Interannual variation in rainfall modulates temperature sensitivity of carbon allocation and flux in a tropical montane wet forest.

Tropical forests exert a disproportionately large influence on terrestrial carbon (C) balance but projecting the effects of climate change on C cycling in tropical forests remains uncertain. Reducing this uncertainty requires improved quantification of the independent and interactive effects of variable and changing temperature and precipitation regimes on C inputs to, cycling within and loss from tropical forests. Here, we quantified aboveground litterfall and soil-surface CO2 efflux ("soil respiration"; FS ) in nine plots organized across a highly constrained 5.2°C mean annual temperature (MAT) gradient in tropical montane wet forest. We used five consecutive years of these measurements, during which annual rainfall (AR) steadily increased, in order to: (a) estimate total belowground C flux (TBCF); (b) examine how interannual variation in AR alters the apparent temperature dependency (Q10 ) of above- and belowground C fluxes; and (c) quantify stand-level C allocation responses to MAT and AR. Averaged across all years, FS , litterfall, and TBCF increased positively and linearly with MAT, which accounted for 49, 47, and 46% of flux rate variation, respectively. Rising AR lowered TBCF and FS , but increased litterfall, with patterns representing interacting responses to declining light. The Q10 of FS , litterfall, and TBCF all decreased with increasing AR, with peak sensitivity to MAT in the driest year and lowest sensitivity in the wettest. These findings support the conclusion that for this tropical montane wet forest, variations in light, water, and nutrient availability interact to strongly influence productivity (litterfall+TBCF), the sensitivity of above- and belowground C fluxes to rising MAT (Q10 of FS , litterfall, and TBCF), and C allocation patterns (TBCF:[litterfall+TBCF]).

Non-native mangroves support carbon storage, sediment carbon burial, and accretion of coastal ecosystems.

Mangrove forests play an important role in climate change adaptation and mitigation by maintaining coastline elevations relative to sea level rise, protecting coastal infrastructure from storm damage, and storing substantial quantities of carbon (C) in live and detrital pools. Determining the efficacy of mangroves in achieving climate goals can be complicated by difficulty in quantifying C inputs (i.e., differentiating newer inputs from younger trees from older residual C pools), and mitigation assessments rarely consider potential offsets to CO2 storage by methane (CH4 ) production in mangrove sediments. The establishment of non-native Rhizophora mangle along Hawaiian coastlines over the last century offers an opportunity to examine the role mangroves play in climate mitigation and adaptation both globally and locally as novel ecosystems. We quantified total ecosystem C storage, sedimentation, accretion, sediment organic C burial and CH4 emissions from ~70 year old R. mangle stands and adjacent uninvaded mudflats. Ecosystem C stocks of mangrove stands exceeded mudflats by 434 ± 33 Mg C/ha, and mangrove establishment increased average coastal accretion by 460%. Sediment organic C burial increased 10-fold (to 4.5 Mg C ha-1  year-1 ), double the global mean for old growth mangrove forests, suggesting that C accumulation from younger trees may occur faster than previously thought, with implications for mangrove restoration. Simulations indicate that increased CH4 emissions from sediments offset ecosystem CO2 storage by only 2%-4%, equivalent to 30-60 Mg CO2 -eq/ha over mangrove lifetime (100 year sustained global warming potential). Results highlight the importance of mangroves as novel systems that can rapidly accumulate C, have a net positive atmospheric greenhouse gas removal effect, and support shoreline accretion rates that outpace current sea level rise. Sequestration potential of novel mangrove forests should be taken into account when considering their removal or management, especially in the context of climate mitigation goals.

Ecosystem carbon storage does not vary with mean annual temperature in Hawaiian tropical montane wet forests.

Theory and experiment agree that climate warming will increase carbon fluxes between terrestrial ecosystems and the atmosphere. The effect of this increased exchange on terrestrial carbon storage is less predictable, with important implications for potential feedbacks to the climate system. We quantified how increased mean annual temperature (MAT) affects ecosystem carbon storage in above- and belowground live biomass and detritus across a well-constrained 5.2 °C MAT gradient in tropical montane wet forests on the Island of Hawaii. This gradient does not systematically vary in biotic or abiotic factors other than MAT (i.e. dominant vegetation, substrate type and age, soil water balance, and disturbance history), allowing us to isolate the impact of MAT on ecosystem carbon storage. Live biomass carbon did not vary predictably as a function of MAT, while detrital carbon declined by ~14 Mg of carbon ha(-1) for each 1 °C rise in temperature - a trend driven entirely by coarse woody debris and litter. The largest detrital pool, soil organic carbon, was the most stable with MAT and averaged 48% of total ecosystem carbon across the MAT gradient. Total ecosystem carbon did not vary significantly with MAT, and the distribution of ecosystem carbon between live biomass and detritus remained relatively constant across the MAT gradient at ~44% and ~56%, respectively. These findings suggest that in the absence of alterations to precipitation or disturbance regimes, the size and distribution of carbon pools in tropical montane wet forests will be less sensitive to rising MAT than predicted by ecosystem models. This article also provides needed detail on how individual carbon pools and ecosystem-level carbon storage will respond to future warming.

Mycorrhizal feedbacks influence global forest structure and diversity.

One mechanism proposed to explain high species diversity in tropical systems is strong negative conspecific density dependence (CDD), which reduces recruitment of juveniles in proximity to conspecific adult plants. Although evidence shows that plant-specific soil pathogens can drive negative CDD, trees also form key mutualisms with mycorrhizal fungi, which may counteract these effects. Across 43 large-scale forest plots worldwide, we tested whether ectomycorrhizal tree species exhibit weaker negative CDD than arbuscular mycorrhizal tree species. We further tested for conmycorrhizal density dependence (CMDD) to test for benefit from shared mutualists. We found that the strength of CDD varies systematically with mycorrhizal type, with ectomycorrhizal tree species exhibiting higher sapling densities with increasing adult densities than arbuscular mycorrhizal tree species. Moreover, we found evidence of positive CMDD for tree species of both mycorrhizal types. Collectively, these findings indicate that mycorrhizal interactions likely play a foundational role in global forest diversity patterns and structure.

Impacts of insect frass and cadavers on soil surface litter decomposition along a tropical forest temperature gradient.

Insect herbivores play important roles in shaping many ecosystem processes, but how climate change will alter the effects of insect herbivory are poorly understood. To address this knowledge gap, we quantified for the first time how insect frass and cadavers affected leaf litter decomposition rates and nutrient release along a highly constrained 4.3°C mean annual temperature (MAT) gradient in a Hawaiian montane tropical wet forest. We constructed litterbags of standardized locally sourced leaf litter, with some amended with insect frass + cadavers to produce treatments designed to simulate ambient (Control = no amendment), moderate (Amended-Low = 2 × Control level), or severe (Amended-High = 11 × Control level) insect outbreak events. Multiple sets of these litterbags were deployed across the MAT gradient, with individual litterbags collected periodically over one year to assess how rising MAT altered the effects of insect deposits on litter decomposition rates and nitrogen (N) release. Increased MAT and insect inputs additively increased litter decomposition rates and N immobilization rates, with effects being stronger for Amended-High litterbags. However, the apparent temperature sensitivity (Q 10) of litter decomposition was not clearly affected by amendments. The effects of adding insect deposits in this study operated differently than the slower litter decomposition and greater N mobilization rates often observed in experiments which use chemical fertilizers (e.g., urea, ammonium nitrate). Further research is required to understand mechanistic differences between amendment types. Potential increases in outbreak-related herbivore deposits coupled with climate warming will accelerate litter decomposition and nutrient cycling rates with short-term consequences for nutrient cycling and carbon storage in tropical montane wet forests.

Mean annual temperature influences local fine root proliferation and arbuscular mycorrhizal colonization in a tropical wet forest.

Mean annual temperature (MAT) is an influential climate factor affecting the bioavailability of growth-limiting nutrients nitrogen (N) and phosphorus (P). In tropical montane wet forests, warmer MAT drives higher N bioavailability, while patterns of P availability are inconsistent across MAT. Two important nutrient acquisition strategies, fine root proliferation into bulk soil and root association with arbuscular mycorrhizal fungi, are dependent on C availability to the plant via primary production. The case study presented here tests whether variation in bulk soil N bioavailability across a tropical montane wet forest elevation gradient (5.2°C MAT range) influences (a) morphology fine root proliferation into soil patches with elevated N, P, and N+P relative to background soil and (b) arbuscular mycorrhizal fungal (AMF) colonization of fine roots in patches. We created a fully factorial fertilized root ingrowth core design (N, P, N+P, unfertilized control) representing soil patches with elevated N and P bioavailability relative to background bulk soil. Our results show that percent AMF colonization of roots increased with MAT (r 2 = .19, p = .004), but did not respond to fertilization treatments. Fine root length (FRL), a proxy for root foraging, increased with MAT in N+P-fertilized patches only (p = .02), while other fine root morphological parameters did not respond to the gradient or fertilized patches. We conclude that in N-rich, fine root elongation into areas with elevated N and P declines while AMF abundance increases with MAT. These results indicate a tradeoff between P acquisition strategies occurring with changing N bioavailability, which may be influenced by higher C availability with warmer MAT.

The impacts of degradation, deforestation and restoration on mangrove ecosystem carbon stocks across Cambodia.

Mangrove forest conservation can help reduce global C emissions. Despite this benefit to climate change mitigation and adaptation, mangrove forests are being deforested or degraded at an alarming rate, though restoration efforts may offset these losses. The impacts of deforestation to C stocks are relatively intuitive and result in significant decreases in C stocks. It remains unclear how degradation from selective harvesting of trees affects C stocks or how effective restoration efforts are at restoring C stocks. Furthermore, total ecosystem C (TEC) stocks of pristine mangroves can significantly vary spatially. To address these issues, we conducted an intensive, national assessment of mangrove forests across Cambodia using a grid approach to: 1) examine how land use land cover (i.e., pristine, deforested, degraded, and restored forests) impacts TEC stocks, and 2) how TEC stocks vary spatially across the country. TEC stocks from deforested mangroves were always lower than pristine forests, resulting in an overall loss of 60% C (480 Mg C ha-1). However, TEC stocks from degraded and 25-year-old restored mangroves forests did not differ from pristine forests. Mean TEC in mangroves was 784.7 ± 30.1 Mg C ha-1, decreasing from 957.2 ± 32.8 Mg C ha-1 in the northern region to 628.9 ± 33.1 Mg C ha-1 in the central region to 386.2 ± 19.1 Mg C ha-1 in the southern region of Cambodia. Intensive sampling in mangroves across Cambodia verified impacts of deforestation reported elsewhere, revealed the lack of degradation impacts on TEC stocks, and demonstrated the effectiveness of restoration on TEC stocks after only 25 years. Our gridded sampling approach was able to capture spatial variability across Cambodia and provide a more realistic TEC stock information that can be used for national reporting or participation in C markets.

Impact of Mean Annual Temperature on Nutrient Availability in a Tropical Montane Wet Forest.

Despite growing understanding of how rising temperatures affect carbon cycling, the impact of long-term and whole forest warming on the suite of essential and potentially limiting nutrients remains understudied, particularly for elements other than N and P. Whole ecosystem warming experiments are limited, environmental gradients are often confounded by variation in factors other than temperature, and few studies have been conducted in the tropics. We examined litterfall, live foliar nutrient content, foliar nutrient resorption efficiency (NRE), nutrient return, and foliar nutrient use efficiency (NUE) of total litterfall and live foliage of two dominant trees to test hypotheses about how increasing mean annual temperature (MAT) impacts the availability and ecological stoichiometry of C, N, P, K, Ca, Mg, Mn, Fe, Zn, and Cu in tropical montane wet forests located along a 5.2°C gradient in Hawaii. Live foliage responded to increasing MAT with increased N and K concentrations, decreased C and Mn concentrations, and no detectable change in P concentration or in foliar NRE. Increases in MAT increased nutrient return via litterfall for N, K, Mg, and Zn and foliar NUE for Mn and Cu, while decreasing nutrient return for Cu and foliar NUE for K. The N:P of litterfall and live foliage increased with MAT, while there was no detectable effect of MAT on C:P. The ratio of live foliar N or P to base cations and micronutrients was variable across elements and species. Increased MAT resulted in declining N:K and P:K for one species, while only P:K declined for the other. N:Ca and N:Mn increased with MAT for both species, while N:Mg increased for one and P:Mn increased for the other species. Overall, results from this study suggest that rising MAT in tropical montane wet forest: (i) increases plant productivity and the cycling and availability of N, K, Mg, and Zn; (ii) decreases the cycling and availability of Mn and Cu; (iii) has little direct effect on P, Ca or Fe; and (iv) affects ecological stoichiometry in ways that may exacerbate P-as well as other base cation and micronutrient - limitations to tropical montane forest productivity.

Changes in soil bacterial community diversity following the removal of invasive feral pigs from a Hawaiian tropical montane wet forest.

Nonnative, invasive feral pigs (Sus scrofa) modify habitats by disturbing soils and vegetation, which can alter biogeochemical processes. Soil microbial communities drive nutrient cycling and therefore also play important roles in shaping ecosystem structure and function, but the responses of soil microbes to nonnative ungulate removal remains poorly studied. We examined changes in the soil bacterial community over a ~25 year chronosequence of feral pig removal in tropical montane wet forests on the Island of Hawai'i. We extracted bacterial eDNA from soil samples collected inside and outside of ungulate exclosures along this chronosequence and sequenced the eDNA using the Illumina platform. We found that ungulate removal increased diversity of soil bacteria, with diversity scores positively correlated with time since removal. While functional and phylogenetic diversity were not significantly different between pig present and pig removed soils, soil bulk density, which decreases following the removal of feral pigs, was a useful predictor of dissimilarity among sites and correlated to changes in functional diversity. Additionally, increases in soil porosity, potassium, and calcium were correlated to increases in functional diversity. Finally, sites with greater mean annual temperatures were shown to have higher scores of both functional and phylogenetic diversity. As such, we conclude that feral pigs influence overall bacterial community diversity directly while influencing functional diversity indirectly through alterations to soil structure and nutrients. Comparatively, phylogenetic differences between communities are better explained by mean annual temperature as a climatic predictor of community dissimilarity.

Climate sensitive size-dependent survival in tropical trees.

Survival rates of large trees determine forest biomass dynamics. Survival rates of small trees have been linked to mechanisms that maintain biodiversity across tropical forests. How species survival rates change with size offers insight into the links between biodiversity and ecosystem function across tropical forests. We tested patterns of size-dependent tree survival across the tropics using data from 1,781 species and over 2 million individuals to assess whether tropical forests can be characterized by size-dependent life-history survival strategies. We found that species were classifiable into four 'survival modes' that explain life-history variation that shapes carbon cycling and the relative abundance within forests. Frequently collected functional traits, such as wood density, leaf mass per area and seed mass, were not generally predictive of the survival modes of species. Mean annual temperature and cumulative water deficit predicted the proportion of biomass of survival modes, indicating important links between evolutionary strategies, climate and carbon cycling. The application of survival modes in demographic simulations predicted biomass change across forest sites. Our results reveal globally identifiable size-dependent survival strategies that differ across diverse systems in a consistent way. The abundance of survival modes and interaction with climate ultimately determine forest structure, carbon storage in biomass and future forest trajectories.

Home range use and movement patterns of non-native feral goats in a tropical island montane dry landscape.

Advances in wildlife telemetry and remote sensing technology facilitate studies of broad-scale movements of ungulates in relation to phenological shifts in vegetation. In tropical island dry landscapes, home range use and movements of non-native feral goats (Capra hircus) are largely unknown, yet this information is important to help guide the conservation and restoration of some of the world's most critically endangered ecosystems. We hypothesized that feral goats would respond to resource pulses in vegetation by traveling to areas of recent green-up. To address this hypothesis, we fitted six male and seven female feral goats with Global Positioning System (GPS) collars equipped with an Argos satellite upload link to examine goat movements in relation to the plant phenology using the Normalized Difference Vegetation Index (NDVI). Movement patterns of 50% of males and 40% of females suggested conditional movement between non-overlapping home ranges throughout the year. A shift in NDVI values corresponded with movement between primary and secondary ranges of goats that exhibited long-distance movement, suggesting that vegetation phenology as captured by NDVI is a good indicator of the habitat and movement patterns of feral goats in tropical island dry landscapes. In the context of conservation and restoration of tropical island landscapes, the results of our study identify how non-native feral goats use resources across a broad landscape to sustain their populations and facilitate invasion of native plant communities.

Leaf litter decomposition rates increase with rising mean annual temperature in Hawaiian tropical montane wet forests.

Decomposing litter in forest ecosystems supplies nutrients to plants, carbon to heterotrophic soil microorganisms and is a large source of CO2 to the atmosphere. Despite its essential role in carbon and nutrient cycling, the temperature sensitivity of leaf litter decay in tropical forest ecosystems remains poorly resolved, especially in tropical montane wet forests where the warming trend may be amplified compared to tropical wet forests at lower elevations. We quantified leaf litter decomposition rates along a highly constrained 5.2 °C mean annual temperature (MAT) gradient in tropical montane wet forests on the Island of Hawaii. Dominant vegetation, substrate type and age, soil moisture, and disturbance history are all nearly constant across this gradient, allowing us to isolate the effect of rising MAT on leaf litter decomposition and nutrient release. Leaf litter decomposition rates were a positive linear function of MAT, causing the residence time of leaf litter on the forest floor to decline by ∼31 days for each 1 °C increase in MAT. Our estimate of the Q 10 temperature coefficient for leaf litter decomposition was 2.17, within the commonly reported range for heterotrophic organic matter decomposition (1.5-2.5) across a broad range of ecosystems. The percentage of leaf litter nitrogen (N) remaining after six months declined linearly with increasing MAT from ∼88% of initial N at the coolest site to ∼74% at the warmest site. The lack of net N immobilization during all three litter collection periods at all MAT plots indicates that N was not limiting to leaf litter decomposition, regardless of temperature. These results suggest that leaf litter decay in tropical montane wet forests may be more sensitive to rising MAT than in tropical lowland wet forests, and that increased rates of N release from decomposing litter could delay or prevent progressive N limitation to net primary productivity with climate warming.