice algal species occurs under conditions prevailing in the disintegrating ice cover during summer, which differ drastically from those generally referred to as characteristic of the sea ice habitat at large (a combination of low temperature, low light and high salinity).

Relating patterns of species abundance to the processes that determine them at local and regional spatial scales is one of the central aims of ecological research. To examine the relationship between distribution and growth in tropical rain forest, seasonal and annual changes in tree diameter of two tree species with contrasting distribution patterns, Celtis mildbraedii and Strombosia glaucescens, were monitored across topographic gradients in moist semi-deciduous and moist evergreen forest in Ghana over 2 years, 1997-9. Concurrent measurements were made of soil water availability, and during 1997/8, of rainfall, nutrient availability and photosynthetically active radiation (PAR). The two sites received similar, low, total rainfall during 1997/8, but soil water availability in the dry season at the evergreen site exceeded that at the semi-deciduous forest site. Soil water availability was higher during 1998/9 at both sites. During 1997/8, PAR was similar at the two sites. The evergreen site had soils of lower pH, available P, Mg, Ca and K concentrations than the semi-deciduous forest site. Water availability and the sand content of the soil increased and concentrations of total N and C decreased, from summit to valley positions at both sites. Celtis mildbraedii had significantly faster growth rates than S. glaucescens, and growth of both species was greater at the semi-deciduous than the evergreen forest site during the wet year (1998/9) but not during the dry year (1997/8). Celtis mildbraedii in semi-deciduous forest grew less in valley than summit and slope positions. We conclude that in the absence of a severe dry season, growth is higher in semi-deciduous than evergreen forest, and is related to the higher soil fertility in more seasonal forest. The patterns are consistent with the hypothesis that concentrations of available P and/or cations in the soil limit growth in evergreen forest, while concentrations of N in valley soils limit growth of C. mildbraedii in semi-deciduous forest. There was no evidence that variation in PAR influenced growth rates in these forests. A reduction in growth rate on less fertile soils may be a factor determining the distribution limit of dry forest species, such as C. mildbraedii, in evergreen forest.

Four common fallow tree species were tested for a response to nitrogen and phosphorus fertilization. Seedlings or cuttings of Melicope glabra (Rutaceae), Macaranga gigantea (Euphorbiaceae), Persea romosa (Laureaceae), Peronema canescens (Verbenaceae) were grown in pots of a 50:50 mix of native soil and sand under 18% full sun for 18 weeks. Every 2 weeks, plants received either added N, added P, added N+P, or no added nutrients (control). Persea, a shade-tolerant species, and Macaranga, a light-demanding pioneer, improved relative growth rate with the addition of both N and P. Neither responded to N or P alone. Peronema and Melicope demonstrated luxury consumption of both N and P but did not show enhanced growth. Two of the four species tested (Persea and Melicope) were more limited by P than N. Macaranga was equally limited by both and Peronema was more limited by N. Along with previous studies, evidence for positive growth response and luxury consumption among light-demanding species suggests that P, rather than N, should limit seedling performance and may ultimately influence tree diversity in young secondary tropical forests.

Relationships between phytoplankton growth and supplies of phosphorus and nitrogen have been variously investigated in three lakes in two contrasted English lake areas using physiological nutrient indices (alkaline phosphatase activity, phosphorus debt, surplus phosphorus, phosphorus uptake kinetics, dark ammonium uptake, ammonium uptake kinetics, stimulation of 14C uptake, and batch bioassay) and by inspection of water chemistry. None of the lakes was significantly affected by wastewater effluent but all lay in agricultural catchments. Physiological indices and water chemistry suggested limitation of phytoplankton biomass mostly by phosphorus, and to some extent by nitrogen in summer in North Ormesby Broad, by nitrogen in summer and to some extent by phosphorus in Lily Broad and by nitrogen in White Mere. Indices did not always accord with one another, though were not severely misleading. They added little to conclusions that could be drawn from water chemistry alone. Reasons for the differential relative importance of nitrogen and phosphorus in the two lake areas are discussed. The paradigm of phosphorus limitation, though unchallenged in upland waters and those on poorly weathered rocks, may be less relevant in lowland lakes.

The growth of the grass Brachypodium pinnatum (L.) Beauv. in Dutch nutrient-poor chalk grasslands increases with enhanced nitrogen supply, whereas other grass species also require an enhanced phosphorus supply for a similar response (e.g. Dactylis glomerata L.), or are competitively suppressed at any increase in nutrient supply (e.g. Briza media L.). We investigated whether this interspecific variation in response to N and P supply is caused by differences in P productivity (PP), i.e. the instantaneous rate of biomass production per unit of P present in the plant. We hypothesized that PP is highest in Brachypodium pinnatum, in contrast to N productivity which is known to be the highest in Dactylis glomerata. Phosphorus productivity and its components were studied using a growth analysis with four exponential P addition rates of 0·03, 0·06, 0·09 and 0·11/0·15 mg P mg−1 P d−1.

Although Brachypodium pinnatum allocated more P to its leaf blades, it had a lower P productivity at high N and low P supply than did Dactylis glomerata. This was associated with a higher productivity per unit leaf P in Dactylis glomerata. Across all species and treatments, leaf PP showed a distinct negative correlation with P concentration per leaf area, regardless whether the variation in area-based leaf P concentration was caused by variation in leaf thickness, leaf tissue mass density or mass-based P concentration. A possible explanation for this would be a positive correlation between leaf chlorophyll concentration and P concentration, leading at high concentrations to shading within the leaf and to a low photosynthetic rate per unit leaf P. We conclude that a high PP is determined by the ability of a plant to distribute its P over a large leaf area, rather than by greater allocation of P to the leaves. Interspecific relationships for P productivity are similar to those known for N productivity.